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

VOLUME 73 BAND 73

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

VOLUME 73 BAND

THE TRUSTEES OF THE DIE TRUSTEES VAN DIE
SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM
CAPE TOWN_ KAAPSTAD

1977

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

440

LIST OF CONTENTS

CLARK, A. M.

The South African Museum’s Meiring Naude cruises, Part 4. Echinoderms.
(Published June 1977.)

EASTWOOD, E. B.

Notes on the scorpion fauna of the Cape, Part 2. The Parabuthus capensis (Ehren-
berg) species-group; remarks on taxonomy and bionomics (Arachnida,
Scorpionida, Buthidae). (Published August 1977.)

EstTEs, R.

Relationships of the South African fossil frog Eoxenopoides reuningi (Anura,
Pipidae). (Published June 1977.) a on nf 53

GRIFFITHS, C. L.
Deep-sea amphipods from west of Cape Point, South Africa. (Published June 1977.)

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

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

Cretaceous faunas from Zululand and Natal, South Africa. The ammonite family
Tetragonitidae Hyatt, 1900. (Published August 1977.) ..

KLINGER, H. C.
Cretaceous deposits near Bogenfels, South West Africa. (Published June 1977.) ..

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

Cretaceous faunas from Zululand, South Africa, and southern Mozambique. The
Aptian Ancyloceratidae (Ammonoidea). (Published December 1977.)

MILLARD, N. A. H.

Hydroids from the Kerguelen and Crozet shelves, collected by the cruise MD. 03 of
the Marion- Dufresne. (Published April 1977.) : y hye &

MILLARD, N. A. H.
The South African Museum’s Meiring Naude cruises, Part 3. Hydroida. (Published
June 1977.) Ae is xe pt Ee Be ae De

WOOLDRIDGE, T.

A new species of Halicyclops (Copepoda, Cyclopoida) from estuaries in Transkei,
southern Africa. (Published November 1977.) :

Page

133

199

49

93

149

81

215

105

361

NEW GENERIC AND SUBGENERIC NAMES PROPOSED
IN THIS VOLUME

Page
Helicancyloceras Klinger & Kennedy, 1977 ae ot Le ae os @-o2)
Helicancyloceras (Helicancyloceras) Klinger & Kennedy, 1977 ee ca a we OST
Helicancyloceras (Nonyaniceras) Klinger & Bee 1977 ae see a rs P27 |

Uniscyphus Millard, 1977 oe ae ae fe a ae Fe bse Roe! 274

VOLUME 73 PART 1 APRIL 1977 | ISSN 0303-2515

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

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

FiscHER, P.-H., DuvAL, M. & 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. 19600. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4): 1-51.

THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
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(continued inside back cover)

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

Volume. 73 Band
April 1977 April
Part 1 Deel

HYDROIDS FROM THE KERGUELEN
PEC ROZET SHELVES, COLLECTED BY
THE CRUISE MD.03 OF THE MARION-DUFRESNE

By

N. A. H. MILLARD

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material
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Obtainable from the South African Museum, P.O. Box 61, Cape Town 8000

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OUT OF PRINT/ UIT DRUK

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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

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES,
COLLECTED BY THE CRUISE MD.03 OF THE MARION-DUFRESNE

By
N. A. H. MILLARD
South African Museum, Cape Town

(With 12 figures and 2 tables)

[MS. accepted 28 December 1976]

ABSTRACT

Thirty-four species of hydroids are described from the Kerguelen and Crozet groups of
islands, including two new species: Halecium dufresneae and Zygophylax crozetensis. Among
them are five cosmopolitan species. The rest show affinities partly with the cold-temperate
Magellan Region of South America, and partly with Antarctica. The fauna is considered to
be subantarctic in nature.

CONTENTS

PAGE
Introduction é : : : : 1
Station list . ; ; 3 : ’ 2
Systematic section . : : : 3
Discussion . : ; . : ; 44
Acknowledgements . : : : 46
References . : , ; : i 46

INTRODUCTION

The cruise MD.03 of the Marion- Dufresne during the period 18 March to
28 April 1974 to the Kerguelen and Crozet groups of islands was intended as
a preliminary reconnaissance of the area to serve as a basis for a more detailed
survey. The cruise was highly successful in the quantity and variety of benthic
invertebrates taken, and although the subsequent cruises MD.04 (1975) and
MD.08 (1976) have now been completed, it may be some time before the hydroid
material is all sorted and worked up. It was felt that the publication of the
present results should not be delayed.

The cruise was financially and logistically supported by the Terres Australes
et Antarctiques Frangaises, Paris. The bulk of the material and the types of the
new species will be deposited in the Muséum National d’Histoire Naturelle, and
a few duplicates are being retained at the South African Museum.

The cruise MD.03 was described by the Chef de la Campagne, J. C. Hureau
(1976), who also gave a complete station list and a chart of the voyage. The
positions of the stations yielding hydroid material are, however, repeated here
for convenience.

1

Ann. S. Afr. Mus. 73 (1), 1977: 1-47, 12 figs, 2 tables.

Nm

ANNALS OF THE SOUTH AFRICAN MUSEUM

The Kerguelen Shelf lies in the southern Indian Ocean at approximately
50°S and 70°E. Kerguelen is the largest island, Heard and MacDonald lying
about 644 km to its south-east. The depths from which hydroids were taken
range from 6 to 790 m, the abyssal dredgings from over the edge of the plateau
yielding no hydroid material.

The Crozet Shelf (including among others the Ile de la Crozet, Ile de la
Possession and Ile aux Cochon) lies some | 930 km to the west of Kerguelen,
and hydroids were dredged from depths of 110 to 400 m.

STATION LIST

2/6. E. Kerguelen; 4.4.74; 49°30,7’'S 70°44,7’E; 115 m: Zygophylax crozetensis, Sym-
plectoscyphus subarticulatus.

2/7. E. Kerguelen; 4.4.74; 49°33,2’S 70°47,1’E; 130 m: Symplectoscyphus subarticulatus.

3/10. E. Kerguelen; 5.4.74; 49°28,5’S 71°52,8’E; 650 m: Eudendrium tottoni, Modeeria
rotunda, Opercularella belgicae, Lafoea dumosa, Plumularia insignis.

3/11. E. Kerguelen; 5.4.74; 49°25,4’S 71°51,7’E; 620-650 m: Eudendrium tottoni, Modeeria
rotunda, Filellum serratum, Lafoea dumosa, Symplectoscyphus subdichotomus, Plumu-
laria insignis.

7/22. N.E. Heard; 7.4.74; 52°12,7’S 75°38,4’E; 525-560 m: Staurotheca dichotoma, Plumu-
laria insignis.

8/24. N. Heard; 8.4.74; 52°58,0’S 73°42,0’E; 123 m: Campanularia norvegiae, Staurotheca
dichotoma, Sertularella picta, Symplectoscyphus elongatus, S. mawsoni.

8/25. N.E. Heard; 8.4.74; 52°59,4’S 73°38,0’E; 90 m: Modeeria rotunda, Staurotheca —
dichotoma, Symplectoscyphus elongatus, S. subdichotomus, Schizotricha unifurcata.

9/26. Heard, Atlas Cove; 8.4.74; 15-20 m: Sertularella picta, Schizotricha unifurcata.

9/27. Heard, Atlas Cove; 8.4.74; 6 m: Sertularella picta.

10/30. S.S.E. MacDonald; 9.4.74; 53°06,7’S 72°50,1’E; 255 m: Modeeria rotunda, Symplecto-
scyphus mawsoni, Schizotricha unifurcata.

11/31. W. Heard; 9.4.74; 53°20,3’S 72°29,2’E; 790 m: Staurotheca dichotoma.

14/44. W. Kerguelen; 13.4.74; 49°48,4’S 64°57,9’E; 250 m: Symplectoscyphus plectilis,
S. subdichotomus.

14/45. W. Kerguelen; 13.4.74; 49°45,8’S 64°50,6’E; 262 m: Hydrodendron arborea, Stauro-
theca dichotoma, Symplectoscyphus subarticulatus.

17/50. N.W. Kerguelen; 14.4.74; 47°24,9’S 66°04,0’E; 585 m: Kirchenpaueria triangulata,
Plumularia insignis.

18/52. N.W. Kerguelen; 15.4.74; 47°42,2’S 68°07,1’E; 243 m: Modeeria rotunda, Plumu-
laria insignis.

21/57. N.E. Kerguelen; 15.4.74; 48°29,7’S 70°55,4’E; 345-360 m: Zygophylax crozetensis,
Staurotheca dichotoma, Symplectoscyphus subdichotomus, Plumularia insignis.

22/58. N.E. Kerguelen; 16.4.74; 48°58,5’S 70°51,1’E; 90-105 m: Eudendrium rameum,
Halecium tenellum, Zygophylax crozetensis, Staurotheca dichotoma, Sertularella picta,
Symplectoscyphus subdichotomus, Schizotricha unifurcata.

23/59. S.E. Kerguelen; 16.4.74; 49°59,2’S 70°01,9’E; 158 m: Eudendrium rameum, Stauro-
theca dichotoma, Plumularia insignis, Schizotricha unifurcata.

24/61. S.E. Kerguelen; 17.4.74; 50°10,7’S 69°48,7’E; 195 m: Tulpa diverticulata, Stauro-
theca dichotoma, Plumularia insignis.

26/63. Crozet, chenal des Orques; 20.4.74; 46°21,5’S 51°55’E; 230 m: Modeeria rotunda,
Phialel. chilensis, Halecium dufresneae, Filellum serratum, Zygophylax crozetensis,
Campanularia sp., Staurotheca dichotoma, Symplectoscyphus curvatus,
S. subdichotomus.

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES 3

26/64. Crozet, chenal des Orques; 20.4.74; 46°24’S 51°59’E; 180 m: Eudendrium rameum,
Modeeria rotunda, Phialella chilensis, Hydrodendron arborea, Grammaria abietina,
Halisiphonia ?nana, Hebella striata, Zygophylax crozetensis, Campanularia sp., Tulpa
diverticulata, Staurotheca antarctica, S. dichotoma, Sertularella picta, Symplecto-
scyphus curvatus, S. elongatus, S. subdichotomus, Oswaldella bifurca, Plumularia
insignis.

26/65. Crozet, chenal des Orques; 21.4.74; 46°23,3’S 51°58,3’E; 165 m: Symplectoscyphus
subdichotomus.

28/71. Entre Possession et Cochons; 22.4.74; 46°18,1’S 51°29’E; 400 m: Halecium deli-
catulum, H. jaederholmi, Sertularella picta, Symplectoscyphus subdichotomus.

30/73. Entre Possession et Cochons; 22.4.74; 46°02,3’S 50°50,2’E; 187 m: Tulpa diverticu-
lata, Staurotheca antarctica, S. dichotoma, Symplectoscyphus subdichotomus, Plumu-
laria insignis.

31/74. Entre Possession et Cochons; 22.4.74; 45°57,2’S 50°32,8’E; 110 m: Modeeria rotunda,
Phialella chilensis, Halecium jaederholmi, Hydrodendron arborea, Grammaria abietina,
Zygophylax crozetensis, Silicularia rosea, Tulpa diverticulata, Staurotheca antarctica,
S. dichotoma, Sertularella geodiae, S. picta, Symplectoscyphus subdichotomus.

SYSTEMATIC SECTION
Family Eudendriidae
Eudendrium rameum (Pallas, 1766)

Eudendrium rameum: Allman, 1888: 4, pl. 2 (figs 1-2). Vervoort, 1946: 150, figs 60-61.
Vervoort, 1972a: 21.

Stations
22/58-C; 23/59-E; 26/64-P. All fertile.

Description

Stiff, bushy colonies with thick, fascicled stems reaching a maximum height
of 120 mm and a diameter of 5 mm at base, branching irregularly and in all
planes. Maximum diameter of unfascicled stems 0,26 mm. Perisarc annulated
on origin of branches, partly or completely on hydranth pedicels, and at other
irregular intervals, brown in colour. Hydranth with about 20 tentacles, with
abundant large nematocysts on hypostome and in nettle ring.

Male gonophores borne on non-atrophied hydranths, one- to two-
chambered. Female gonophores borne on non-atrophied hydranths, young
ones with unbranched spadix.

Large nematocysts: microbasic euryteles very similar to those illustrated
by Weill (1934, fig. 66); 19,8 x 9,0 — 25,8 x 10,8 um, discharged shaft 20,4-24,0
pm.

Remarks

Totton (1930) considered that all Antarctic material of Eudendrium should
be included in one species: E. antarcticum Totton = E. tottoni Stechow.
Totton included Jaderholm’s (1905) material from South Georgia doubtfully
in this species; and it presumably follows that Allman’s (1888) material from

4 ANNALS OF THE SOUTH AFRICAN MUSEUM

Kerguelen Island should be included too, since Jaderholm said that the two
samples were very similar.

In the present collection, however, there are almost certainly two species
of Eudendrium, a delicate form with a flexuous stem (E. fottoni) and a stiff
bushy form which the author has assigned to E. rameum. Unfortunately the
former is not well enough preserved for examination of nematocysts, which
must provide the final evidence, but the latter possesses abundant large micro-
basic euryteles, a type which according to J. Bouillon (personal communication)
is characteristic of E. rameum. The bushy colony is also very similar to Ver-
voort’s (1946) illustration of E. rameum.

Eudendrium tottoni Stechow, 1932

Eudendrium ramosum: Vanhoffen, 1910: 288, fig. 13.
Eudendrium antarcticum Totton, 1930: 140.
Eudendrium tottoni: Rees & Thursfield, 1965: 62.

Stations
3/10-E; 3/11—A. (Both fertile.)

Description

Delicate colonies reaching a height of about 16 mm. Stems unfascicled
or weakly fascicled at base, flexuous, difficult to distinguish from hydrorhiza,
branching irregularly, about 0,08 mm in diameter. Groups of annulations —
present on origins of branches and rarely at other points. Hydranths poorly
preserved.

Female gonophores with unbranched spadix arching over a single egg.
Male gonophores one- to two-chambered.

Remarks

The hydranths in this species are too poorly preserved to distinguish
structure or number of tentacles, though the gonophores have survived. The
tentacles of the fertile hydranths appear to be persistent. E. tottoni is known
from a number of localities in the Antarctic.

Family Campanulinidae

Modeeria rotunda (Quoy & Gaimard, 1827)

Stegopoma fastigiatum: Vervoort, 1972a: 42.
Modeeria rotunda: Edwards, 1973: 573, figs 1-3. Millard, 1975: 137, fig. 45A.

Stations
3/10—-C; 3/11—G; 8/25-D; 10/30-C; 18/52-B; 26/63-B; 26/64—G; 31/74-M.
Description

Many infertile colonies epizootic on other hydroids. Hydrotheca and
pedicel extremely variable in size.

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES 5

Remarks
This is a new record for this area. The species has, however, been recorded
from the south-western Atlantic at a very similar latitude (Vervoort 1972a).

Opercularella belgicae (Hartlaub, 1904)
Figure 1B

Campanulina belgicae Hartlaub, 1904: 10, pl. 1 (figs 8-9). Vanhoffen, 1910: 308, fig. 28.
Opercularella belgicae: Leloup, 1974: 4, fig. 2.
Opercularella sp. Vervoort, 1972a: 42, fig. 11b, c.
Station

3/10-F.
Description

A sparse colony without gonothecae growing on a polyzoan. Colony
stolonic. Pedicels of very variable length, with 2-4 spiral annulations at base,
merging smoothly into hydrotheca. Hydrotheca deep, widest at opercular base,
with about eight opercular segments and a delicate diaphragm.

Measurements (mm)

Total height, pedicel + hydrotheca .. - 1,00-6,90

Hydrotheca, height from diaphragm to tip oF eseetge a 0,46-0,57

maximum diameter As ie + Ae re 0,16—-0,20
Remarks

Vervoort’s material (1972a: Opercularella sp.) from Magellan is very
similar to the present material, though with shorter pedicels. It appears to fall
within the range of variation of O. belgicae.

The generic name of this species is still provisional pending knowledge of
the gonophore.

Phialella chilensis (Hartlaub, 1905)
Figure 1A

Campanulina chilensis Hartlaub, 1905: 589, fig. L?, M?, N?. Naumov & Stepaniants, 1962:
76, fig. 3. Leloup, 1974: 3, fig. 1.
Phialella chilensis: Vervoort 1972a: 38, fig. 10.

Stations
26/63—H; 26/64-Q; 31/74_N.
Description

Slender stems, a few lightly fascicled at base, branching sympodially and
reaching a maximum height of 10,8 mm. Stem closely annulated in basal half
of each internode and smooth in distal half, as illustrated by Vervoort (1972a)
rather than by Hartlaub (1905). Hydrothecal pedicels completely annulated
or with a smooth area in the centre of the longer ones. Hydrotheca as in previous
descriptions. Gonothecae absent.

6 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 1.

Phialella chilensis (Hartlaub). A. Part of stem with hydrothecae.
Opercularella belgicae (Hartlaub). B. hydrothecae.
Halecium delicatulum Coughtrey. C. Hydrophore. D. Gonophore.
Scale in mm/10.

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES a

Measurements (mm)

Hydrotheca, height from diaphragm to tip of operculum .. 0,21-0,29
maximum diameter oF a ae B ae 0,11-0,14

Remarks

This species is well known from the Antarctic and Subantarctic, especially
from the South American area. This is the first record from the Kerguelen
region.

Family Haleciidae

Halecium delicatulum Coughtrey, 1876
Figure 1C-—D

Halecium delicatulum: Ralph, 1958: 334, figs 1le, h-n, 12a—p. Naumov & Stepaniants, 1962:
94, fig. 16. Vervoort, 19725: 341, fig. 2a.

Halecium antarcticum Vanhoffen, 1910: 317, fig. 34. Billard, 1914: 7, fig. 5. Totton, 1930:
144, fig. 4.

Station
28/71-B.

Description

A number of stems growing on a gorgonian and reaching 32 mm in height.
Stem fascicled and stiff, branching irregularly. Internodes separated by oblique
nodes sloping in alternate directions, with one or two annulations at the base
of each. Primary hydrophores free from stem, smooth, sometimes containing
a pseudodiaphragm. Secondary hydrophores annulated, at least in basal region.
Hydrothecal margin strongly flared outward.

Female gonothecae present, compressed, oval in broad view, with terminal
aperture surrounded by an internal collar of perisarc, containing 6-10 eggs.

Measurements (mm)

Internode length .. a i ed a br se 0,53-0,72
Hydrotheca, depth to diaphragm m oe . 0,03-0,06
diameter at margin ee a sia sis sat 0,17-0,20
Gonotheca, length ae 7. a vs ae 0,71-1,12
maximum diameter ee oe es aes as 0,35—0,52
Remarks

The author has bowed to the opinion of Naumov & Stepaniants (1962)
and Vervoort (19725) in including H. antarcticum in the synonymy of H. deli-
catulum, although she feels that there is a distinct Antarctic form in which the
female gonotheca is smaller and more slender and without the ‘ears’ of typical
H. delicatulum as illustrated by Millard (1975). The gonothecae in the present
material are similar to those illustrated by Totton (1930) and Vervoort (19725).

8 ANNALS OF THE SOUTH AFRICAN MUSEUM

H. delicatulum is well known from the Antarctic, particularly in the area
south of America, and was reported from Marion Island by Allman (1888,

as H. flexile).

Halecium dufresneae sp. nov.
Figure 2A—D

Material

Holotype: from Station 26/63-A. Part in Muséum National d’Histoire
Naturelle, Paris, and part in the South African Museum (SAM-H2785).

Description of holotype

One colony, with a thick rootstock flattened below and a thick fascicled
stem 245 mm in height. Stem branching and rebranching irregularly but mainly
in one plane. Final branches unfascicled, divided into internodes by oblique
nodes sloping in alternate directions, each internode bearing a hydrotheca
from an apophysis at distal end.

Primary hydrophore not clearly demarcated from apophysis, with a stout
pseudodiaphragm in base, adnate to internode almost to diaphragm. Secondary
hydrophores rather rare in this material, smooth and not constricted after
origin, asymmetrical with adcauline wall longer than abcauline, with or without
a pseudodiaphragm. Hydrotheca shallow, with adcauline surface free from _
stem, with margin usually everted.

Gonothecae borne on sides of hydrophores, the two series together forming
a single row on the anterior surface of the stem. Gonotheca flattened, in broad
view widening to the truncated distal end, with one or two large embryos
discharged into a marsupium which may be as large as, or larger than, itself.

Measurements (mm)

Internode, length Ag ce cP <. = ie 0,51-—1,00
Hydrotheca, depth from diaphragm .. Pn i se 0,03-0,07
diameter at margin ae os ch es a 0,19-0,23
Gonotheca, depth =f = ue tA - a 0,94-1,14
maximum diameter ae es si if te 0,38—0,64
Marsupium, depth co se ie . =e 52 0,66-1,14
Remarks

Female gonothecae with external marsupia are known from two other
species of Halecium, namely H. pallens Jaderholm, 1904, from South Georgia,
and H. marsupiale Bergh, 1887, from the Arctic.

The author feels that in these two, as in the present species, the marsupium
is an important diagnostic character, and she cannot agree with Naumov &
Stepaniants (1962) and Vervoort (19725) who include H. pallens in the synonymy
of H. delicatulum. A marsupium has never been seen in the latter; moreover

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES 9

Fig.2:

Halecium dufresneae sp. nov. from the holotype. A-—C. Parts of stem with hydrophores:
C with secondary hydrophore. D. Gonophore with two embryos in marsupium.
Hydrodendron arborea (Allman). E. part of glomulus with female gonophores. F. Part of stem
with hydrophores and a giant nematotheca on bottom left. G. normal nematothecae.

Scale in mm/10.

10 ANNALS OF THE SOUTH AFRICAN MUSEUM

the shape of the mature female gonotheca is different, being wider in broad
view and ‘eared’ in H. delicatulum.

It follows that it is not possible to identify most of the species of Halecium
from sterile material alone.

The present species differs from H. pallens in the shorter pedicel of the
primary hydrophore and in the marsupium, which is almost as large as, or
larger than, the gonotheca and contains only one or two eggs.

Halecium jaederholmi Vervoort, 1972
Figure 3A—B

Halecium jaederholmi Vervoort, 1972a: 21, fig. 2 (synonymy).

Stations
28/71-A; 31/74-A.

Description

The first colony (28/71—A) is medium-brown in colour, with thick fascicled
stems 4 mm in diameter at the base and 160 mm in maximum height, branching
irregularly and in all planes. The second colony (31/74-A) is dark brown and
has an even thicker main stem (8 mm diameter at base) and is more stiff and "
rigid in habit. It reaches a height of 140 mm.

In both colonies the unfascicled parts together with their internodes and
hydrophores have a structure exactly like that illustrated by Vervoort (1972a)
except that no pseudodiaphragmata are present nor any secondary hydrophores.
The hydrothecae are very shallow and adnate to the internodes.

Typical female gonothecae are present in the first colony and contain up
to six larvae; some of the gonothecae are damaged and broken off transversely
just beyond the aperture. In the second colony all the gonothecae (female) are
damaged in this way.

Measurements (mm)

Internode, length ae he a ae Ea 0,50-0,94
Hydrotheca, depth to diaphragm a o is aoe 0,03-0,04
diameter at margin on < a os a 0,17-0,23
Gonotheca, length Re i ae ac sa ae 1,49-1,85
maximum diameter. . a ae 3 fs a 0,59-0,71
Remarks

The distribution of this species has been summarized by Vervoort; it occurs
in the Antarctic and Subantarctic, but this is the first record from the Kerguelen
area.

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES 11

Halecium tenellum Hincks, 1861
Halecium tenellum: Millard, 1975: 156, fig. 5SOF—L. Cornelius, 1975: 409, fig. 12.

Station
22/58—D.

Description
Colony epizootic on a polyzoan, with sparsely branched stems reaching
8 mm in height, and with female gonophores.

Remarks
This cosmopolitan species has been reported from the south-western
Atlantic and from Antarctica. This is the first record from the Kerguelen area.

Hydrodendron arborea (Allman, 1888)
Figure 2E-G

Halecium robustum Allman, 1888: 10. Vervoort, 1972a: 23.
Halecium arboreum Allman, 1888: pl. 4. Hickson & Gravely, 1907: 27, pl. 4 (figs 27-29).
Ophiodissa arborea: Totton, 1930: 142, fig. 2a. Vervoort, 1972a: 25.

Stations
14/45-B; 26/64-N; 31/7-J. All colonies fertile.

Description

Shrubby colonies with thick, fascicled stems reaching a maximum height
of 105 mm. Branching very irregular; final branches divided into internodes
by oblique nodes slanting in alternate directions. Each internode bearing one
hydrophore on an apophysis at distal end. Primary hydrophore not distinctly
demarcated from apophysis, adnate to internode almost, or completely, to level
of diaphragm.

Primary hydrotheca with free adcauline wall, with margin sometimes
everted on adcauline side. Secondary hydrophores usually quite symmetrical,
with no constriction after origin.

Nematothecae goblet-shaped, normally one on each internode on side
opposite to hydrotheca, but often broken off leaving only the aperture, and
sometimes absent without trace; also present on tubes of gonosome.
Occasionally giant nematothecae present, about five times the size of normal
ones.

Gonothecae borne in glomulus (term from Naumov 1960: 442), strongly
curved, with tubular neck, containing eggs or planulae. Glomuli forming large
masses about 15 mm in diameter.

Measurements (mm)

Internode, length “a fie 43 a! . is 0,42-0,93
Hydrotheca, depth to diaphragm of ae ae d's 0,04—0,07
diameter at margin we ae a Re ot 0,21-0,28

12 ANNALS OF THE SOUTH AFRICAN MUSEUM

Measurements (mm)

Normal nematotheca, depth .. a ie ae i 0,06-0,10
Giant nematotheca, depth i o ee ue a 0,44-0,48
Gonotheca, maximum length .. a : oe a 0,78-1,02
maximum diameter nts oe 5A . os 0,47—0,60
Remarks
This species is widely distributed in the Antarctic; its type locality is
Kerguelen.

As other authors have remarked, the nematothecae are variable in occur-
rence and not invariably present on each internode. The’ giant nematothecae
have not been recorded before. The regenerated (secondary) hydrophores are
very distinctive and clearly distinguish the species from forms such as H. beanii.

Family Lafoeidae

Filellum serratum (Clarke, 1879)
Filellum serratum: Vervoort, 1972a: 51, fig. 14a—b. Millard, 1975: 178, fig. S9YA—C.

Stations
3/11-F; 26/63-J: infertile colonies epizootic on polyzoa and other hydroids.

Remarks

This cosmopolitan species has not yet been reported from the Kerguelen
area, though it is known from New Zealand and South America.

Grammaria abietina (Sars, 1850)
Figure 3D-E
Grammaria abietina: Cornelius, 1975: 382, fig. 3 (synonymy).

Stations
26/64-K; 31/74-L.

Description

Two colonies with thick, fascicled stems reaching a maximum height of
83 mm and branching in an alternate, subopposite or opposite manner in one
plane. Hydrothecae forming six longitudinal rows on stem and branches, but
not strictly regular in arrangement.

Hydrotheca curved outwards to a varying degree, with margin everted
and either parallel or oblique to axis of stem; free part 0,34-0,51 mm in length;
0,28-0,35 mm in diameter at margin.

A single coppinia present, about 15 mm iong and 5 mm wide. Gonothecae
tightly adpressed, flask-shaped, with terminal aperture on a short tubular
neck. Accessory tubes about three times length of gonothecae, very strongly
curved.

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES 13

Bue

’
ee ee ee ee ee oe |

Fig. 3.

Halecium jaederholmi Vervoort. A. Part of stem with two hydrophores. B. Gonophore.
Halisiphonia ?nana Stechow. C. Hydrothecae.
Grammaria abietina (Sars). D. Part of stem with six rows of hydrothecae. E. Part of t.s. of
coppinia, showing gonothecae, protective structures, and arrangement of thecal bases in
three’s in centre of stem.

Scale in mm/10.

14 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks

The synonymy of this species was recently revised by Cornelius (1975),
who has included as synonyms G. stentor Allman, 1888, G. magellanica Allman,
1888, and G. insignis Allman, 1888.

The species has a bipolar distribution and is widely known from the Arctic
and Subarctic as well as from the Antarctic and Subantarctic. The type locality
of G. stentor is Kerguelen Island.

Halisiphonia ?nana Stechow, 1921
Figure 3C
Halisiphonia nana: Stechow, 1925: 452, fig. 22.

Station
26/64-S.

Description

An infertile colony epizootic on Sertularella picta. Pedicel arising from
hydrorhiza and merging smoothly into hydrotheca which widens evenly to
margin. No diaphragm or annular thecal thickening. Margin sometimes slightly
everted.

Measurements (mm)

Pedicel + hydrotheca, height .. ab iby Es ba 0,64—0,95

Pedicel, diameter at base we ee a me ae 0,04—0,12

Hydrotheca, diameter at margin ne ae Ee - 0,14-0,23
Remarks

This material is assigned with some doubt to H. nana. The dimensions are
more or less in agreement with those of Stechow, and are too small for H. mega-
lotheca Allman, but the pedicel is often rather wide at the base and the thecal
margin may be everted. H. nana has been reported only once, from east of
Bouvet Island; the gonothecae are unknown.

Hebella striata Allman, 1888

Hebella striata Allman, 1888: 30, pl. 15 (figs 3, 3a). Vanh6offen, 1910: 313. Vervoort, 1972a:
62, fig. 17b-c.

Station
26/64-R.

Description

An infertile colony epizootic on Grammaria abietina. Details exactly as in
Vervoort’s material, but dimensions a little larger.

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES iS

Measurements (mm)

Pedicel, length .. me cS ce i a es 0,44-0,87

Hydrotheca, depth we os ce ee na eS 0,77-1,10

diameter a Ra ae ba a os 0,24-0,31
aks

This species is known mainly from the Subantarctic near South America,
but was reported from Kerguelen by Vanhdffen (1910).

Lafoea dumosa (Fleming, 1820)

Lafoea fruticosa: Millard, 1975: 187, fig. 61A—F.
Lafoea dumosa: Cornelius, 1975: 385, fig. 4 (synonymy).

Stations
3/10—B; 3/11-B. Both colonies infertile.

Description

Stems flexuous, reaching a maximum height of 78 mm, with many
fragments.

Remarks

The author has followed Cornelius (1975) in uniting L. fruticosa (M. Sars,
1851), L. gracillima (Alder, 1856) and L. dumosa under the last name.

Zygophylax crozetensis sp. nov.

Figure 4
Material

Holotype: from station 26/64-B. Part in Muséum National d’Histoire
Naturelle, Paris, and part in the South African Museum (SAM-H2779).

Other stations: 2/6-A (infertile); 21/56—-C (infertile); 22/58—B (male);
26/23-D (fertile); 31/74-F (fertile).

Description of holotype

A large, branching colony 220 mm in height. Rootstock a large mass of
interwoven fibres flattened below and reaching 45 mm in diameter. Stem
fascicled and thick, giving off thick primary branches in an irregular fashion
and mainly in one plane, these branches giving off secondary and tertiary
branches which are strictly in one plane. Final branches (hydrocladia) sub-
opposite in arrangement, lightly fascicled or unfascicled. Hydrocladia and
final terminations of other branches segmented where exposed, with one or two
hydrothecae to an internode, but with many irregular regenerative nodes.
Hydrothecae normally alternate in arrangement with the two rows more or less
in one plane, and this arrangement characteristic of unfascicled hydrocladia

16 ANNALS OF THE SOUTH AFRICAN MUSEUM

a is B ‘<a

Fig. 4.

Z. ygophylax crozetensis sp. nov. from the holotype. A. Part of stem showing origins of hydro-
cladia. B-C. Hydrothecae and nematothecae. D. Surface view of coppinia showing hoods
of gonothecae and protective structures. E. Gonothecae from t.s.

Scale in mm/10.

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES lg

and smaller branches, but in fascicled branches supplementary hydrothecae
may arise from peripheral tubes in any plane and in any position imparting a
very irregular appearance to the whole.

Hydrothecae tubular, widening to margin which is slightly everted, curved
outwards, those in axils of hydrocladia less curved and practically straight.
Diaphragm distinct and oblique.

Nematothecae two-chambered, with a short basal chamber and a long
tubular distal chamber, one or two on each hydrotheca-bearing apophysis,
that/those of the axillary hydrotheca shifted on to the apophysis next to it,
various irregular supplementary ones present on peripheral tubes of stem and

branches.
Coppiniae numerous and clothing most of the larger branches completely

to a width of 3-5 mm, so that it is not possible to distinguish one from another.
Coppinia consisting of adpressed gonothecae with between them irregular
branching structures bearing nematothecae and rarely hydrothecae as well.
Gonotheca slender, widening distally to top of adnate part, then narrowing
to a free, pointed, and sometimes curved, hood or horn bearing an aperture on
one side. Sex not determinable.

Measurements of holotype (mm)

Hydrothecal pedicel, height .. e se ie is 0,06-0,13

Hydrotheca, height abcauline .. m, oe ve : 0,32-0,38

diameter at margin ye os fe os S 0,16—0,20

Pedicel + hydrotheca, height .. in ¥ Ms bes 0,40-0,47

Nematotheca, height .. = oe & ye — 0,09-0,19

diameter at margin m we a Ee ce 0,05-0,08
Remarks

In the four fertile samples all the gonothecae are of the same shape. Most,
however, are empty. Contents are present in 22/58—B only, and these are male.

The hydrothecae of this species are similar to those of Z. africana Stechow,
1923, and a number of related species, and particularly to ‘Lafoea’ halecioides
Allman, 1874 (= ‘Lafoea’ pinnata Sars, 1873). The last-mentioned, however,
has an open coppinia with unfused gonothecae.

Nutting (1905) described and figured a closed coppinia with one-horned
gonothecae for a species which he called Lictorella halecioides. This is apparently
a synonym for Zygophylax antipathes (Lamarck, 1816), and not for Z. pinnata.
It is probably this account which prompted Totton (1930: 166) to mention
‘hooded apertures’ for Z. antipathes and not the account of Trebilcock (1928)
who does not mention the species. However, Z. antipathes has somewhat
broader hydrothecae than the present material, and the gonothecae as depicted
by Nutting have terminal apertures.

It appears that a number of southern hemisphere species of Zygophylax
have similar trophosomes but different gonosomes. The gonothecae of the

18 ANNALS OF THE SOUTH AFRICAN MUSEUM

present species are almost exactly like those of Cryptolaria pectinata (Allman,
1888).

Family Campanulariidae

Campanularia norvegiae Broch, 1948
Figure 5A-E
Campanularia norvegiae Broch, 1948: 16, fig. 5.

Station
8/24-E.

Description

A dense fertile colony growing on an empty tube. Hydrorhiza reticular.
Pedicel sometimes spirally grooved throughout, but very variable and often
smooth in certain areas, always with a segment of lesser diameter at distal end.
Hydrotheca funnel-shaped, oval in section and thickened, more so at the narrow
ends; thickened at, or just below, margin.

Gonotheca irregularly bottle-shaped, with a short annulated pedicel,
narrowing distally to terminal aperture, often twisted or irregularly corrugated,
circular in section, empty or containing planulae.

Measurements (mm)

Pedicel + hydrotheca, length .. xn ne a a 1,70—8,50
Hydrotheca, depth - oe oa ; 0,48-0,72
maximum diameter (broad view) .. oe = ee 0,38-0,56
Gonothecal pedicel, depth pe oe Le 3 7 0,24-0,32
Gonotheca, depth - ae as leg 2 ss 1,58—2,28
maximum diameter a > Le 433 sa 0,52-0,65
Remarks

This species is distinguished from the closely related C. integra Mac-
Gillivray, 1842, by the shape of the gonotheca. Since the gonothecae contain
planulae it is obvious that no medusoid is released.

C. norvegiae was originally described from South Georgia.

Campanularia sp.
Figure 5F
Stations
26/63-G; 26/64-R.

Description

Infertile colonies epizootic on other hydroids. Pedicels of variable length,
with one terminal spherule of lesser diameter, generally roughly corrugated

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES 19

Fig. 5.

Campanularia norvegiae Broch. A-C. Hydrothecae. D-E. Gonophores.
Campanularia sp. F. Hydrothecae.
Tulpa diverticulata Totton. G. Gonotheca. H. Hydrotheca.

Scale in mm/10.

20 ANNALS OF THE SOUTH AFRICAN MUSEUM

immediately below this and at base. Hydrotheca deep, with parallel sides, with
an annular thickening near base, with 8-10 slender, bluntly rounded marginal
teeth.

Measurements (mm)

Pedicel, length .. oe Ate on * we is 0,48-1,07
Hydrotheca, depth fs ie ss a ah we 0,60—-0,90
diameter at margin Ke eS bs ave eo: 0,25—0,33
diameter/depth a: oe - e. i. mc 0,34-0,51
Remarks

In the absence of gonophores this species cannot be definitely diagnosed.
The two most commonly reported species of Campanularia from the Antarctic
are C. hicksoni Totton, 1930, and C. tincta Hincks, 1861. (C. cylindrica Allman,
1876, from Kerguelen is probably a synonym of the former.) These two species
are distinguished mainly by their gonothecae—smooth in the former, annulated
in the latter; however, none of the Antarctic material attributed to C. tincta has
yet been found with gonothecae and possibly only one species is involved.

Silicularia rosea Meyen, 1834

Silicularia rosea: Ralph, 1956: 293. Millard, 1968: 259. Millard, 1971: 405.
Silicularia bilabiata: Ralph, 1956: 285, figs 2-3. Ralph, 1957: 842.

Station
31/74-C.

Description

A rich, fertile colony on brown alga.

Remarks

S. rosea is known from Crozet and Kerguelen Islands and many other
localities in the Antarctic and Subantarctic. The distribution was summarized
by Ralph (1957).

Tulpa diverticulata Totton, 1930
Figure 5G-H

Tulpa diverticulata Totton, 1930: 145, fig. 5. Ralph, 1957: 844, fig. 7l-n.
Campanularia diverticulata: Naumov & Stepaniants, 1962: 72.

Stations
24/61-C; 26/64-H; 30/73-A; 31/74-B.
Description

Hydrorhiza creeping on other hydroids, polyzoa and worm tubes, but
often becoming free to form a tangled mass. Hydrothecal pedicels of very

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES 21

variable length, unsegmented, though often with regeneration nodes. Hydro-
thecae as in previous descriptions; regenerated margins sometimes present.
Gonothecae arising from hydrorhiza on very short pedicels, irregularly oval,
with terminal aperture on long narrow neck.

Measurements (mm)

Hydrothecal pedicel, length .. A a i i! 0,60—2,80
Hydrotheca, length 7: ne a a Se nfs 3,00-4,10
diameter at margin ee a eS ie - 0,95-1,35
Gonotheca, length including pedicel .. i sd ef 1,85—3,45
maximum diameter oe ne oe s a 0,89-1,50
Remarks

Some of these colonies are very rich, and several stolons may run together
or twine round each other simulating a fascicled stem. However, the pedicels
always arise at right angles, and there is never any question of an erect branching
stem with oblique pedicels as in 7. tulipifera (Allman, 1888). It is clear that
these are two separate species.

Very fine cross-striations may be present on some of the hydrothecae and
gonothecae in certain colonies.

Young gonothecae are cone-shaped and similar to those described by
Fraser (1944) for T. speciosa (Clarke).

T. diverticulata is so far known only from New Zealand and to the south
of it. It is a new record for Kerguelen and Crozet Islands. The closely related
T. tulipifera occurs in Heard Island (Allman 1888) and in the south-western
Atlantic (Vervoort 1972a).

Family Syntheciidae
Staurotheca antarctica Hartlaub, 1904

Staurotheca antarctica Hartlaub, 1904: 16, pl. 1 (fig. 4), pl. 2 (fig. 4). Totton, 1930: 176, 178,
fig. 28, pl. 2 (fig. 6). Vervoort, 1972a: 198, figs 67, 68b.

Stations
26/64-M (fertile); 30/73-B (fertile); 31/74-D.

Description

Luxurious colonies with unfascicled stems which branch dichotomously
and reunite to form an elaborate reticulum in one plane. Nodes irregular.
Hydrothecae generally arranged in decussate pairs, but rarely in groups of three,
and with many irregularities.

Male and female gonothecae present, as in previous descriptions.

22 ANNALS OF THE SOUTH AFRICAN MUSEUM

Measurements (mm, all without regenerations)

Hydrotheca, depth abcauline
depth adcauline, adnate part
depth adcauline, free part ..
adnate part/total adcauline depth
diameter at mouth

Gonotheca, male, length
maximum diameter

Gonotheca, female, length
maximum diameter (including spur)

Remarks

0,34—0,44
0,38-0,50
0,10—0,20
0,66-0,83
0,17-0,20
0,84-1,12
0,55-0,66
1,30-1,48
0,98-1,21

There is little to add to the description of this well-known Antarctic and
Subantarctic species. Vervoort (1972a) has summarized the distribution.

Staurotheca dichotoma Allman, 1888

Staurotheca dichotoma Allman, 1888: 76, pl. 36 (fig. 1). Billard, 1910: 27, figs lus 12. Billard,

1914 >-15, ne. 9 votton, 1930: 175, 178.tie. 27;

Stations

7/22-B; 8/24-A (fertile: female gonophores); 8/25-B (fertile: male and
female gonophores); 11/31-A; 14/45—A; 21/57-B (fertile: female gonophores); —

22/58—G; 23/59-A (fertile: male gonophores); 24/61-B;

-C; 26/64-L;

30/73-C (fertile: female gonophores); 31/74—-E (fertile: female gonophores).

Description

Luxuriant colonies. Stems thick, fascicled at base in larger colonies,

branching in a subdichotomous manner, reaching a maximum height of 130 mm.
Hydrothecae generally 3 to a whorl (and forming 6 longitudinal rows), but
some of the thickest stems have 4 to a whorl (and 8 longitudinal rows) and the
thinner terminal branches only 2 (4 longitudinal rows). Hydrothecae adnate
for almost their entire length. Male and female gonothecae as described by
Billard (1910) and Totton (1930).

Measurements 8/25—B (mm)

Hydrotheca, depth abcauline .. ay ah ae er 0,59-0,67
depth, adcauline, adnate part ri a » me 0,78—-0,96
depth adcauline, free part . ; Me i bo 0,03-0,08
adnate part/total adcauline tent os as Bo 0,91-0,97
diameter at margin i: a me e es 0,27-0,32

Gonotheca, female, length a Ay a% Fe oa 1,49-2,17
maximum diameter a oe a ay %y 1,11-1,29

Gonotheca, male, length oe so oe a ae 1,23-1,60

maximum diameter ae 9 ai m1 i 0,78—1,02

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES 23

Remarks

As previous authors have remarked, S. dichotoma is clearly distinguished
from S. antarctica by the structure of the female gonotheca and by its greater
dimensions. The species is widely distributed in the Antarctic and Subantarctic.
Its type locality is Marion Island, not far from the present localities.

Family Sertulariidae

Sertularella geodiae Totton, 1930
Figure 6E-F

Sertularella geodiae Totton, 1930: 196, fig. 43, pl. 3 (figs 7-8). E. A. Briggs, 1939: 37. Ralph,
1961: 831, fig. 24c, g. Vervoort, 1972a: 120, fig. 37.

Station
31/74-K.

Description

A number of infertile stems and fragments, reaching a maximum height
of 40 mm. Stem stiff, fascicled and straight at base, geniculate in terminal
regions only, branching rather sparsely and in one plane. Branches unfascicled,
arising below hydrothecae and, when several are present, below every third
hydrotheca.

Hydrotheca adnate for about half adcauline wall, bent outwards (a perpen-
dicular dropped through centre of margin passes through adcauline wall), with
a few undulations on free part of adcauline wall, with wide mouth. No internal
teeth.

Measurements (mm)

Internode length .. mn re io e ie ode 1,01-1,52
diameter at node .. xe yd ee oe ae 0,20-0,32
Hydrotheca, length abcauline .. ser o Se ais 0,66—0,77
length adcauline, adnate part ne te on “i 0,49-0,61
length adcauline, free part a Se £e ae 0,46-0,60
adnate part/total adcauline length Bi se is 0,46-0,55
diameter at margin ay $3 ae ie ee 0,35-0,40
Remarks

This material is in many ways intermediate between S. geodiae and S. conica
Allman, 1877, considering especially Vervoort’s description of the latter (1972a:
123). However, S. conica has an unfascicled stem and a hydrotheca which
narrows more markedly to the mouth; its distribution is mainly tropical.
The author has therefore assigned this material to S. geodiae in spite of the
absence of gonothecae; this is a species known from New Zealand, Tasmania

24 ANNALS OF THE SOUTH AFRICAN MUSEUM

Ss

©

A, DF eet
B,C le
UO
Mm
“TI

Fig. 6.

Sertularella picta (Meyen). A. Part of stem showing origin of a hydrocladium. B—C. Hydro-
thecae. D. Gonotheca.
Sertularella geodiae Totton. E. Hydrotheca. F. Part of stem showing origins of two

hydrocladia.
Scale in mm/10.

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES 25

and the area around the southern tip of South America, but which has not been
reported from Kerguelen.

Sertularella picta (Meyen, 1834)
Figure 6A—D

Sertularella picta: Hartlaub, 1905: 645, fig. L*. Billard, 1922: 106, fig. 2B. Stechow, 1923:
187, fig. B’. Millard, 1971: 405, fig. 6A, B. Vervoort, 1972a: 111, figs 34, 35.

Stations
8/24-D; 9/26-A; 9/27-A; 22/58-E; 26/64-F; 28/71-D; 31/74-H. The
last three fertile.

Description

Stems slender and straggling, branching repeatedly in an irregular fashion
to produce tangled colonies often intertwined with polyzoans and with Symplec-
toscyphus subdichotomus. Stem normally unfascicled, but rarely with one or
two supplementary tubes in basal region, usually with one or two annulations
at base and on origin of branches, geniculate, reaching a maximum height of
38 mm, often reuniting by stolons with other stems. On most stems the two
rows of hydrothecae are in the same plane, but in other stems, or even in other
parts of the same stem, they may be shifted on to the anterior surface so that
the two rows subtend an obtuse angle between them. Branches arising below
hydrothecae, with one or two annulations at origin and a long first internode,
otherwise similar to stem and of same diameter. Internodes slender, very
variable in length, separated by oblique nodes sloping in alternate directions,
usually with an annulation immediately above each node.

Hydrotheca fusiform, adnate for less than half adcauline height, narrowed
below margin. In most hydrothecae the marginal teeth are equally developed
and the margin is perpendicular to the axis (a perpendicular dropped through
centre of margin passes through basal thickening of adcauline wall), but in
some the abcauline marginal tooth is produced so that the margin is tilted
towards the stem (and a perpendicular dropped through centre of margin
passes through the hydrophore or base of the abcauline wall). All intergradations
may occur in one colony. The free part of the adcauline wall usually has two to
four indistinct annulations, but the latter are of variable development—they
may form distinct striations which pass round the sides of the hydrotheca
almost to the abcauline wall, or they may be almost obsolete. Small internal
teeth are present in most hydrothecae, but again the degree of development is
variable; the full complement appears to be three (one median abcauline and
two latero-adcauline), but one or two, or even all three, may be absent. In a
few stems there is no trace of internal teeth.

Gonothecae borne on front of stem immediately below hydrothecae,
obovate, with three to four distinct annulations in distal half, with three or four
blunt marginal spines, with external marsupium. Sex not determinable.

26 ANNALS OF THE SOUTH AFRICAN MUSEUM

Measurements (mm)

Internode, length a - ot . a ay 0,77-1,34
diameter at node .. As ae R Me a 0,12-0,18
Hydrotheca, length abcauline .. - i vit a 0,55-0,70
length adcauline, adnate part ie a a i 0,24—0,35
length adcauline, free part i op ae ae 0,40-0,51
adnate part/total adcauline length * us fi 0,33-0,44
diameter at margin 2 ae os se e 0,20-0,26
Gonotheca, length a mn ne Se a mY 1,50-2,15
maximum diameter ve ee oF Ms Le 1,30—1,52
Remarks

The identification of this species has been based mainly on Vervoort’s
work (1972a), who also found considerable variability in structure. The form
of the colony, the shape of the hydrotheca and the gonotheca are very similar
to his material.

The species is known from the Falkland/Tierra del Fuego region, and was
also reported from Marion Island (Millard 1971). The Marion Island material
differs in the smooth hydrothecal wall and the greater displacement of the two
rows of hydrothecae.

Symplectoscyphus curvatus (Jaderholm, 1917)
Figure 7A—C

Sertularella curvata Jaderholm, 1917: 9, pl. 1 (figs 11-12).
Symplectoscyphus curvatus: Totton, 1930: 192, fig. 40, pl. 2 (figs 1-3). E. A. Briggs, 1939: 31.
Sertularella curvatus: Naumov & Stepaniants, 1972: 45, fig. 7.

Stations
26/63—-E; 26/64—-C. Both infertile.

Description

Many unrooted, unfascicled stems and fragments yellow-brown in colour
and reaching a maximum length of 44 mm. Most stems pinnate, geniculate,
giving off alternate branches at each elbow and immediately below every third
hydrotheca. On the proximal part of the stem the nodes are not clearly demar-
cated and the hydrothecae are well spaced and do not overlap, but on the
distal part and on the branches the nodes are distinct and the tip on one hydro-
theca overlaps the base of the next. The two rows of hydrothecae and branches
in one plane.

Hydrotheca tubular, large, adnate for less than half adcauline length,
curved outwards, not narrowing to mouth and with diameter in centre region
approximately equal to that at margin, marginal teeth well developed, no
internal teeth. No gonothecae.

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES 27

A
B
D

En
4 <p a
aS
cA Vy

Pes te

| Dd

Fig. 7.

Symplectoscyphus curvatus (Jaderholm). A. Distal part of hydrocladium. B. Hydrotheca.
C. Part of stem showing origin of hydrocladium.
Symplectoscyphus elongatus (Jaderholm). D. Part of stem showing origins of hydrocladia.
E. Hydrotheca.

Scale in mm/10.

28 ANNALS OF THE SOUTH AFRICAN MUSEUM

Symplecto-
scyphus
exsertus

Measurements (mm) 26/64-C (Allman)
Internode length .. si bs Ms o 0,80—1,32 0,43-0,60
Hydrotheca, length abcauline .. ne ao 0,62-0,70 0,28-0,32

length adcauline, adnate part a ms 0,36-0,44 0,14-0,17
length adcauline, free part a oe 0,59-0,68 0,22-0,28
adnate part/total adcauline length ie 0,35-0,41 0,35-0,44
diameter at mouth ee a re 0,41-0,49 0,12-0,14

Remarks

The measurements of the hydrothecae agree with those of Jaderholm and
Totton; they are a little greater than those of Briggs and Naumov & Stepaniants.

S. curvatus shows resemblances to S. exserta (Allman, 1888) from Heard
Island. However, the author has examined type material of the latter species
and finds that it is much smaller in all its dimensions. The measurements are
included above for comparison.

Symplectoscyphus elongatus (Jaderholm, 1904)
Figure 7D-E

Sertularia articulata Allman, 1888: 61, pl. 29 (figs 3, 3a).

Sertularella articulata: Jaderholm, 1905: 29, pl. 11 (fig. 4), pl. 12 (figs 1-3). Naumov & Stepani-
ants, 1972: 42, fig. 5A.

?Sertularella spiralis Hickson & Gravely, 1907: 19, pl. 3 (figs 19-20).

Sertularella elongata: Naumov & Stepaniants, 1962: 80, fig. 5. Naumov & Stepaniants, 1972:
42, fig. 5B.

Symplectoscyphus articulatus: Rees & Thursfield, 1965: 127.

Symplectoscyphus elongatus: Vervoort, 1972a: 136, figs 43, 44a (synonymy).

Stations
8/25—C; 26/64-D; 8/24-C.

Description

Three samples, the first with several infertile stems reaching 67 mm, the
second with numerous infertile stems and fragments reaching a maximum
height of 52 mm, and the third with numerous stems and fragments reaching a
maximum height of 58 mm. None of the stems rooted. Stems golden brown in
colour, unfascicled, geniculate. Internodes of stem very long, up to 3,3 mm,
each bearing a prominent apophysis on distal end on which is seated one
hydrotheca and on each side of it a hydrocladium, the apophyses with their
paired hydrocladia arranged in a loose spiral. Nodes very distinct.

Hydrocladia bearing alternate hydrothecae, and commonly alternate sub-
branches arising opposite every third hydrotheca, the two rows of hydrothecae
and branches usually in one plane but occasionally very slightly displaced

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES 29

towards one surface. Internodes of variable length, those of hydrocladium
bearing one or two hydrothecae, those of its sub-branches one.

Hydrothecae tubular, adnate for about half adcauline wall, abcauline wall
straight or curved slightly outwards, with three prominent marginal teeth, but
no internal teeth.

A few empty and damaged gonothecae present in 8/24-C, of the same
shape and size as described by Hickson & Gravely (1907).

Measurements (mm)

Hydrotheca, length abcauline .. bs mn ne im 0,39-0,52
length adcauline, adnate part a: se ns a 0,21-0,35
length adcauline, free part ie se = ce 0,18-0,36
adnate part/total adcauline length sd ae oe 0,37-0,65
diameter at mouth .. oe id i a ne 0,17-0,23

Remarks

This species has been reported many times from Antarctic and Subantarctic
waters; Kerguelen Island is the type locality. The synonymy of the species is
fully discussed by Vervoort (1972a). This synonymy is accepted here, though
with some reservations about the inclusion of Sertularella spiralis Hickson &
Gravely, 1907, a form with internal hydrothecal teeth.

Symplectoscyphus mawsoni Briggs, 1939
Figures 8—9

Non Sertularella biformis Jaderholm, 1905: 28, pl. 11 (figs 1-3).

Sertularella biformis: Totton, 1930: 199, fig. 45, pl. 2 (fig. 8). Naumov & Stepaniants, 1972:
47, fig. 9.

Symplectoscyphus mawsoni E. A. Briggs, 1939: 35, fig. 2, pl. 16 (figs 1-2). Naumov & Stepani-
ants, 1972: 46, fig. 8.

Stations
10/30-A; 8/24-B.

Description

The first colony (10/30—A) consists of two stems 56 mm in maximum
length, and a number of fragments. Basal part of stem lightly fascicled for
16-30 mm, the rest unfascicled and straight. Stem divided into internodes by
distinct oblique nodes sloping in alternate directions. Each internode with a
distal apophysis, alternately on the left and right, which bears a hydrotheca
and two hydrocladia, one on each side of the hydrotheca, the two rows of
hydrothecae in one plane. Many of the hydrocladia and hydrothecae of the
lower half of the stem missing or damaged, those in the upper half better
preserved. Each pair of hydrocladia forming an angle of about 40° with the
stem and about 90° with each other. Hydrocladium divided into regular inter-

30 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 8.
Symplectoscyphus mawsoni Briggs. A. Thick part of stem in side view showing origins of
hydrocladia. B. Hydrotheca. C. Gonotheca with external marsupium. D. Part of hydro-
cladium showing bifurcation. E. A hydrotheca from Totton’s material (BM 1929.10.28.158)
attributed by him to Sertularella biformis, here included in S. mawsoni.
Scale in mm/10.

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES Sif

nodes by straight nodes, each bearing a hydrotheca near the distal end; dividing
dichotomously at the distal end of the fourth internode, one limb of the dicho-
tomy being slightly thicker than the other; both limbs sometimes dividing
dichotomously once or twice more. The two rows of hydrothecae on the hydro-
cladium not in one plane but forming an acute angle between them. Hydrotheca
completely free from internode, very delicate and often crumpled in basal part,
curved outwards, widening towards aperture, with three well-developed mar-
ginal teeth, one adcauline and two latero-abcauline. No internal teeth. Oper-
culum of three valves. Floor similar to that of Sertularella (Fig. 8A, B, D;
Fig. 9 left).

The second colony (8/24—-B) is in a very fragmentary condition, consisting
mainly of small pieces of hydrocladia. There is, however, a stem of 110 mm
with a few stumps of hydrocladia left; it is fascicled for the first 45 mm. Another
portion of stem 46 mm long bears regular pairs of hydrocladia as in 10/30-A.
The hydrocladia branch up to five times. This colony is fertile and bears

Fig. 9.

Symplectoscyphus mawsoni Briggs. A stem from the Marion-Dufresne collection (Station
10/30—A) on the left, compared with Totton’s specimen attributed to Sertularella biformis
on the right (x 2,5). Photo: D. Gerneke.

32 ANNALS OF THE SOUTH AFRICAN MUSEUM

numerous gonothecae on the hydrocladial internodes. Female gonotheca
elongated oboval, with external marsupium, containing one planula (Fig. 8C).

Measurements (mm) 10/30-A 8/24-B
Stem, internode length .. ns a a 2,20-3,95 2,50—4,10
diameter oy = we ‘- - 0,52-0,70 0,50-0,62
Hydrocladium, internode length a Ae 0,57—1,09 0,67-1,15
Hydrotheca, depth abcauline .. a be 0,71-0,82 0,74-0,83
depth adcauline.... Y Re a 0,87—1,00 0,92-1,01
diameter at margin ae oe a 0,24-0,32 0,29-0,32
Remarks

Several species have been recorded from the Antarctic and Subantarctic
which resemble one another in their method of branching, aptly described by
Naumov & Stepaniants (1972) as ‘panicle-shaped’. These can be divided into
two groups:

1. Those where the hydrothecae are adnate for half or more of their height
and where the ‘panicle’ is flexuous: included under the synonymy of Symplecto-
scyphus elongatus by Vervoort (1972a) and in this paper.

2. Those where the hydrothecae have little or no adnate part and where
the ‘panicle’ is stiff.

It is the second group that concerns us here and it includes material recorded
under the names of Symplectoscyphus (or Sertularella) biformis and mawsoni.
Previous and present records of these species, together with the more important
measurements, are listed in Table 1 (the depth of the hydrotheca is not included
because of doubt as to exactly how it was measured by different authors).

The type material of Sertularella biformis Jaderholm, 1905, was examined
(Fig. 1OE-F), and Jaderholm’s description is confirmed. The material differs
from all other records in Table 1 in the absence of hydrocladia or any vestiges
thereof, in the absence of nodes in most of the stem, in the grouping of the
distal hydrothecae in three’s, and in the shape of the hydrotheca which narrows
towards the margin. Totton (1930) assigned material to this species (S. biformis)
assuming that the specimens ‘would correspond with the apical part missing
from Jaderholm’s type . . .’, and that the stem progresses in complexity from
base to apex, both assumptions being unwarranted. Totton’s material, on
examination, was found to differ from Jaderholm’s in all the characters listed
near the beginning of this paragraph (Fig. 8E, Fig. 9 right). It is clearly a different
species altogether. Jaderholm’s species has not been rediscovered to date, and
the type material remains the sole specimen.

The remaining records in Table 1 seem to be conspecific and to form a
series in the order shown, starting with the type material of Symplectoscyphus
mawsoni Briggs and ending with Totton’s material. In this series the complexity
of branching in the hydrocladia increases, and the diameter of the hydrotheca

33

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES

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[ aTav]

34 ANNALS OF THE SOUTH AFRICAN MUSEUM

decreases. The stem internodes increase in length except for Totton’s material
where they are relatively short and wide so that the hydrocladia are closer
together and give a dense ‘bottle-brush’ appearance to the colony. The material
of Naumov & Stepaniants (1972) attributed with a query to Sertularella biformis
differs from the rest in the absence of hydrothecae on the stem, but it is very
possible that these have been damaged or eroded away. (Totton omitted to
state that his material possesses cauline hydrothecae.)

The present material from the Kerguelen area lies about mid-way in the
series resembling Briggs’ material in the appearance of the colony and the
structure of the stem internodes, but resembling Totton’s material in the shape
of the hydrothecae.

It seems that the first five samples in Table 1 cannot be separated on any
definite characters and it is proposed to unite them under the name Symplecto-
scyphus mawsoni Briggs, 1939. The gonothecae were first described by Naumov
& Stepaniants (1972, fig. 9).

Symplectoscyphus plectilis (Hickson & Gravely, 1907)
Figure 11A—C

Sertularella plectilis Hickson & Gravely, 1907: 20, pl. 3 (fig. 21). Ritchie, 1913: 30, figs 8, 9, 11.
Broch, 1948: 11, fig. 2f-g.
Symplectoscyphus plectilis: Vervoort, 1972b: 354, fig. 8b—c.

Station
14/44-C.

Description

A single, small infertile colony of about 12 stems growing on Halecium sp.
Stem unfascicled, reaching a maximum height of 9 mm, slender, slightly geni-
culate, branching sparingly and in one plane. Branches arising from apophyses
immediately below hydrothecae at a wide angle (95-125°). Nodes usually indi-
cated by indentations of perisarc only. The two rows of hydrothecae in one
plane.

Hydrotheca very small, tubular, curved outwards, adnate for about one-
third adcauline length, narrowing to margin. Marginal teeth well developed.
No internal teeth.

Measurements (mm)

Internode, length R. Se ie ast ie oy 0,39-0,63
Hydrotheca, length abcauline .. ae 4 ie a 0,23-0,29
length adcauline, adnate part ie: oR a8 <i 0,09-0,14
length adcauline, free part 2 se a a 0,22-0,28
adnate part/total adcauline length a a a 0,26-0,36

diameter at margin 2 Me ey 8 es 0,08-0,12

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES 35

Remarks

The hydrothecae in this material are of a similar size to those described by
Ritchie (1913); they are a little longer than those described by most other
authorities.

S. plectilis is known from a number of localities in the Antarctic and
Subantarctic, the nearest to the present locality being Bouvet Island (Broch
1948). The distribution and synonymy are summarized by Vervoort (19725).

Symplectoscyphus subarticulatus (Coughtrey, 1875)
Figure 10OA—D

Thuiaria sub-articulata Coughtrey, 1875: 287, pl. 20 (figs 32-34).
Symplectoscyphus subarticulatus: Ralph, 1961: 801, 811, figs 14g-h, 15a—c (synonymy).
Blanco, 1968: 213, pl. 4 (figs 1-3). Leloup, 1974: 41, fig. 38.

Stations
2/6—B (fertile); 2/7—A; 14/45-C.

Description

Fascicled, pinnate stems reaching a maximum height of 105 mm and a
maximum diameter at the base of 1,5 mm. Hydrothecae completely obscured
by the peripheral tubes in the proximal part of the stem, with only the margins
visible in the central region, uncovered in the distal unfascicled region. In the
distal region nodes fairly clearly marked and sloping in alternate directions,
each internode bearing three hydrothecae and a hydrocladium arising imme-
diately below the third hydrotheca. The two rows of hydrothecae and hydro-
cladia in one plane.

Hydrocladium separated from stem apophysis by a distinct node, narrower
than stem. Nodes scarce, oblique, not well marked, sometimes not visible
for the entire length. Hydrothecae closely set, more so in the distal region where
the margin of one overlaps the base of the next, most hydrothecae with a
fenestra immediately below the base on one side.

Hydrotheca tubular, curved outwards, adnate for about two-thirds
adcauline height.

Gonothecae borne on hydrocladia below hydrothecae, long and carrot-
shaped, widest at distal end, strongly annulated with 15-17 circular crests (not
spiral), with terminal aperture on a raised collar.

Measurements (mm)

Hydrotheca, length abcauline .. a ve ba fe 0,40-0,47
length adcauline, adnate part - “ed a mS 0,38—0,60
length adcauline, free part on zi sh ye 0,23-0,31
adnate part/total adcauline length ue af ae 0,57—0,70
diameter at margin - i Le ry - 0,31-0,33

Gonotheca, length re wt as se iis “a 2,35-3,10

maximum diameter i a e7 ap isk 0,86—-1,24

36 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 10.
Symplectoscyphus subarticulatus (Coughtrey). A-B. Hydrothecae. C. Distal part of stem
showing origins of hydrocladia. D. Gonotheca.

Symplectoscyphus biformis (Jaderholm). E. A single hydrotheca. F. A group of three hydro-
thecae. Drawn from the holotype.

Scale in mm/10.

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES 37

Remarks

The hydrothecae of this material are larger than those of Ralph but smaller
than those of Blanco. The details of structure and the method of branching
agree exactly with those described by Ralph. The only hesitation in the identi-
fication lies in the gonotheca, which is about twice the length of that described
by Ralph and has more annulations (15-17 as against 5-8). However, the
shape of the gonotheca is exactly like that depicted by Coughtrey (he does not
give the size, but the diagram shows 10 annulations).

S. subarticulatus is best known from New Zealand, but recent reports
show that it also occurs in the southern Argentine (Blanco) and in Chile (Leloup).
It is a new record from the Kerguelen area.

Symplectoscyphus subdichotomus (Kirchenpauer, 1884)

Figure 11D-—F

Sertularella subdichotoma Kirchenpauer, 1884: 46, pl. 16 (fig. 1).
Symplectoscyphus subdichotomus: Blanco, 1969: 49, figs 1-18. Vervoort, 1972a: 140, figs 44b—d,
45 (synonymy). Leloup, 1974: 42, fig. 40.

Stations

Small form: 8/25—E; 26/64-E (fertile); 26/65—A (fertile); 28/71-C (fertile);
30/73-D; 31/74-G.
Large form: 3/11—D; 14/44-B; 21/57-D; 22/58-J; 26/63-F.

Description

Colonies flexuous and straggling, growing profusely with polyzoans and
Sertularella picta, and often intertwined to form a bushy mat. Stem unfascicled,
reaching 33 mm in height (small form) or 58 mm (large form), generally geni-
culate and giving off a branch at each elbow immediately below every third
hydrotheca, but many irregularities present. The two rows of hydrothecae and
branches in one plane. Branches of same diameter as stem, forming an angle
of 85—-108° with it, and often rebranching producing a subdichotomous effect.
Most of the nodes not clearly defined and recognizable only by an indentation
of the perisarc, but those at the origin of each branch, and on the stem imme-
diately above each axillary hydrotheca, more clearly defined and usually
forming a definite septum. Internodes always longer than the hydrothecae,
which do not overlap each other.

Hydrotheca adnate for over half adcauline height, curved outwards, widest
in centre and narrowing to margin, with three well-developed marginal teeth
and no internal teeth.

Gonotheca obovate (female), with 10-11 strong and crested circular
annulations (not spiral) and a long flaring mouth-funnel, containing two or
three large planulae.

38 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 11.

Symplectoscyphus plectilis (Hickson & Gravely). A. Part of stem showing origin of hydro-
cladium. B—C. Hydrothecae.

Symplectoscyphus subdichotomus (Kirchenpauer). D. Stem of small form showing origins
of hydrocladia and a gonotheca. E. Hydrotheca of small form. F. Hydrotheca of large form.

Scale in mm/10.

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES

Measurements (mm)

39

Small form Large form

Internode length .. 0,47-0,74 0,74-1,27
Hydrotheca, length abcauline 0,26-0,31 0,28-0,38
length adcauline, adnate part 0,25—0,30 0,25—0,39
length adcauline, free part 0,13-0,21 0,14-0,28
adnate part/total adcauline length 0,58—0,68 0,48-0,71
diameter at mouth 0,13-0,17 0,20—-0,30
Gonotheca, length 1,06—1,55
maximum diameter 0,58-0,84

Remarks

As far as the detailed measurements are concerned this material includes
two size-ranges—a small and a large—the smaller fitting best with the dimensions
given by Vervoort, Blanco and earlier workers. The growth-form and structure
of the two are the same, but there are no gonothecae to confirm the identifica-
tion of the larger form. |

The synonymy of S. swbdichotomus has been reviewed by Vervoort (1972a),
who also gave some excellent diagrams. There is no doubt at all that the present
material (at any rate the smaller form) belongs to the same species. Because
of the presence of a number of closely allied species of dubious validity, the
geographical range is uncertain—the species is certainly common in the Falk-
lands/South American region, but has apparently not been reported from
Kerguelen.

Family Plumulariidae

Kirchenpaueria triangulata (Totton, 1930)
Plumularia triangulata Totton, 1930: 225, fig. 61.
Kirchenpaueria triangulata: Millard, 1975: 375, fig. 119E—-H.
Station
17/50-B.

Description

A rich, fertile colony epizootic on Plumularia insignis. Stems reaching
18 mm in height, unfascicled, with nodes absent or very indistinct, otherwise
details as in previous descriptions. Gonophores female, reaching 2 mm in
length, triangular in section and containing a single layer of eggs over a central
spadix.

Remarks and distribution

This is a new record for the Kerguelen area. The species was previously
known from South Africa (in depths greater than 100 m) and from New Zealand.

40 ANNALS OF THE SOUTH AFRICAN MUSEUM

Oswaldella bifurca (Hartlaub, 1904)
Figure 12A—C

Schizotricha bifurca Hartlaub, 1904: 16, pl. 3 (figs 4-8).
Oswaldella bifurca: Totton, 1930: 208, fig. 50. Naumov & Stepaniants, 1962: 98.

Station
26/64-J.

Description

Several infertile stems reaching a maximum height of 22 mm. Stem divided
by transverse nodes into internodes which usually bear one hydrocladium
each, but rarely two. Cauline nematothecae very difficult to distinguish and
sometimes missing; at the maximum development there is one mamelon on
the hydrocladial apophysis and one nematotheca in the axil, with another
nematotheca (or a naked sarcophore) on the main stem above the axil.

Hydrocladia unbranched near the proximal and distal ends of colony,
branched in the central region. Branches arising from below hydrothecae and
to one side, as many as four from one hydrocladium, and these may be all on
the same side, on alternate sides or in opposite pairs. One secondary hydro-
cladium has a tertiary branch.

The two rows of primary hydrocladia not in the same plane but displaced —
towards anterior surface of stem. Secondary hydrocladia similarly displaced,
so that the whole arrangement is in at least six different planes and presents a
stiff and spiky appearance. Since the hydrothecae are borne on the anterior
surface of the hydrocladia, those of primary and secondary hydrocladia do not
face one another.

Hydrocladia generally heteromerous with alternating athecate and thecate
internodes, but many variations; the athecate internodes may be absent or
duplicated, long or short, and may or may not bear a median nematotheca.
Thecate internodes usually with one median inferior and one median superior
nematotheca, but the nematotheca is often missing leaving only a naked sarco-
phore, and this is especially common in the first thecate internode of the hydro-
cladium. Nematothecae, when present, curved and scoop-shaped. Median
superior nematotheca situated behind distal part of adcauline thecal wall
which is not completely adnate.

Hydrotheca wider than deep, with more or less straight abcauline wall
and margin at least slightly oblique.

Measurements (mm)

Thecate internode, length an i ds oe a 0,42-0,96
Hydrotheca, depth abcauline .. ¥ - ng ad 0,12-0,19
diameter at margin oe oe es 7 a 0,17-0,22

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES 4]

F

A-C
Dh le

Fig. 12.
Oswaldella bifurca (Hartlaub). A-B. Normal parts of hydrocladia. C. Proximal end of
hydrocladium showing origin from stem and first thecate internode which is without a

nematotheca.
Plumularia insignis Allman. D. Part of stem in thinner region showing origins of two branches

and their preceding hydrocladia. E. Gonothecae. F. Part of hydrocladium.

Scale in mm/10.

42 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks

This material is assigned to O. bifurca with some hesitance, the decision
being based mainly on the shape of the hydrotheca, which is wider than deep,
and on the method of branching. As regards the latter feature Totton’s words
(1930) are pertinent, when he says for O. bifurca ‘Hydrothecae of hydroclade
and branch not strictly opposed, but facing slightly upwards’. On the other
hand the features used by Totton as diagnostic for O. bifurca, namely ‘once-
branched hydrocladia’ and the absence of an inferior nematotheca on the first
hydrocladial internode, are found to be variable and of dubious value. The
presence of athecate internodes in the hydrocladium and the number of hydro-
cladia to a stem internode are also variable characters. This species is known
from various localities in the Antarctic; it has not been reported before from
Kerguelen.

Plumularia insignis Allman, 1883
Figure 12D-F

Plumularia flabellum* Allman, 1883: 19, pl. 1 (figs 1-4).

Plumularia insignis Allman, 1883: 21, pl. 2. Billard, 1910: 32, fig. 14.
Plumularia abietina Allman, 1883: 21, pl. 3.

Plumularia insignis v. flabellum: Billard, 1910: 34, fig. 15.
Plumularia insignis v. abietina: Billard, 1910: 35.

Plumularia sp. Naumov & Stepaniants, 1962: 99, fig. 19.

Stations
3/10—-A; 3/11-—C; 7/22—A; 17/50—A; 18/52—A; 23/59-D; 24/61-A; 26/64-A;
30/73-E.

Description

Many magnificent colonies, the tallest reaching 940 mm. Rootstock a
mass of interwoven fibres suitable for anchoring in mud. Stem long, flexuous
or fairly stiff, fascicled for the greater part, giving off branches in all planes,
unsegmented. Immediately below each branch a single hydrocladium arises
from the stem and from the same component tube, otherwise cauline hydro-
cladia occur only rarely. Component tubes of stem bearing longitudinal rows
of nematothecae.

Branches fascicled or unfascicled, pinnate, divided by transverse nodes at
irregular intervals into internodes bearing one to four alternate hydrocladia.
Hydrocladial apophysis with two axillary nematothecae (one anterior and one
posterior) with a mamelon between them, and one (or rarely two) nematothecae
on distal end. Nematothecae on main axis rather irregular, but at least one
between every two successive hydrocladia and several below the first
hydrocladium.

* Although P. flabellum has page priority over P. insignis, from Article 24(a) of the Code of

Zoological Nomenclature, Billard (1908: 759) counts as the ‘first reviser’ and he has clearly
chosen P. insignis as the name of the composite species.

een aise

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES 43

In some colonies the hydrocladia bear practically only thecate internodes,
each with one hydrotheca and three nematothecae (one median inferior and
one pair of laterals), athecate internodes occurring only sporadically and
obviously resulting from regeneration after injury. In other colonies intermediate
athecate internodes are present throughout, each with one median nematotheca
and two internodal septa. Thecate internodes very rarely have two median
inferior nematothecae. Internodal septa usually not so well developed as in
the type material: 1-2 below hydrotheca, 0-3 behind it, and 0-1 above it.

Gonothecae arising from branches next to hydrocladia, elongated, widening
distally, with a terminal oblique opening, as illustrated by Allman (1883) for
P. abietina.

Measurements (mm)

Thecate internode, length v = Ne < ae: 0,54-0,80
Hydrotheca, height abcauline .. ot: . He Pa 0,21-0,34
diameter at margin A 2 on =? fa 0,14-0,21
Gonotheca, length 43 ie a a ne A 1,40-1,95
maximum diameter ie ae = Ae ue 0,35-0,58
Remarks

This species occurs in several varieties, of which three (described by Allman
as P. insignis, P. flabellum and P. abietina) are known from the Subantarctic.
In the general appearance of the colony and in the detailed measurements,
most of the present material closely resembles the nominal variety illustrated
by Allman (1883: pl. 2). One colony, however (26/64—A), is of stouter build
and more irregular branching and is more like Allman’s diagram of P. flabellum
(1883: pl. 1). Of the three, Allman showed hydrocladia on the main stem only
in P. flabellum, Nowhere has the singular character of one hydrocladium below
each branch been described, and if this character is indeed peculiar to the
Kerguelen material it would be worth naming a separate variety.

Most of this material is unusual in possessing as the ‘normal’ condition
only thecate internodes in the hydrocladia, athecate internodes being rare and
sporadic. One colony alone (26/64-A, mentioned also above) has regular
intermediate athecate internodes. Billard (1910) has commented on the irregu-
larity of occurrence of athecate internodes which are often the result of regenera-
tion after injury. Naumov & Stepaniants (1962) described some infertile material
from Kerguelen and Heard Islands without athecate internodes as Plumularia sp.
This is without doubt the same species.

The species is known from Prince Edward Island, Marion Island, Kerguelen
and Heard Islands, and there are two varieties from the East Indies.

Schizotricha unifurcata Allman, 1883

Schizotricha unifurcata Allman, 1883: 28, pl. 7 (figs 1-3). Stechow, 1925: 498. Totton, 1930:
231, fig. 65, pl. 3 (fig. 4). Naumov & Stepaniants, 1972: 54. Blanco & De Miralles, 1972:
21, pl. 5 (figs 3440).

Polyplumaria unifurcata: Billard, 1910: 41, fig. 18.

44 ANNALS OF THE SOUTH AFRICAN MUSEUM

Stations
8/25-A; 9/26-B (fragments); 10/30-B; 22/58-A; 23/59-C.

Description

Stems with tangled rootstock for mud-penetration and reaching 175 mm
in height. Hydrocladia branching one to three times, 10-18 mm in length.
Details of hydrocladia as in type material.

Male gonothecae present on first colony, arising from thecate internodes
below hydrothecae, pear-shaped, with oblique distal aperture, bearing two to
four nematothecae on basal region.

Remarks

Stechow (1925) combined the species S. unifurcata Allman, 1883; S. tur-
queti Billard, 1906; S. anderssoni Jaderholm, 1904; and S. glacialis (Hickson
& Gravely, 1907) under one name. Totton (1930) added S. multifurcata Allman,
1883. All of these species are Antarctic or Subantarctic in distribution.

This material is very similar to the type material and indeed, two of the
colonies come from the type locality (Kerguelen Island).

DISCUSSION

Of the 33 species here recorded (omitting Campanularia sp.) 5 are cos-
mopolitan, namely Modeeria rotunda, Halecium delicatulum, Halecium tenellum, -
Filellum serratum and Lafoea dumosa (Table 2).

The remaining 28 species show only a very slight affinity with the nearest
continental masses, for only 1 (3,6%) also occurs in South Africa (Kirchen-
paueria triangulata) and only 8 (28,6°%) also occur in the Australasian region.

The affinities of these 28 species are in fact mainly with the South American
region (or Magellan Province, including the Falklands and South Georgia)
and with the Antarctic Continent, for 19 of the species (67,9%) also occur in
the former and 14 (50,0°%) in the latter.

Surprisingly, there is little obvious affinity with Bouvet Island (3 species:
10,7°%) or Marion and Prince Edward Islands (5 species: 17,9%) which lie
at much the same latitude, but this is probably merely a reflection of the inade-
quate knowledge of these islands.

This analysis thus supports the classification of the Kerguelen and Crozet
groups of islands as Subantarctic (J. C. Briggs 1974), with a fauna intermediate
between that of the Cold Temperate South American Region and the Antarctic
Region. As has been remarked by Naumov & Stepaniants (1962), it is not
possible to draw a sharp line of demarcation between the fauna of the Ant-
arctic and that of the Subantarctic.

There is at this stage a little evidence of a divergence between the fauna
of the Kerguelen group on the one hand and the Crozet group on the other.
Although they share 12 (42,9°%%) of the non-cosmopolitan species, 8 occur only
in Kerguelen and 8 only in Crozet.

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES 45

TABLE 2
The distribution of the 33 species recorded in this paper.

z
—
s 3 E
bie Sige Tee = <
S -h 8: « oe
ee a ay ie ee eS
aos Peete gs ee
Gig Wa Ei ee wae gs Se
o 5 w = oS oS = = 5
O wo A a = Se Co 2
Eudendrium rameum x x a
E. tottoni x x x
Modeeria rotunda x x x x (cosmop.)
Opercularella belgicae x x SA
Phialella chilensis * ys sé
Halecium delicatulum xX x x x x x x x (cosmop.)
H. dufresneae WA
H.jaederholmi . x x x
H. tenellum : x x oe * x (cosmop.)
Hydrodendron arborea xX NA x Se Me
Filellum serratum x x x x x (cosmop.)
Grammaria abietina. xX mK x
Halisiphonia ?nana x x
Hebella striata . % x x
Lafoea dumosa . : x ~ 4 x K x (cosmop.)
Zygophylax crozetensis X
Campanularia
norvegiae : ; Xx Xx
Silicularia rosea x Me Sc x <
Tulpa diverticulata x rm x
Staurotheca antarctica x x < we
S. dichotoma Done x x x x
Sertularella geodiae . x S x
S. picta . x * x x
Symplectocyphus
curvatus . s< x
S. elongatus . xX x x o<
S. mawsoni < x
S. plectilis . x * x x
S. subarticulatus x SS x
S. subdichotomus . xX x x x x
Kirchenpaueria
triangulata . ; x x x
Oswaldella bifurca . x x
Plumulariainsignis . x x x
Schizotricha
mayurcata” tt S< Se oe x

Total (less cosmop.) 20 19 19 3 5 1 14 1 8
map icss.cosmop.) 71:4 “67,9 67,9 - 10,7 17,9 ° 3,6 50:0 3,6 28,6

46 ANNALS OF THE SOUTH AFRICAN MUSEUM

ACKNOWLEDGEMENTS

The author wishes to thank Dr J. C. Hureau of the Muséum National
d’Histoire Naturelle, Paris, and Dr P. M. Arnaud of the Station Marine
d’Endoume, Marseille, for the privilege of examining and reporting on this
collection.

The British Museum (Natural History) granted permission for the exami-
nation of Challenger material of Sertularia (Symplectoscyphus) exserta and
Terra Nova material of ‘Sertularella biformis’, while the Naturhistoriska Riks-
museum, Stockholm, permitted examination of type material of Sertularella
biformis from the Swedish Southpolar Expedition. This help is gratefully
acknowledged.

REFERENCES

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ALLMAN, G. J. 1888. Report on the Hydroida dredged by H.M.S. Challenger during the years
1873-76. Part II. The Tubularinae, Corymorphinae, Campanularinae, Sertularinae and
Thalamophora. Rep. Voy. Challenger 1873-76, Zoology 23 (70): 1-90. -

BILLARD, A. 1908. Sur les Plumulariidae de la collection du ‘Challenger’. C.r. hebd. Séanc.
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BILLARD, A. 1910. Revision d’une partie de la collection des Hydroides du British Museum.
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BILLARD, A. 1914. Hydroides. Deux. Expéd. Antarct. franc. 1914: 1-34.

BILLARD, A. 1922. Note critique sur quatre espéces de Sertularella. Revue suisse Zool. 30:
103-114.

BLANco, O. M. 1968. Neuva contribucion al conocimiento de la fauna marina Hidroide.
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BLANco, O. M. 1969. Simplectoscyphus subdichotomus (Kpr.), con especial referencia a la
morfologia gonangial (Celent. Hydrozoa). Neotropica 15: 49-53.

BLANCO, O. M. & DE MIRALLEs, D. A. B. 1972. Hidrozoos de la Isla Pedro I. Contrnes Inst.
Antart. argent. 145: 3-43.

Briccs, E. A. 1939. Hydroida. Sci. Rep. Australasian Antarct. Exped. (C) 9 (4): 1-46.

BriacoGs, J. C. 1974. Marine zoogeography. New York et al.: McGraw-Hill.

Brocu, H. 1948. Antarctic hydroids. Scient. Results Norw. Antarct. Exped. 28: 1-23.

CORNELIUS, P. F. S. 1975. A revision of the species of Lafoeidae and Haleciidae (Coelenterata:
Hydroida) recorded from Britain and nearby seas. Bull. Br. Mus. nat. Hist. (Zool.) 28:
375-426.

COUGHTREY, M. 1875. Notes on the New Zealand Hydroideae. Trans. Proc. N.Z. Inst. 7:
281-293.

Epwarps, C. 1973. The medusa Modeeria rotunda and its hydroid Stegopoma fastigiatum,
with a review of Stegopoma and Stegolaria. J. mar. biol. Ass. U.K. 53: 573-600.

FRASER, C. McL. 1944. Hydroids of the Atlantic coast of North America. Toronto: Univ.
Toronto Press.

HARTLAvB, C. 1904. Hydroiden. Résu/t. Voyage S.Y. Belgica: 1-19.

HarTLaAvus, C. 1905. Die Hydroiden der magalhaensischen Region und chilenischen Kite.
Zool. Jb. Suppl. 6 (3): 497-714.

Hickson, S. J. & GRAVELY, F. H. 1907. Coelenterata. I]. Hydroid zoophytes. Nat. Antarct.
Exped. nat. Hist. 3: 1-34.

HureEau, J. C. 1976. La campagne d’océanographie biologique MD 03/ICHTYO a bord du
‘Marion-Dufresne’ (18 Mars—28 Avril 1974). Résultats préliminaires et liste des stations.
CNFRA 39: 3-25.

JADERHOLM, E. 1905. Hydroiden aus antarktischen und subantarktischen Meeren gesammelt
von der schwedischen Siidpolarexpedition. Wiss. Ergebn. schwed. Siidpolarexped. 5: 1-41.

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES 47

JADERHOLM, E. 1917. Hydroids from the south seas. Redog. Norrkdépings Ldéroverk Lésaret
1916-1917: 1-23.

KIRCHENPAUER, G. H. 1884. Nordische Gattungen und Arten von Sertulariden. Adbh. Geb.
Naturw. Hamburg 8 (3): 1-54.

LeLoup, E. 1974. Hydropolypes calyptoblastiques du Chili. Report no. 48 of the Lund Uni-
versity Chile Expedition 1948-1949. Sarsia 55: 1-61.

MILLarp, N. A. H. 1968. South African hydroids from Dr Th. Mortensen’s Java—South Africa
Expedition, 1929-1930. Vidensk. Meddr dansk naturh. Foren. 131: 251-288.

MILLARD, N. A. H. 1971. Hydrozoa. In: ZINDEREN BAKKER, E. M. VAN, WINTERBOTTOM, J. M. &
Dyer, R. A., eds. Marion and Prince Edward Islands: 396-408. Cape Town: A. A. Balkema.

MILLARD, N. A. H. 1975. Monograph on the Hydroida of southern Africa. Ann. S. Afr. Mus.
68: 1-513.

Naumov, D. V. 1960. Hydroids and Hydromedusae of the marine, brackish and freshwater
basins of the U.S.S.R. Opred. Faune SSSR 70: 1-585. (In Russian; translated by Israel
Program for Scientific Translations, 1969.)

Naumov, D. V. & STEPANIANTS, S. D. 1962. Hydroidea (Thecophora) collected by the Soviet
Antarctic Expedition on the M/V ‘Ob’, in antarctic and subantarctic waters. Biol. Rep
Soviet Antarct. Exped. (1955-1958), I. Studies of marine fauna 1 (9): 69-104. (In Russian;
translated by Israel Program for Scientific Translations, 1966.)

Naumov, D. V. & STEPANIANTS, S. D. 1972. Marine invertebrates from Adelie Land, collected
by the XIIth and XVth French Antarctic Expedition. 3. Hydroida. Tethys Suppl. 4: 25-60.

NuttTIna, C. C. 1905. Hydroids of the Hawaiian islands collected by the steamer ‘Albatross’
in 1902. Bull. U.S. Fish Comm. 23: 931-959.

RAtpH, P. M. 1956. Variation in Obelia geniculata (Linnaeus, 1758) and Silicularia bilabiata
(Coughtrey, 1875) (Hydroida, F. Campanulariidae). Trans. R. Soc. N.Z. 84: 279-296.

RAtpH, P. M. 1957. New Zealand thecate hydroids. Part I. Campanulariidae and Campanu-
linidae. Trans. R. Soc. N.Z. 84: 811-854.

RALPH, P. M. 1958. New Zealand thecate hydroids. Part II. Families Lafoeidae, Lineolariidae,
Haleciidae and Syntheciidae. Trans. R. Soc. N.Z. 85: 301-356.

RA.pH, P. M. 1961. New Zealand thecate hydroids. Part III. Family Sertulariidae. Trans. R.
Soc. N.Z. 88: 749-838.

Rees, W. J. & THURSFIELD, S. 1965. The hydroid collections of James Ritchie. Proc. R. Soc.
Edinb. (B) 69: 34-220.

Ritcuig, J. 1913. The hydroid zoophytes collected by the British Antarctic Expedition of
Sir Ernest Shackleton, 1908. Proc. R.Soc. Edinb. (B) 33: 9-34.

STECHOW, E. 1923. Zur Kenntnis der Hydroidenfauna des Mittelmeeres, Amerikas und anderer
Gebiete. II. Teil. Zool. Jb. (System. Abt.) 47: 29-270.

STECHOW, E. 1925. Hydroiden der deutschen Tiefsee-Expedition. Wiss. Ergebn. dt. Tiefsee-
Exped. ‘Valdivia’ 17: 383-546.

ToTTon, A. K. 1930. Coelenterata. Part V. Hydroida. Nat. Hist. Rep. Br. Antarct. Terra Nova
Exped. 5: 131-252.

TREBILCOCK, R. E. 1928. Notes on New Zealand Hydroida. Proc. R. Soc. Vict. 41: 1-31.

VANHOFFEN, E. 1910. Die Hydroiden der deutschen Siidpolar-Expedition 1901-1903. Dr.
Stidpol.- Exped. 11: 269-340.

VERVOORT, W. 1946. Hydrozoa (Cl). .A Hydropolypen. Fauna Ned. 14: 1-336.

VERVOORT, W. 1972a. Hydroids from the ‘Theta’, ‘Vema’ and ‘Yelcho’ cruises of the Lamont-—
Doherty Geological Observatory. Zool. Verh., Leiden 120: 1-247.

VERVOORT, W. 19726. Hydroids from submarine cliffs near Arthur Harbour, Palmer Archi-
pelago, Antarctica. Zodl. Meded. 47: 337-357.

WEILL, R. 1934. Contribution a l’étude des Cnidaires et de leurs nématocystes. I. Recherches
sur les nématocystes. II. Valeur taxonomique du cnidome. Trav. Stn zool. Wimereux 10:
1-347; 11: 349-701.

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Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (figs 8a—b).

Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:
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N. A. H. MILLARD

HYDROIDS FROM THE KERGUELEN J

AND CROZET SHELVES, COLLECTED BY
THE CRUISE MD.03 OF THE MARION-DUFRESNE

JME 73 PART 2 JUNE 1977 ISSN 0303-2515

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| OF THE SOUTH AFRICAN
MUSEUM

ZAPE TOWN

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

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

FISCHER, P.-H., DuvaL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. 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.

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

THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270.

(continued inside back cover)

i wat: si a

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

Volume 73 Band
June 1977 Junie
Part 2 Deel

RELATIONSHIPS OF THE SOUTH AFRICAN
FOSSIL FROG
EOXENOPOIDES REUNINGI (ANURA, PIPIDAE)

By

RICHARD ESTES

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material
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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

RELATIONSHIPS OF THE SOUTH AFRICAN FOSSIL FROG
EOXENOPOIDES REUNINGI (ANURA, PIPIDAE)

By
RICHARD ESTES

Department of Zoology, San Diego State University,
San Diego, California

(With 11 figures)

LMS. accepted 29 December 1976]

ABSTRACT

The relationships of the South African fossil frog, Eoxenopoides reuningi, originally
described as a pipid, have been questioned and several authors have placed it in a separate
family Eoxenopoididae. Restudy of the original material for the first time since its original
description shows this frog to be, in fact, a pipid as originally described and to have a remarkable
mixture of primitive and derived character states. The anterior position of parietal foramen,
scapula length, unfused carpus and expanded transverse processes of posterior vertebrae
indicate E. reuningi as more primitive than any living pipid. Highly specialized in lacking teeth,
and particularly in reduction and fusion in the vertebral column that results in possession of
only six presacral vertebrae, it has nevertheless only one derived character state in common
with the extant specialized Hymenochirus and South American pipids. Its greatest number of
derived character states are in common with species of the living pipid Xenopus, but there is
no indication of greater proximity to any one of them and E. reuningi appears to have diverged
from the ancestor of the living genus some time prior to the late Cretaceous.

Review of available radiometric and palaeontological evidence suggests a Palaeogene age
(perhaps late Eocene or Oligocene) for Eoxenopoides reuningi rather than the original suggestion
of late Cretaceous or early Tertiary.

CONTENTS
PAGE
Introduction . : : ‘ é : : : . fl ee
Geological age and provenance of Eoxenopoides reuningi . 51
Materials : : : é : : 5 : : : ; SS
Redescription of materials. f : : : ; : a OOS
Comparisons . : ‘ ; ‘ : : ; , : oi 469
Relationships of Eoxenopoides reuningi ; : ‘ : sph ly aac
Taphonomy and palaeoecology . : : : : : ae.
Acknowlegments . i : : : : : : : Si PES
References. : : ; : : ‘ P E : : Neola
Abbreviations . : ; : : : : : : : a OU
INTRODUCTION

Large collections of fossil frogs are rare. One such collection, made in the
early 1930s first by E. Reuning and later by L. Boonstra in Namaqualand, north-
western South Africa, has received little attention since its original description
(Haughton 1931). Over the last several years, the author has been concerned with

49

Ann. S. Afr. Mus. 73 (2), 1977: 49-80, 11 figs.

50 ANNALS OF THE SOUTH AFRICAN MUSEUM

the fossil record of pipid frogs, and need for revaluation of this important
collection was seen through comparisons made during study of fossil Xenopus
from Brazil (Estes 1975a, 19755).

Haughton referred all the frogs in the collection to a single genus and species
Eoxenopoides reuningi, and he prepared a careful description of his material.
Most of the fossil pipid frogs (¢ndeed most fossil frogs in general) have been
described since that time, and Haughton was thus unable to evaluate his material
as thoroughly as it is possible to do now. The most recent summary of the frog
fossil record is given by Estes & Reig (1973), and a further evaluation of fossil
pipids may be found in Estes (1975a); only such literature and discussion as is
pertinent to the revaluation of Eoxenopoides will be repeated here.

Although Haughton’s description was carefully done, his drawings were
somewhat diagrammatic and the nature of the material led to some misinterpreta-
tion of bones. It is no surprise, therefore, that the very little written about the
genus subsequent to Haughton’s account is incorrect for the most part and has
confused rather than clarified the status of the animal; so far as the author is
aware no one of these subsequent papers has been based on personal observation
of the fossil material.

Kuhn (1941) cited it as belonging to a ‘fam. opisthocoelorum nov.’, and as
‘recht aberrante Opisthocoelia . . .”. He did not at that time name a new family,
but suggested relationships with his middle Eocene genus Opisthocoelellus from
Germany. Reig (1958), in discussing a proposed new classification of frogs, noted -
that Eoxenopoides ‘pertenece a una familia seguramente nueva, que también debe
relacionarse estrechamente con la de los pipidos’, but gave no explanation as to
why a new family was suggested. Shortly after this, Casamiquela (1960, 1961)
described the frog Shelania pascuali from the Eocene of Patagonia, giving pre-
sumed resemblances between Eoxenopoides and Shelania; he used these resem-
blances in 1961 to define a new family Eoxenopoididae to include the two genera,
and suggested a more primitive but close relationship to Pipidae. He was
unaware, apparently, that the family name Eoxenopoididae had already been
used the previous year by Parodi Bustos et al. (1960), who referred late Cretaceous
frogs from northern Argentina to a new species Eoxenopoides? saltensis. The
name Sal/tenia ibanezi had already been applied to this taxon by Reig (1959),
based on fossils from the same locality; Reig referred S. ibanezi to the Aglossa.
Parodi Bustos et a/. seemed not to have been aware of Reig’s paper initially, but
after receiving it Parodi Bustos & Kraglievich (1960) defended with additional
force their generic identification, stating that ‘el material . . . no puede separarse
genericamente de Eoxenopoides’. Later, Parodi Bustos (1962) agreed with
Casamiquela (1961) that Shelania pascuali, Eoxenopoides? saltensis and E.
reuningi belonged to the same family. Meanwhile, however, Kuhn (1961) had
also named a new family Eoxenopoididae, including only Eoxenopoides reuningi.
He apparently had not seen the papers of Parodi Bustos but suggested that
Saltenia ibanezi Reig belonged to yet another new family, which he did not name.
The family Eoxenopoididae was thus named independently three times, by Parodi

RELATIONSHIPS OF THE SOUTH AFRICAN FOSSIL FROG 3!

Bustos et al. (1960), by Casamiquela (1961) and by Kuhn (1961). Hecht (1963)
mentioned Eoxenopoides as constituting evidence for wide distribution of the
Aglossa during the Cretaceous (in combination with other aglossan fossils) but
made no taxonomic judgements.

Nevo (1968) was the first to attempt clarification of this issue. He correctly
pointed out that the character states used by Casamiquela (1961) to set up the
family Eoxenopoididae were ‘inappropriate as taxonomic criteria at the family
level’. He returned Eoxenopoides reuningi to the Pipidae, following the original
decision made by Haughton in 1931; this allocation is supported and documented
in the present paper. Current work on Shelania cited by Estes (1975a) has shown
it to be congeneric with Xenopus and hence a pipid as well (Baez & Estes in
preparation). Baez (in ms. as noted by Estes & Reig 1973, and Estes 1975a) has
confirmed that Eoxenopoides? saltensis Parodi Bustos et al. is a junior synonym
of Saltenia ibanezi Reig, and that the taxon is unquestionably referable to the
Pipidae. As the present study and that of Baez demonstrates, Saltenia and
Eoxenopoides are not closely related, although both are pipids.

It will be attempted to show in this paper that Eoxenopoides reuningi is a
pipid closely related to Xenopus, perhaps closest to the relatively generalized
species X. muelleri, but specialized in a number of features, notably the short
vertebral column (six presacrals) and in lack of teeth on the maxilla and
premaxilla.

GEOLOGICAL AGE AND PROVENANCE OF EOXENOPOIDES REUNINGI

The specimens of Eoxenopoides reuningi were taken from sediments filling
one of a series of volcanic pipes on a farm near Banke (or Banker) in the Gamoep
area near Platbakkies, Namaqualand, Republic of South Africa (30°22’S 18°26’E,
3018 AD Platbakkies sheet, first edition, Geological Map of South Africa,
1: 50 000). The pipe in question is probably one of the group immediately south
of the Kimberlite group marked on the 1970 edition of the Geological Map of
the Republic of South Africa, Kingdoms of Lesotho and Swaziland, | : 1 000 000.

The specimens occur both in black carbonaceous and greenish clays
deposited in a temporary lake that developed in the eroded basin or crater of
a volcanic pipe about 280-325 metres in diameter, following cessation of the
volcanism. The volcanic pipe cuts crystalline metamorphic rocks of Precambrian
age. At least 33 metres of clays were deposited in this particular trap, and fossil
frogs occur throughout this thickness at the type locality although most of the
material came from dumps around the edge of the pipe and was consequently
from the upper part of the section. Above the black and green clays a white
clayey deposit occurs, and in a near-by excavation a white clayey sandstone
contains fragments of fragile green mudstone like that in which the frogs occur.
The sandstones are derived from weathering of local basement rocks (gneisses)
and although they cannot be traced laterally, Haughton (1931) noted their
similarity to the Kangnas sands near by, below which a specimen of the iguano-

52 ANNALS OF THE SOUTH AFRICAN MUSEUM

dont dinosaur Kangnasaurus coetzeei was found; he therefore suggested that the
clays containing the frogs might be “contemporaneous with or slightly later’ than
the dinosaur-bearing sediment and thus of late Cretaceous or early Cenozoic age.
The dinosaur remains are not reworked, but Anthony Tankard (1976 in /itt.) has
confirmed that there is no direct evidence for this correlation; it should therefore
be discounted in an age determination for Eoxenopoides reuningi.

Recent radiometric dates on an olivine melilitite pipe on the farm Dikdoorn,
south of Garies, gave an age of 38,5 m.y. (Kroner 1973) or about late Eocene age.
Andrew Moore (1976 in /itt.) notes that as yet there has been no date on the
similar melilitite cluster of pipes in the Gamoep area although they are assumed
to be of the same age. The Banke pipe, capped by sediments that yielded the flora
and fossil frogs, however, seems to have a distinctly different chemical imprint
from the olivine melilitite suite and thus may be of different age from that given
above. Moore further notes that a late Cretaceous to early Cenozoic age for the
pipe at Banke, as suggested by Haughton, would indicate that the pipe is of
similar age to a phase of volcanic activity that produced pipes on the southern
margin of the continent dated at 58 m.y. (Palaeocene) (Dingle & Gentle 1972).
Unfortunately, both Haughton’s and Moore’s suggestions are circumstantia], and
lacking a radiometric date on the Banke pipe its age and that of its overlying
sediments must on geological grounds still be regarded as speculative.

Along with the publication of Haughton’s paper, three papers on associated
plant material appeared (Rennie 1931; Adamson 1931; Kirchheimer 1934); none
of them gave any specific age determination for the sediments in which both frogs
and plants were found. More recent knowledge indicates, however, that a some-
what more specific determination is possible. The pollen described by Kirch-
heimer has been mentioned by Cranwell (1961), who noted that similar pollen
types have been found in ‘rather younger’ (Oligocene) deposits of Kerguelen
Archipelago (Cookson 1947). The leaf flora offers more specific limits. D. I. Axel-
rod (1976 in Jitt.) believes that the size relationships of the leaves and their sclero-
phyllous nature indicate a relatively dry climate, with rain limited to the warm
season. Based on comparisons with other known fossil floras, the leaf flora is
unlikely to be older than later Eocene and not much younger than Miocene,
according to Axelrod’s determinations.

If the rather circumstantial evidence from the floral remains is accepted, the
age range for Eoxenopoides reuningi would be late Eocene through Oligocene, a
determination that is in accord with the radiometric dates of 38,5 m.y. on the
Garies pipes, as well as with the rather tenuous correlation with the Kerguelen
sequence suggested by Cranwell (1961). As these suggestions are all compatible
this range is taken as the most acceptable one at present, one based on both
geological and fossil data. If Moore’s suggestion that difference in chemical
imprint between the Banke and olivine melilitite pipes indicates an earlier age
can be coniirmed by a radiometric date, this estimate of age must be revised
downward; a Palaeogene age, however, seems to be compatible with both
suggestions.

RELATIONSHIPS OF THE SOUTH AFRICAN FOSSIL FROG 53

MATERIALS

The collection has not been increased beyond the materials available to
Haughton (1931), and includes the original series collected by E. Reuning as well
as the later systematic collection of L. D. Boonstra that was correlated more or
less with depth of excavation. Haughton gave his series the informal numbers
1-44; of these only 3-7, 9-14, 16-18, 20-21, 23, 35, 40 and 43 still retain these
numbers and could be checked against Haughton’s description. Subsequently the
best specimens have been given South African Museum (SAM) numbers 9938-
9965, K4596-K4628, and K4633-K4636; each number may include the remains
of more than one individual. There are in addition perhaps 50 fragmentary
specimens that are not well enough preserved to be informative. In the discussion
below, the SAM numbers are used; Haughton’s informal numbers are appended
in cases in which reference to his description is made.

Some of Haughton’s specimens have been covered with a layer of shellac
that has hardened and darkened over the years to an extent that confirmation of
his findings on individual specimens is now often difficult or impossible. There is,
fortunately, abundant material not so treated and in which preservation is excel-
lent; thus few significant aspects of the osteology of Eoxenopoides remain in
question.

REDESCRIPTION OF MATERIALS
Holotype

SAM-9938 in the South African Museum, Cape Town. Complete adult
frog, Banke, near Platbakkies, Namaqualand, Republic of South Africa.
Collected by E. Reuning.

The general proportions of Eoxenopoides are shown in Figure 1; it is a
relatively compact-bodied frog with relatively large hind feet. Skull proportions
of the most common size group appear in the restorations of Figure 2; age
changes in skull proportions are shown in Figure 3.

Premaxilla

Haughton’s description of these bones in SAM—9948 (H6) was actually
based on the nasals (see below). The premaxillae are poorly preserved on all
specimens but are best seen on SAM-—9941. Teeth are absent (e.g. SAM—K4635).
The bones are paired, their median suture overlapped ventrally by the anterior
end of the parasphenoid. Their lateral expansions complete the arcade of the
upper jaw by fitting into excavations on the anteromedial sides of the maxillae.
The prominent alary (nasal) processes are separated on the midline by the
anterior ends of the nasals, and appear to be somewhat expanded dorsally
(Fig. 2).

Maxilla

The maxilla is well preserved on many specimens, e.g. SAM—9940, 9941 and
K4633 (Fig. 2). It is a very long bone, extending far anteriorly in front of the

54 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 1. Eoxenopoides reuningi. Restoration of skeleton in dorsal view. Cleithrum and lower
jaw removed on right. Dashed line indicates extension of elements of shoulder girdle ventral
to skull and vertebral column.

premaxillae to the alary processes of the latter, and posteriorly reaching or
exceeding the posterior border of the palatal vacuity. It is flattened dorsally and
expands anteriorly into a spoon-shaped process, the cavity of which opens
posteriorly into a large circular foramen extending into the body of the bone
(Fig. 4A—B). Its ventral edge is toothless but forms a sharp ridge (pars dentalis)
that marks the oral margin. Its posterior end is pointed.

Lachrymal

No lachrymal is present. The element so identified by Haughton is a part
of the ossified planum antorbitale (see below, sphenethmoid).

Nasal

The nasals are best shown on SAM—-K4596a, K4605a, K4615 and K4616.
They are semicircular paired bones lying on the borders of the nares and partly
overlapping the frontoparietal (Fig. 2A). Laterally they are flattened and
teardrop-shaped; on the midline, their closely approximated tips curve ventrally,
forming two narrow, strap-shaped processes that separate the alary processes of
the premaxillae.

Haughton identified what is actually the cracked anterior end of the fronto-

RELATIONSHIPS OF THE SOUTH AFRICAN FOSSIL FROG 3D

parietal in SAM-—9939 (H21) as fused nasals, and identified the actual nasals of
SAM-9948 as premaxillae. The ‘lateral spurs’ of Haughton’s presumed nasals,
however, are in fact those bones, representing their maxillary (lateral) processes.
The median foramen on the ‘nasals’ figured by Haughton (1931, fig. 1) is the
parietal foramen, in its normal position on the anterior end of the frontoparietal.

Frontoparietal

This large bone is visible on most specimens, but only a few examples are not
cracked or distorted; best preservation is seen on SAM-9965, K4596, K4597,
K4599, K4601, K4602 and K4608c. It is a single bone rather than being paired
as indicated by Haughton, broadly ovoid, bluntly rounded anteriorly and some-
what more pointed posteriorly (Figs 2A, 3). Its dorsal surface is relatively smooth
and flat, with a faint thickening of the lateral edges, but in younger specimens
the frontoparietal surface may be faintly convex. Haughton cited the fronto-
parietal as ‘depressed longitudinally in the middle’, but this is the result of
crushing. The parietal foramen is prominent and placed far forward, about
13-15 per cent of the total length from the anterior end.

SAM-—9939, on which Haughton’s figure | was based, has been severely
damaged by dried, cracked and exfoliated shellac and is difficult to interpret.
It is clear, however, that the posterior area suggested by him as having contained
a cartilaginous supraoccipital is actually a crack and that the bone he suggested
as being a scapula or opisthotic is actually a fragment of cleithrum.

Otoccipital

The shape of the paired otoccipitals is shown on many specimens, in par-
ticular SAM-—9941, 9944, 9946b, 9965, K4609, K4610b, K4620f and K4626b.
They are large and concave dorsally, with outlines of the semicircular canals
prominent (Fig. 2). Ventrally the otoccipitals do not meet on the midline
anteriorly (SAM-—9941), and each otic region is inflated into a hemispherical
chamber anterior to which there is a prominent channel for the eustachian tube.
Lateroventrally in SAM-K4610b the fenestra ovalis is visible, partially occluded
on the right by the columella. Posteriorly the otoccipitals meet on the midline
dorsally (SAM-—9946, K4603d) and fuse ventrally, forming the occipital condyles
(SAM-—K4610b). Small protuberances for cervical muscle attachment occur on
the posteromedial ends of the otic capsules (SAM-—K4603d, K4609a). No fora-
mina are visible.

Columella

The plectrum of the columella is shown in few specimens, but in SAM-
K4610b it is visible ventrally, in natural position in the fenestra ovalis (see
Fig. 2). A boomerang-shaped bone, it extends anterolaterally from the fenestra
ovalis, then abruptly bends anteriorly and extends forward perhaps as far as the
posterior tip of the maxilla (SAM—9965, although some forward displacement
of the columella has taken place here). No trace of operculum is present.

56

ANNALS OF THE SOUTH AFRICAN MUSEUM

RELATIONSHIPS OF THE SOUTH AFRICAN FOSSIL FROG 57

Sphenethmoid

The sphenethmoid is well developed (Fig. 2B); in all examples it is a single
ossification that extends posteriorly almost to the posterior borders of the palatal
vacuities (SAM-9941, K4600d, K4635). Just anterior to this point, large optic
foramina are visible (SAM—9941, K4609b, K4635). Crushing has usually
obscured the contours of the sphenethmoid, but in SAM—9941 and 9940 particu-
larly it is clear that its apparent wideness (observable on many specimens) is not
the result of crushing. In these two specimens the lateral sides are seen to con-
verge ventrally from their contact with the frontoparietals, and form a broad
wedge-shaped bone whose laterally-keeled, flattened ventral surface is capped by
the long, narrow parasphenoid. Anteriorly, as the sphenethmoid expands laterally
into the planum antorbitale, there is a constricted region (SAM-9941, K4597c,
K4609b). Anterolaterally, the arms of the planum antorbitale extend to the
maxilla (SAM—9955, K4597c). The smooth ventral and posterior surfaces of the
planum antorbitale suggest that this region is ossified, with the formation of a
lamellar surface layer, rather than being calcified. On SAM-—9940 a cross-section
reveals the presence anteriorly of paired internal channels for the olfactory tracts,
lined with smooth lamellar bone.

Palatine

No trace of this bone can be seen, although it may be present and fused to
the planum antorbitale.

Vomer

In SAM-K4614c, 9940 and K4609b, cross-sections show that the rod-like
cultriform process of the parasphenoid, anterior to the sphenethmoid constric-
tion noted above, overlies a roughly polygonal layer of bone that can only be
fused vomers, distinct but fused to sphenethmoid and parasphenoid (Fig. SA).
In another specimen (SAM-—K4635) no bone underlies the ventral surface of the
parasphenoid in this region, indicating that the vomer may sometimes remain
unfused (in this case lost).

Parasphenoid

Shape and form of the entire parasphenoid is shown well only in relatively
young metamorphosed (SAM-K4603b) or in late tadpole (SAM-—K46]12a,
K4613a) stages (Fig. 6A). It is a long, narrow bone, expanded very slightly in
its midportion in tadpole stages but more so in adults (SAM-—K4597a), an expan-

Fig. 2. Eoxenopoides reuningi. A. Dorsal view of restored skull. B. Ventral view. Based

primarily on SAM—9940, 9941, K4596a, K4606a. Lower jaw removed on right side of dorsal

view, left side of ventral view. Squamosal outlines and anterior extent of columella somewhat

conjectural (see text). Dotted-dashed line indicates approximate position of eustachian tube.

Note impression of eyeball and iris. C. Lower jaw in dorsal view. Composite of K4599a,

dentary, and K4606a, postdentary complex; position of large internal dentary foramen shown
in dotted line (see Fig. 4A for comparable foramen in maxilla).

58 ANNALS OF THE SOUTH AFRICAN MUSEUM

sion that does not exceed one-half the maximum width of the frontoparietal in
any individual (Fig. 2B). Its anterior end is narrow and pointed, overlapping the
premaxillae; posteriorly the bone narrows between the otoccipitals and termi-
nates near the posterior border of the skull in a spatulate tip (SAM—K4602b).

Pterygoid

This is a well-developed bone (SAM-9940, 9941, 9955) extending anteriorly
as a wide curved blade; as it reaches the maxilla it narrows, forming a channelled
tip (SAM-9940) that overlies the maxilla, curving along with the latter until it
terminates at or beyond the level of the posterior border of the planum antorbi-
tale (Fig. 2). Posteriorly a thin process extends towards the quadrate region,
curving ventrally into a sharp ridge (SAM-9955, H14). Posteromedially, a
flattened process extends from this ridge to floor the eustachian tube region; it
terminates near the midline in a point (SAM—9940), underlying (or perhaps abut-
ting) the pseudobasal process.

Quadrate

In most cases the quadrate is obscured by the lower jaw, but it appears to
have been well ossified (SAM-—9940), and not unusual in general aspect (Fig. 2).

Squamosal

Indistinct fragments are all that can be seen of the squamosal; crushing and
displacement of what must have been a lightly ossified bone has left little to
interpret. The bone was small and seems to have encircled the columella; no
processes can be seen, but on SAM—9965, K4596a and K4601 it seems to have
formed a conch within which the columella lay (see Fig. 2).

Mandible

This is best shown in SAM-9940, 9941, K4597c, K4599a, K4606a and
K4614c (see Fig. 2). The mandible is usually rather poorly preserved and crushed
in articulated specimens and is often difficult to distinguish from surrounding
bones. Examples of this condition are seen in SAM-—9940 and 9941. A perfectly
preserved isolated dentary has been extracted from specimen SAM—K4599a.
Posteriorly the dentary is expanded and keeled laterally (Fig. 2); anteriorly it is
widened at the symphysis, rather flattened both dorsally and ventrally, and per-
forated with numerous small foramina. No symphysials (mentomeckelians) were
observed on any specimen and the close approach to each other of the dentaries
at the symphysis region indicates that they were not present. The prearticular is
well shown in a number of specimens, in particular a perfectly preserved isolated
specimen SAM-K4606a (Fig. 2c). It is long and sharply pointed anteriorly and
complex posteriorly, where it includes the fused articular. Its posterior end
(SAM-9940, K4597a, K4606a) is flattened and expanded, the adductor muscle
attachment surface prominently developed with a squared-off anteromedial

RELATIONSHIPS OF THE SOUTH AFRICAN FOSSIL FROG

a eee

| |
\ f | |
leek Roe Sey ‘ ee ue Vas
A B ;

Fig. 3. Eoxenopoides reuningi. Relative growth of skull. A. SAM—K4608c, skull length 8,8 mm.
B. SAM-—9965, skull length 9,4 mm. C. K4596a, skull length 10,9 mm.

Imm

2mm

Fig. 4. Left maxillae of various pipids in dorsal view. A. Eoxenopoides reuningi, SAM-9940.
B. The same, SAM-K4596c. C. Xenopus muelleri. D. Xenopus tropicalis. E. Xenopus laevis.
F. Xenopus vestitus (the same configuration is seen in X. gilli, X. clivii and X. fraseri).

59

60 ANNALS OF THE SOUTH AFRICAN MUSEUM

border. Anteriorly the bone extends almost to the symphysis as shown in
articulated material.

Vertebral column

The method of vertebral articulation in articulated specimens cannot be
seen, but SAM-—4623b, an isolated vertebra, shows the presence of an anterior
condyle, indicating an opisthocoelous column. SAM-K4618a, the cross-section
of an isolated vertebra, indicates that the vertebrae were probably epichordal, as
no significant expansion of the centrum region occurs.

There are only six presacral vertebrae in all specimens in which the full
column is present (Fig. |). Haughton believed that either six or seven might be
present, but careful examination of all specimens available failed to disclose more
than six. The first vertebra is elongated and bears a long transverse process,
indicating that the true first vertebra (cervical vertebra or ‘atlas’) is fused with the
second presacral vertebra. This is confirmed in one of the tadpole specimens
(SAM-K4613a), which has a suture separating the two vertebrae (Fig. 6B—C).
In subsequent discussion, the fused cervical and second presacral will be referred
to as the first presacral to avoid confusion. The transverse processes of the first
three presacral vertebrae are elongated ; the tips of most examples are broken but
the second and third are complete and subequal in length in SAM-—9940 and
K4626b, the third complete in K4609a. In K4626b expansions of the transverse
processes near the centrum perhaps indicate the points of rib fusion but separate
ribs do not occur. Because of closeness to the skull, the shape of the first trans-
verse process Is generally not clear, but on SAM-9946, K4609a, and K4626b it
appears to be shorter than the second and third, and curved anteriorly, fitting
closely to the skull. Transverse processes of vertebrae 4—6 are relatively shorter,
anteriorly directed, somewhat expanded at the base and sharply pointed distally
(SAM-K4609a).

The centra and neural arches are blocky, the zygopophyses simple and not
strongly projecting. The ventral surfaces of the centra appear somewhat bulbous
on each side of a shallow midline depression. Dorsally, there is no trace of a
neural spine or crest on the neural arch surface in most specimens (e.g. SAM—
9939) but occasionally (SAM-—K4634) the posteromedial border of the neural
arch forms a minute projection. The posterolateral borders of the arch are slightly
concave whether or not this projection is present.

The sacrum and urostyle are fused and there is no trace of a postsacral
vertebra except on SAM-K4636 (Fig. 7C). The sacral diapophyses are widely
expanded on all specimens, at their tips approximating the length of the last three
presacrals.

Haughton discussed a number of ‘morphological stages’ that represent
variation in the vertebral and sacral region. His stage ‘A’ is misinterpreted; the
single specimen on which it is based (SAM-—9945, H17) has a series of cracks in
the sacral diapophysis rather than a series of diapophyses, and as noted above,
there are always six presacrals rather than ‘not greater than seven’. Haughton’s

RELATIONSHIPS OF THE SOUTH AFRICAN FOSSIL FROG 61

Fig. 5. A. Eoxenopoides reuningi, SAM—K4614c, dorsal view of ventral snout region, split in
a plane immediately dorsal to the parasphenoid and planum antorbitale. Note vomer ankylosed
to ventral surface of parasphenoid but remaining distinct from it. B. Ventral view of posterior
region of pelvis of Xenopus muelleri. C. The same view.of Eoxenopoides reuningi, SAM-9945.

62 ANNALS OF THE SOUTH AFRICAN MUSEUM

stage ‘B’ is the most common (85 % of specimens examined in which this region
could be interpreted clearly; n = 27); it is the unmodified condition, with normal
development of sacroiliac articulation and transverse processes (Fig. 1). As
Haughton indicated, there are in this stage only six presacral vertebrae. Examples
of this configuration are many, the best ones including SAM-—9938, 9942, 9956,
9964, K4596a, K4597d, K4603, K4609a, K4611, K4620, K4622, K4626 and
K 4627. Haughton’s stage ‘C’ could not be confirmed on the three specimens for
which he cited it: SAM—9948 (H6) and 9960 (H7); H22 could not be found and
lacks a SAM number. In SAM-—9948 the left transverse process of the last pre-
sacral is somewhat longer than usual, but is not sacralized. In K4596c, however,
a specimen not mentioned by Haughton, the left transverse process is expanded
and sacralized, that on the right somewhat enlarged. This is the closest approxi-
mation now available for stage ‘C’ (Fig. 7B). Stage ‘D’ of Haughton is valid,
based only on the specimen SAM-9946 (H23; Fig. 7A). There is a double
sacrum, the sixth (last) presacral having been completely sacralized yet remaining
suturally separate from the true sacrum. This is similar to specimen F24 of the
early Cretaceous pipid Thoraciliacus in which it appears in 2 of 247 specimens
(0,8 %; Nevo 1968). Stage ‘E’ of Haughton was cited as present in SAM-9942
(H11) and was presumed to represent a stage having only two. vertebrae with
short, anteriorly directed transverse processes rather than three, and with a post-
sacral vertebra showing transverse processes. This specimen is actually a stage
‘B’ with six presacrals and no postsacral vertebra visible. The closest approxima-
tion to this stage that could be found was in SAM—K 4636, in which a fused post-
sacral vertebra with large transverse processes is visible, the right one fused to the
sacrum; in this specimen the normal six presacrals are present so that this does
not represent a stage “‘D’ (Fig. 7C). Haughton’s ‘anomalous’ stage, with
asymmetry of sacral transverse processes, is well shown in the single specimen
on which he described it (SAM-—9965, H40); only the right transverse process of
the last presacral and the left transverse process of the sacrum are sacralized.
This is a common anomaly in frogs, both recent and fossil; Nevo (1968) has
described it in the Cretaceous pipid Thoraciliacus (specimen F128), finding it in
3 of 247 specimens (0,8 %), and Ritland (1955) observed it in living Ascaphus.

Shoulder girdle

The coracoid and clavicles are widely divergent and the clavicles do not meet
in a broad suture; the girdle was therefore presumably arciferal, although without
the cartilaginous epicoracoid horns this cannot be confirmed. The coracoids are
very slightly expanded at their medial ends (Figs 1, 8). A prominent groove is
present on the dorsal surface of the clavicles. The junction of scapula and clavicle
is difficult to interpret owing to poor preservation in all specimens, but is
relatively clear in SAM-—9940 and 9948 (H6). The scapula is relatively long and
somewhat expanded anteriorly; the presence of a cleft is not determinable. On
SAM-—9940 it appears that there was a suture between scapula and clavicle, but
this may be a crack, for several other cracks are present in the body of the clavicle

RELATIONSHIPS OF THE SOUTH AFRICAN FOSSIL FROG 63

‘Li el
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ez

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ow
aac
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me
ae on 7
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fee
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r
Uli.

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Fig. 6. Eoxenopoides reuningi, tadpoles. A. SAM—K4612a, ventral view of frontoparietal and

parasphenoid. B. SAM—K4613a, vertebral column showing partial fusion of first two presacral

vertebrae. C. Ventral view of same specimen, showing partially ossified otic capsule, outline
of body cavity, and hind limbs.

(see below). The cleithrum is widely forked and shown well on SAM-K4596a,
K4615c and K4626b. The questionable cleithrum noted by Haughton is correctly
identified, but is seen folded over on the scapula rather than its normal more

dorsal position; it is thus not applied to the anterior margin of the scapula as
Haughton supposed.

Humerus

In crushed form, the humerus is visible on many specimens (Fig. 1).
Fortunately, an uncrushed isolated left humerus was prepared from SAM-
K4610a and shows that the humeral head has a prominent trochlea for articula-

64 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 7. Eoxenopoides reuningi, variation in sacro-urostyle (see text). A. SAM—9946. B. SAM-
K4596c. C. SAM-K 4636.

2mm

Fig. 8. Eoxenopoides reuningi. A. Restoration of shoulder girdle, rotated so that all bones
appear in the same plane. Based primarily on SAM-—9940 and 9948, cleithrum from SAM—
K4615c. B. K4610a, left humerus in ventral view.

RELATIONSHIPS OF THE SOUTH AFRICAN FOSSIL FROG 65

tion with the scapula, from which a prominent crest extends distally to meet the
deltoid crest on the shaft. A deep channel occurs on the medial surface of the
deltoid crest, set off by another crest lateral to it. Distally the humeral ball is
small, as are the epicondyles; the fossa cubitus ventralis 1s prominently marked
(Fig. 8B).

Radioulna

The radioulna is of usual anuran form (Fig. 1), fused, slightly expanded
distally, and with a prominent fossa proximally for the ball of the humerus.

Carpus

SAM-—K4611 (H11), in which Haughton described the carpus, no longer has
that region preserved. Fortunately, K4599b, K4609b (Fig. 9A), K4615c and
K 4627 show reasonably well-preserved carpal arrangements in both ventral and
dorsal views. Two main elements in the proximal row of the carpals represent
radiale and a large ulnare; the latter has a prominent ventromedial process shown
well on two specimens that probably represents the fused intermedium (Fig. 9B).
Distally there are again two main elements: a lateral one probably representing
the fourth distal carpal, clearly shown on K4615c to include another element
medially, probably the third centrale, and a medial, subtriangular bone that
represents at least the second centrale. Distal carpals 1-3 are shown on K4627b,
as well as a small medial bone that represents the prepollex (or first centrale).

Metacarpals and phalanges

There are four metacarpals, well shown on SAM-K4599b, K4603a and
K4609b. Distally these three specimens show that the phalanges are relatively
short and that the phalangeal count is 2—2—3—3. Digit length increases pro-
gressively from digits 2-5, but the fingers are not much different in length
(Fig. 10A).

Pelvic girdle

The proportions of the pelvic girdle as a whole are shown on numerous
specimens, among them SAM-—9938, 9945, K4597f and K4609a. The ilium is well
seen in dorsal view on K4609a. No specimen permits a lateral view of the
acetabular region, but the general shape of the ischium and pubis area is shown
in K4609a and 9945 (Fig. SC). The ilium is elongate and there is an iliac symphysis
proximally on the midline. The iliac shaft on most specimens extends far forward,
nearly or actually touching the transverse processes of the third vertebra (SAM-
9965) but in others terminates near the end of the sacrum (SAM-K4596c). The
iliac shaft is robust but only a small iliac crest is present. The dorsal protuberance
is well developed but low and rounded, not protruding far from the shaft. The
ischium is usually obscured by the urostyle, but in SAM-—K4609a it is well
ossified and robust (Fig. SC). The pubis is well ossified as shown in SAM-9945,

66 ANNALS OF THE SOUTH AFRICAN MUSEUM

(See ee on ee eee eee tl
1mm

Fig. 9. A. Xenopus muelleri, right carpus in dorsal view. B. Eoxenopoides reuningi, restoration
of carpus, based primarily on SAM—-K4615c and K4627b. Note position of ventral impressions
of intermedium and central elements three and four, indicated in dotted line.

and forms a prominent transverse crest on the ventral surface of the pelvic girdle;
as in most recent pipids it is smoothly ossified rather than calcified (Fig. 5C).

Femur

The general shape of the femur is visible on many specimens. It is relatively
slender but well ossified. No details that indicate notable features are visible

(Fig. 1).

Tibiofibula

The general shape of the bone Is visible on many specimens. It is not notable
in any way (Fig. 1).

Tarsus

Tibiale and fibulare are intimately fused. This is visible externally on many
specimens, and a split longitudinal section (SAM-—K4598a) shows that this fusion
extends to the interior of the bones as well as to the epiphysial caps; the fused
bones have a common cavity.

A central element associated with the fibulare, one for the prehallux, and a
distal tarsal for the first metatarsal are visible on SAM—9945; centralia associated
with fibulare, prehallux, and metatarsals 1-3 on 9959; on K4609a distal tarsals
|—4 are visible, as well as the centralia for the prehallux and fibulare (Fig. 10C).

RELATIONSHIPS OF THE SOUTH AFRICAN FOSSIL FROG 67

Fig. 10. A-C. Comparison of right dorsal surfaces of hand and left ventral surfaces of foot
of Eoxenopoides reuningi. B—D. The same of Xenopus muelleri. A based on SAM-K4603a,
K4615c and K4627b, C based on K4609a; both specimens are from animals of equal size.

Metatarsals and phalanges

These bones are well shown on many specimens but are completely pre-
served on SAM-K4609a (Fig. 10C) and K4624b. They are elongated but not
unusual In any way.

The phalanges are generally not well preserved but are complete on SAM-
K4609a (Fig. 10C), K4624b and, except for the first digit, on K4627a and

68 ANNALS OF THE SOUTH AFRICAN MUSEUM

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4
:

2

aap

Fig. 11. £Loxenopoides reuningi. A. SAM—9938, holotype. B. SAM—K4609a, natural cast of

dorsal surface of skeleton (except for a few regions, including the hind foot, where bone is

still present). C. SAM-K4613a, tadpole, note outline of body cavity left by melanophore
concentration. Scale in centimetres.

RELATIONSHIPS OF THE SOUTH AFRICAN FOSSIL FROG 69

K4627b. They are relatively short. The prehallux is represented only by a single
phalanx, seen well in SAM-—9945 and 9959 as well as on K4609a. SAM-K4609a
and K4627a are well preserved and show that the phalangeal formula of digits
1-5 is 2-2-3-4-4; K4627b demonstrates with equal clarity that 2-2-3-4-3 may
occur as well. In the latter case the terminal phalanx is reduced and pointed,
indicating that this is of natural rather than of taphonomic origin, although
traumatic loss during the life of the animal cannot be ruled out.

COMPARISONS

Eoxenopoides reuningi possesses the suite of pipid character states given by
Estes (1975a and 19755) and no question exists as to its family allocation, as will
be demonstrated below. Comments on the structure of E. reuningi follow the
sequence of bones given in the description.

The maxillae are remarkable in that their anterodorsal surface is almost
identical with that in Xenopus muelleri and X. tropicalis in having an oral depres-
sion with a large posterior foramen that opens into the body of the bone. No
other species of Xenopus nor species of other pipid genera show this configura-
tion. The absence of teeth on the maxilla is unlike all pipids save some of the
South American species; this is not in any case a character state of more than
specific importance.

The nasals are paired, but otherwise resemble closely those of living Xenopus,
especially in having long, slender processes anteromedially that overlie the nasal
capsule and separate the alary processes of the premaxillae. The nasals may
occasionally remain paired in adult living Xenopus although usually they are
fused.

The frontoparietal is unlike that of any known pipid in detail, but resembles
that of the fossil Xenopus romeri in having an extensive dorsal skull table,
essentially parallel sides, and an anteriorly placed parietal foramen (Estes 19750).
It differs from that of all species of Xenopus, however, and resembles Hymeno-
chirus boettgeri and Pipa spp. in lacking muscle attachment on the dorsal surface
of the frontoparietal. Lambdoid crests are thus not developed except on the
lateral edge of the bone, and weak supraorbital ridges are present.

The otoccipital resembles that of Xenopus in detail, especially ventrally, in
which the expansion of the inner ear region and the configuration of the
eustachian tube channel closely resemble conditions in Recent X. muelleri and
the fossil species XY. romeri and X. pascuali. The pars media plectri of the
columella is well ossified, and bent into a boomerang shape, with otic and
tympanic processes as in Xenopus spp.

The sphenethmoid is widened dorsally, reflecting the width of the dorsal
table of the frontoparietal, which its laterodorsally diverging sides meet. Two
distinct internal channels for the olfactory tracts occur as in pipids in general;
the lack of flattening of these channels and their close approximation is as in
Xenopus spp. rather than in living South American species.

70 ANNALS OF THE SOUTH AFRICAN MUSEUM

The apparent rhomboid, toothless shape of the vomer and its tendency either
to fuse to the sphenethmoid and parasphenoid or to remain separate is as in
Xenopus romeri (Estes 1975a, figs 1-2). Lynch (1973) and Trueb (1973) have
called these bones ‘prevomers’. This designation has been shown to be in error
by Parrington & Westoll (1940), who demonstrated that the original suggestions
of the existence of a separate “prevomer’ in lower vertebrates was a
misinterpretation.

The cultriform process of the parasphenoid is narrow anteriorly as in
Xenopus and the fossil Cordicephalus, Thoraciliacus (Cretaceous, Israel; Nevo
1968), and Saltenia (Cretaceous, Argentina; Baez in preparation), rather than
widened as in Hymenochirus and the living South American pipids.

Posteriorly, the pterygoids resemble those in pipids generally, with expanded
medial processes flooring the common opening of the eustachian tube, but the
long, slender anterior process reaching almost to the level of the planum antor-
bitale is exactly as in Xenopus and Saltenia, lacking the expansion in that region
found in Pipa. Such an anterior extension of the pterygoids is also seen in the
Cretaceous genera Thoraciliacus and Cordicephalus, but curvature of the bones
appears to have been different in the latter, the anterior ends being laterally
concave rather than convex.

The squamosal, while poorly ossified, seems to have enclosed the columella
in the usual pipid manner, essentially as a cylinder for the reception of the
columella and tympanic membrane. In lacking the strong development of a:
zygomatic process some similarity to Hymenochirus and Saltenia occurs.

The dentary differs from that of other pipids in having an expanded sym-
physis region; in other pipids the dentary tapers to a point anteriorly. The post-
dentary compound element (articular and prearticular) resembles that of Xenopus
spp. closely except for being relatively straight at the posterior end rather than
having the articular region twisted laterally.

Aside from the reduction in vertebral number, the vertebral column as a
whole resembles that of many frogs in having a relatively short first transverse
process, posteriorly curved, elongated second and third transverse processes, and
short, anteriorly directed and pointed transverse processes of the fourth, fifth,
and sixth vertebrae. Among pipids, fusion of cervical and second vertebra is
characteristic of all species of Pipa, Hymenochirus and Pseudhymenochirus,
Xenopus tropicalis and X. romeri, and may occur occasionally in large X. /aevis.

The neural arches are imbricate, their surfaces smooth and their posterior
borders slightly concave on each side; the centra are opisthocoelous and probably
epichordal. In all these features they resemble pipids as a group.

The fused sacrum and urostyle region resembles that of Sa/tenia, Thoracilia-
cus and Cordicephalus in having sacral diapophyses only moderately expanded,
to about the length of the last three presacrals rather than equivalent to the
length of five or more vertebrae as in other pipid genera. The sacral vertebra is
clearly not compound; the necks of the sacral diapophyses are narrow and there
is at most one major spinal nerve foramen visible on their posterior surfaces.

RELATIONSHIPS OF THE SOUTH AFRICAN FOSSIL FROG 7)

Thus in Eoxenopoides reduction in vertebral number is partially a result of
synostosis (second presacral and cervical) and partially a case of true reduction by
deletion of the seventh vertebra through the sacrum (Lynch 1973: 141) and
constitutes the second such case described. Tihen (1960) has shown fusion of
cervical and second presacral vertebrae and true reduction of the column to
seven segments (hence six presacral vertebrae) in the bufonid Nectophryne.
Hymenochirus also has cervical and second presacral fused and only five pre-
sacrals in all, as does the bufonid Oreophrynella, but in both of these latter two
cases the reduction of posterior vertebrae has been effected by synostotic fusion
of presacrals with the sacrum, which shows an obvious compound nature, unlike
the situation in Eoxenopoides.

Broad expansion of sacral diapophyses has been considered primitive in
frogs, and Lynch (1973) cites them as ‘characterizing the four living archaic
families’ Ascaphidae, Discoglossidae, Pipidae and Rhinophrynidae. Primitive
fossil members of Ascaphidae and Discoglossidae have quite narrow sacral
diapophyses, however, and Cretaceous pipids as well as Eoxenopoides reuningi
have only moderate expansion (see Estes & Reig 1973). There is thus doubt that
the expanded condition (at least a widely expanded one) is primitive, as the fossil
record does not support this view. The separate sacral ribs of Triadobatrachus
are somewhat expanded, however, and the picture is therefore not entirely clear.
Triadobatrachus was very froglike in some respects but it had not yet achieved
anuran status, in spite of the statement of Wassersug (1975) to the contrary (see
Estes & Reig 1973). It is perhaps best in this case to give greatest weight to true
anuran conditions, and the fossil record suggests that an unexpanded or at most
a moderately expanded form was the most primitive.

The general configuration of the shoulder girdle does not offer much of
comparative interest. Scapula and clavicle are separated by what might be inter-
preted as a suture in SAM-9940; if it is one the clavicle dces not overlap the
scapula as much as in other pipids. It is possible, however, that this presumed
suture is, in fact, a crack for there is some distortion and several other cracks on
the clavicle of this specimen. If scapula and clavicle are fused, the condition is
derived, and resemblance is to Xenopus (excepting X. pascuali); if they are not,
- the condition is primitive. Because the apparent suture cuts transversely across
the bone rather than being on an angle, overlapping the scapula anteriorly as
would be expected in a pipid or other primitive frog, the author interprets the
scapula and clavicle as having been fused, but broken in an area of relatively
weak ossification. This ambiguity is the only one that has affected interpretation
of the Eoxenopoides fossils but it is not a fundamental one.

The humerus is very like that of pipids in general. The specific details of
trochlear head, crests and epicondyle configuration resemble those in Xenopus
spp. rather than those of the living South American pipids or Hymenochirus.
Both of the latter groups have either large epicondyles (Pipa) or strong develop-
ment of specialized humeral crests (Hymenochirus). The humeral ball is larger
and more protuberant than in most living or fossil pipids; only the Miocene

72 ANNALS OF THE SOUTH AFRICAN MUSEUM

Xenopus from Morocco (Vergnaud-Grazzini 1966) and the Recent Xenopus
tropicalis approach it in size. This would be a primitive feature, probably, since
the result is more like that of other groups of living and fossil frogs.

The radioulna offers no points of special interest.

The carpus is as in Xenopus laevis and X. muelleri, the only two species for
which cleared and stained preparations were available. The chief interest that it
offers is the evidence for fusion of carpal elements in pipids, and perhaps for
frogs in general. On the ventral view of the large ulnare of SAM—K4615c there
is a process probably representing a fused intermedium, as has been demonstrated
in living frogs (Trueb 1973). Of more interest is the fourth distal carpal, which
shows very well an element that is probably the attached and fused third centrale,
as suggested by Trueb (1973) to occur in living forms. The radiale shows a com-
parable situation; its internal side has a prominent centre of ossification (not a
process) that must be the fourth centrale, again as postulated by Trueb (1973).
A prepollex or first centrale is visible next to the first distal tarsal. The configura-
tion in Yenopus is somewhat different (Fig. 9A); there has been a shortening and
torsion of the carpus and the fourth distal carpal is much clongated, firmly
abutting and overlapping the radiale, suggesting that the former may include
the fourth centrale or at least have exhibited strong medial growth. The tarsus
of Eoxenopoides in its general configuration and apparent absence of significant
torsion is more primitive than Xenopus, and is generally comparable with (e.g.)
primitive discoglossids (Vergnaud-Grazzini & Wenz 1975, fig. 5).

The phalanges indicate relatively short fingers. Tinsley (1973) has given ratios
for the species of Xenopus; the ratio of first finger length/total lower forelimb
length in Eoxenopoides is 0,45, very close to the result for X. tropicalis. The
meaning of this is not clear as accurate measurements for other fossil and recent
species were not available. Measurements taken from photographs of the
Cretaceous Thoraciliacus and Cordicephalus indicate ratios of about 0,46 for
Thoraciliacus rostriceps and 0,52 for Cordicephalus gracilis. The former thus
resembles Eoxenopoides and X. tropicalis, the latter is closer to the range for the
longer-fingered species X. fraseri and X. laevis. As in Xenopus and many other
frogs, the third digit is the longest, but in Eoxenopoides there is less difference
between digit length than in Recent pipids. These proportions are reminiscent
of the early Cretaceous pipids from Israel and also of Palaeobatrachus spp. as
described by Spinar io72)

The pelvic girdle of Eoxenopoides is interesting in that as preserved, the
iliac shafts of most specimens extend forward to approach, touch or overlap the
transverse processes of the third presacral vertebra. This condition was used by
Nevo (1968) as a generic character for the early Cretaceous species of Thoracilia-
cus, not otherwise strongly separated from the sympatric Cordicephalus, as Nevo
states (Nevo 1968: 272). Whether or not the ilia overlap the ribs has in part to do
with the position of the sacrum relative to the ilia at the time of death or shortly
thereafter, as well as the position that may be characteristic in life. In general the
articulation of the sacrum in forms with narrow sacral diapophyses tends to be

RELATIONSHIPS OF THE SOUTH AFRICAN FOSSIL FROG 3

at the tip of the ilia (acroiliac of Nevo 1968), and in forms with moderately or
widely expanded diapophyses there is of necessity a more posterior articulation
(medioiliac of Nevo 1968), the ilia sometimes projecting in front of the sacrum.
Nevo has correctly rejected the latter as diagnostic of the ‘Eoxenopoididae’ of
Casamiquela (1961). In the author’s experience, specimens of living Xenopus that
have died as a result of desiccation show an extreme ‘medioiliac’ condition while
those in which the tissues remained moist tend towards an ‘acroiliac’ condition.
The critical factor in deciding the utility of a character state involving overlap
of ribs and ilia, then, is the relative length of the ilia as well as the determination
of the preferred orientation of the sacrum in life. Considerable movement is
possible at the sacroiliac joint (Whiting 1961) so that this latter feature is difficult
to assess. Nevo (1968) ascribed a functional significance to the overlap of ilium
and transverse process in the forming of a “synsacrum’. It is doubtful that actual
articulation of ilia with ribs occurred and that overlap is of functional signifi-
cance. In Xenopus overlap of transverse processes and ilia may occur (especially
in cleared and stained preparations made from desiccated specimens) but in this
case neither synsacrum nor joint occurs. There is no evidence that a joint occurred
at this overlap in Eoxenopoides, nor does it seem that an actual joint can be
demonstrated in Thoraciliacus. In deposits containing the latter two genera there
is no sedimentary evidence that the animals died in burrows (and hence might
have dried before final burial by overlying sediments). Nevo (1968) has carefully
quantified the differences between the species of Thoraciliacus and Cordicephalus,
and it is not suggested here that only one genus is represented. In Eoxenopoides
reuningi, however, the overlapping of transverse processes and ilia is not inter-
preted as demonstrating the presence of a synsacrum: the pattern of variation
seen in this species is more easily interpretable as variation in the position of the
sacrum relative to the ilia, a pattern of variation that is comparable to that seen
in Xenopus.

The ossification of the pubis and the development of an iliac symphysis are
as in pipids generally. The posterior extent of ischium is relatively great but does
not exceed that of other pipids.

The femur and tibiofibula offer no features of comparative interest.

The proximal tarsals (tibiale and fibulare) are fused, as in Xenopus and
other pipids, and is a fusion that extends beyond the epiphysial caps to the actual
cavities of the bones. Neither Trueb (1973) nor Lynch (1973) cited pipids as
groups showing fusion of these elements although the synostosis in living forms
is quite clear.

The presence of four distinct distal tarsals in Eoxenopoides is a primitive
condition; so far as the author is aware it is not shown in other frogs, which have
the third and fourth distal tarsals fused. The tarsal area as well as the carpus of
Eoxenopoides thus shows some primitive features.

The presence of a well developed prehallux is primitive and it occurs in
Xenopus and other pipids. Proportions of the foot are exactly as in X. muelleri
and X. /aevis, both of which species have relatively long toes.

74 ANNALS OF THE SOUTH AFRICAN MUSEUM

The metatarsals and phalanges are notable only in having an extra phalanx
in the fifth toe of two out of the three specimens in which the foot is well
preserved. The formula of 2—2—3-4—4 is also present in the fossil palaeobatrachid
frogs (Spinar 1972) and might be considered a derived character state linking
Eoxenopoides with the palaeobatrachids. Although pipids and palaeobatrachids
are believed to be related (Estes & Reig 1973; Estes 1975a) presumably this con-
dition in Eoxenopoides, which is lacking in more primitive pipids, is a parallel
condition. The presence of the extra phalanx gives no increase in length of the
digit when compared with the example having three phalanges in the fifth toe and
the significance of the condition cannot be ascertained at this time. Some living
frogs increased phalangeal count by developing intercalary cartilages, but these
are clearly distinguishable morphologically and functionally from the true
phalanges. The case in palaeobatrachids and Eoxenopoides is one of true hyper-
phalangy and appearance and ossification of all phalanges are similar. Utilizing
Tinsley’s (1973) proportion of tibia length/fifth toe length gives a ratio of about
0,82, almost within the range of the long-limbed Xenopus laevis group (ratios of
0,69-0,77, against 0,97—1,12 for the shorter limbed tropicalis group, Tinsley 1973).

RELATIONSHIPS OF EOXENOPOIDES REUNINGI

From the above comparisons a general assessment of the relationships of
Eoxenopoides reuningi is not difficult to make. Clearly pipid, it has only two
rather weak similarities to the living South American pipids: the lateral position
of muscle crests on the frontoparietal, indicating that adductor musculature did
not encroach on the dorsal skull table, and the fusion of the first two vertebrae.
Neither of these is exclusive to living South American pipids, however; both
may occur in species of Hymenochirus, and fusion of the first two vertebrae occurs
in the fossil Xenopus romeri and living X. tropicalis as well as in some other
groups of frogs (Lynch 1973). No resemblances to Hymenochirus exist beyond
the skull roof and vertebral configurations commented on above.

There is little resemblance to the Cretaceous fossil pipid Saltenia ibanezi
from South America. The fragility and general shape of the squamosal and the
moderate expansion of sacral diapophyses are the only resemblances to this
species, but neither resemblance is significant. Reduction of the squamosal can
be achieved independently and the moderate expansion of the sacral diapophyses
is probably primitive.

The greatest resemblances of Eoxenopoides reuningi lie with species of
Xenopus. As noted by Estes (1975a) understanding of the interrelationships of
these species is at a very early stage of development. Tinsley (1973, 1975) has
attempted morphological analysis of Xenopus based primarily on external
features and proportions. Although there are various degrees of intermediacy,
he has shown that two informal species groups of Xenopus may be recognized.
One group has relatively short limbs, fingers and toes and relatively small eyes,
and includes X. vestitus, X. gilliand X. tropicalis. The other group has relatively

RELATIONSHIPS OF THE SOUTH AFRICAN FOSSIL FROG a5

longer limbs, fingers and toes and larger eyes, and includes two groups of related
species: X. clivii and X. fraseri on the one hand, X. /aevis and X. muelleri on the
other. To a certain extent the two groups are linked by character states of YX.
vestitus. Tinsley has also suggested the possibility of grouping species of Xenopus
on the presence of four claws on the foot (X. fraseri, X. clivii and X. tropicalis)
or three (the remaining species). It is more likely that the four-clawed condition
is primitive and lost in parallel by some members of both groups, or that it is
an independently acquired specialization. The adaptive significance of a difference
in claw number is not known and the character seems not to display the kind of
taxonomic utility expressed by the other states mentioned.

Although there was insufficient osteological material to confirm Tinsley’s
groupings, the small sample available in this study supports his wide separation
of Xenopus tropicalis and X. laevis, and some of his other conclusions. The skull
of X. tropicalis is broad and short, with a wide, flattened braincase; that of
X. laevis is relatively longer and the braincase longer and narrower. In general
skull proportions and morphology X. muelleri seems least modified among
species of Xenopus, but in having a relatively long skull shows similarity to
X. laevis. X. vestitus has a wide frontoparietal as in X. tropicalis. The position
of X. clivii, X. fraseri, and X. gilli is not clarified by the small sample and for the
present Tinsley’s assessment of their relationships is accepted. The Palaeocene
species X. romeri from Brazil and the Miocene YX. stromeri from South West
Africa are related to X. tropicalis and X. muelleri respectively; the affinities of
X. pascuali from the Eocene of Argentina are now being studied; they may be
with X. muelleri rather than with the X. tropicalis group (Estes 1975a).

It remains now to assess the similarities and differences shown by Eoxeno-
poides reuningi to Tinsley’s groupings of the species of Xenopus. First, there are
a group of character states of E. reuningi that are probably primitive for pipids
in general. These include the nasal shape, anterior position of parietal foramen,
narrow cultriform process of parasphenoid, relatively long scapula, configuration
of the humerus, primitive carpus (Estes 1975a) and the flattened and expanded
transverse processes of the posterior vertebrae (Nevo 1968).

A further group of character states emphasizes the distinctiveness of
Eoxenopoides reuningi when compared with Xenopus: these include absence of
teeth in the upper jaw, reduction and fusion in the vertebral column, dentary with
lateral crest and expanded symphysis, and the combination of relatively short
fingers and long toes. With regard to the latter feature the general proportions
and length of hind limb and feet closely resemble those of X. muelleri and
X. laevis, but the shortness of the fingers is more as in_X. tropicalis. The fingers
of E. reuningi are almost subequal in length, reminiscent of conditions in the
Cretaceous pipids from Israel (Nevo 1968) and the palaeobatrachids (Spinar
1972). This combination of character states is found in no other pipid species
and seems distinctive enough to justify generic status for Eoxenopoides.

There remains for discussion only the group of character states in which
Eoxenopoides reuningi resembles various species of Xenopus. The hemispherical

716 ANNALS OF THE SOUTH AFRICAN MUSEUM

expansions of the inner ear region, in the details of their shape and relationship
to the channel for the eustachian tube, are almost precisely duplicated in Xenopus
muelleri and the fossil South American species X. romeri and X. pascuali (Estes
1975a; Casamiquela 1961) and are lacking in other species of Xenopus. Inner ear
expansion is found also in the Cretaceous pipids Thoraciliacus and Cordicephalus
from Israel and in the Northern hemisphere palaeobatrachids. Nevo (1968) has
noted that this expansion in his material from Israel is positively correlated with
size and that it is thus not necessarily a paedomorphic feature. In light of the
detailed resemblance of the otic expansion of E. reuningi to that of the relatively
primitive species of Xenopus noted above, this feature may be scored as a rather
weak resemblance to these species, although there is definite possibility of
parallelism in ear capsule expansion in pipids.

The rhomboidal, toothless shape of the fused vomers closely resembles that
of Xenopus romeri from the Palaeocene of Brazil and the living species YX. /aevis
and YX. clivii. Fused toothed vomers may occur in X. muelleri. The narrow,
dorsally channelled shape of the anterior process of the pterygoid is precisely
duplicated in all species of Xenopus. The shape of this process (if not its far
anterior extension, which is found in pipids in general) is a derived feature
showing resemblance to all species of Xenopus. Presence of an anterior, spoon-
shaped process of the maxilla that leads posteriorly into a large, circular foramen
is a close resemblance to XY. muelleri and X. tropicalis. As Figure 4 shows, there
is perhaps more detailed resemblance of XY. muelleri and Eoxenopoides reuningi.
It is interesting that this maxillary configuration is lacking in the more specialized
X. laevis on the one hand and_X. vestitus and X. gilli on the other, representing
both presumed species groups of Xenopus. Perhaps this condition is primitive,
and like the presence of three or four claws has been lost in parallel in derived
species of Xenopus.

In summary, there are a number of suggestive resemblances of Eoxenopoides
reuningi to Xenopus muelleri and its possible fossil relative X. pascuali on the one
hand, and to X. tropicalis and its fossil relative XY. romeri on the other. The
similarities of maxilla, ear capsule, and foot proportions to those of X. muelleri
suggest relationship to this relatively generalized species of Xenopus. Indications
of resemblance to YX. tropicalis are weaker and include the broad skull shape,
maxillary shape, and fused first two vertebrae; the fused toothless vomer and
fusion of first two vertebrae are like those of XY. romeri (although present in other
species as well).

None of the following character states of Eoxenopoides reuningi occurs in
any living pipid: moderately expanded sacral diapophyses, expanded posterior
presacral transverse processcs, anterior position of the parietal foramen (the latter
also found in the fossil species X. romeri, X. pascuali and X. stromeri), relatively
long scapula, and primitive carpus. This probably indicates an early origin of
Eoxenopoides reuningi relative to the diversification of species of Xenopus, in spite
of specialization of the former in loss of maxillary teeth and reduction of vertebral
number. It is therefore suggested that Eoxenopoides reuningi was an early offshoot

RELATIONSHIPS OF THE SOUTH AFRICAN FOSSIL FROG 77

of a primitive X. muelleri-like ancestor of Xenopus, a species that included the
above-mentioned primitive character states. Presence of these primitive states in
E. reuningi and their absence (excepting parietal foramen position) in species of
Xenopus indicates that origin of E. reuningi must have been within the Cretaceous
to have preceded appearance of Xenopus, the origin of which has been estimated
as prior to 91 m.y. (Cretaceous, late Turonian; Estes 1975a). None of the resem-
blances of E. reuningi to any of the species of Xenopus is specific or clear-cut
enough to suggest an origin within Xenopus although there is a well documented
series of character states confirming its close relationship to the living genus, as
documented above.

TAPHONOMY AND PALAEOECOLOGY

The specimens of Eoxenopoides reuningi occur throughout the sequence of
approximately 33 metres of clays forming the deposit, and occur in both black
carbonaceous and greenish grey rock types. There was no precise stratigraphic
control on the original collection of specimens, many of which came from the
dumps around the edge of the pipe. The second collection made by Boonstra
had more precise horizon data permitting determination of whether or not the
specimen came from high or low in the section. The morphology of the specimens
is, however, highly uniform, and no significant variation other than that noted
in the text could be detected. More precisely controlled collections may in future
demonstrate some stratigraphic progression but the author does not believe that
it would achieve or even approach the specific level.

There has been relatively little disarticulation of material; most specimens
are relatively well articulated and complete although many have lost parts of the
skeleton by breakage in collection. Some maceration occurred, however, as
isolated skulls are present and the sediment contains fragments of bones or
complete elements from time to time. It is possible that more disarticulated
material was originally present but that it was not collected. The excellent
condition of preservation of the specimens indicates that after death the speci-
mens were buried rather quickly in conditions that did not permit extensive
decomposition. The wide distribution of the specimens throughout the section
indicates that no sudden kill was involved although frequent repetitive conditions
such as lake overturn could have caused periodic or seasonal kills.

Most of the frog remains were of adult, metamorphosed individuals; only
a small number of tadpoles was recovered, all of which were rather late stages
with small but well-preserved limbs. The metamorphosed specimens are all well
ossified and the individual elements distinctly and completely formed; few
indicate specimens that had recently metamorphosed, as far as comparisons
with living Xenopus permit this to be determined. Size range of the specimens is
relatively limited. Total skull width ranges from 6,6 to 10,6 mm, M = 8,8,
N = 17, a size close to that of the unmetamorphosed specimens. It is thus
probable that Eoxenopoides reuningi was either a relatively small species or that

78 ANNALS OF THE SOUTH AFRICAN MUSEUM

all specimens were relatively young and do not indicate maximum size. Wide
distribution argues for the former alternative and the author believes that the
expressed size range closely matches that of the species.

Close similarity in body proportions to those of living Xenopus, particularly
the elongated metacarpals and metatarsals and the specialized form of the
humerus, indicate that Eoxenopoides reuningi was as aquatic as the living Xenopus
and that its feeding habits were similar, making use of a stereotyped antero-
posterior motion of arm and elongated hands to create currents that brought
olfactory sensations to the nasal epithelium and to aid in forcing food into the
mouth.

What little of the flora is known indicates a relatively dry climate, with rain
limited to the warm season (Axelred 1976 in Jitt.).

ACKNOWLEDGEMENTS

It is a pleasure to acknowledge here the debt that this paper owes to the
careful account prepared in 1931 by Dr S. H. Haughton, who correctly placed
this interesting fossil frog to family at a time when little of substance on fossil
frogs was available in the literature.

I am grateful for all the assistance graciously provided by Dr Michael Cluver
and his laboratory staff during my stay at the South African Museum, and I also
thank the Director, Dr T. H. Barry, for his courtesy. Drs A. M. Baez, Z. Spinar,
R. Tinsley and R. Wassersug read the manuscript critically. This study was made
possible by a grant from the National Geographic Society, which is gratefully
acknowledged.

REFERENCES

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S. Afr. 19: 255-258.

CASAMIQUELA, R. 1960. Datos preliminares sobre un pipoideo fésil de Patagonia. Actas Trab.
1™° Congr. Sudam. Zool. 4: 17-22.

CASAMIQUELA, R. 1961. Un pipoideo fdsil de Patagonia. Revta Mus. La Plata, Sec. Paleontol.
(n.s.) 4: 71-123.

Cookson, I. 1947. Plant microfossils from the lignites of Kerguelen Archipelago. Rep. Br. Aust.
N. Z. Antarct. Res. Exped. (1929-31) Sr. (A) 2: 127-142.

CRANWELL, L. 1961. Antarctica: cradle or grave for its Nothofagus? In: Proc. 10th Pacif. Sci.
Congr.: Ancient Pacific Floras. University of Hawaii Press.

DINGLE, R. & GENTLE, R. 1972. Early Tertiary volcanic rocks on the Agulhas Bank, South
African continental shelf. Geol. Mag. 109: 127-136.

Estes, R. 1975a. Fossil Xenopus from the Paleocene of South Africa and the zoogeography of
pipid frogs. Herpetologica 31: 263-278.

Estes, R. 19755. Xenopus from the Paleocene of Brazil and its zoogeographic importance.
Nature, Lond. 254: 48-50.

Estes, R. & REIG, O. A. 1973. The early fossil record of frogs: a review of the evidence. Jn:
VIAL, J., ed. Evolutionary Biology of the Anurans. Columbia: University of Missouri Press.

RELATIONSHIPS OF THE SOUTH AFRICAN FOSSIL FROG 79

HauGutTon, S. 1931. On a collection of fossil frogs from the clays at Banke. Trans. R. Soc.
S. Afr. 19: 233-249.

Hecut, M. 1963. A revaluation of the early history of the frogs. Part II. System. Zool. 12:
20-35.

KIRCHHEIMER, F. 1934. On the pollen from the Upper Cretaceous Dysodil of Banke, Namaqua-
land (South Africa). Trans. R. Soc. S. Afr. 21: 41-50.

Kroner, A. 1973. Comments on ‘Is the African plate stationary ?’. Nature, Lond. 243: 29-30.

Kuan, O. 1941. Die eozanen Anura aus dem Geiseltale. Nova Acta Leopoldina (n.F.) 10:
313-376.

Kuan, O. 1961. Die Familien der rezenten und fossilen Amphibien und reptilien. Bamberg:
Meisenbach KG.

Lyncu, J. 1973. The transition from archaic to advanced frogs. /n: VIAL, J., ed. Evolutionary
Biology of the Anurans. Columbia: University of Missouri Press.

Nevo, E. 1968. Pipid frogs from the early Cretaceous of Israel and pipid evolution. Bu//. Mus.
comp. Zool. Harv. 136: 255-318.

Paropi Bustos, R. 1962. Los anuros cretacicos de Puente Morales (Salta) y sus vinculaciones
con Shelania pascuali Casamiquela (Chubut) y E. reuningi Haughton, de Africa del Sur.
Reyvta Fac. Cienc. nat. Salta 1: 37-40.

PARopI Bustos, R. & KRAGLIEVICH, J. 1960. A proposito de los anuros cretacicos descubiertos
en la provincia de Salta. Revta Fac. Cienc. nat. Salta 1: 37-40.

PaARop!I Bustos, R., FIGUEROA CAPRINI, M., KRAGLIEVICH, J. & DEL Corro, G. 1960. Noticia
preliminar acerca del yacimiento de anuros extinguidos de Puente Morales. Revta Fac.
Cienc. nat. Salta 1: 1-20.

PARRINGTON, R. & WESTOLL, T. S. 1940. On the evolution of the mammalian palate. Trans.
R. Soc. Lond. (B) 230: 305-355.

Reic, O. A. 1958. Proposiciones para una nueva macrosystematica de los anuros. Nota pre-
liminar. Physis 21: 109-118.

Reic, O. A. 1959. Primeros datos descriptivos sobre los anuros del Eocretaceo de la Provincia

. de Salta (Rep. Argentina). Ameghiniana 1: 3-7.

RENNIE, J. 1931. Note on fossil leaves from the Banke clays. Trans. R. Soc. S. Afr. 19: 251-253.

RITLAND, R. 1955. Studies on the post-cranial morphology of Ascaphus truei. 1. Skeleton and

. spinal nerves. J. Morph. 97: 119-178.

SPINAR, Z. 1972. Tertiary frogs from Central Europe. The Hague: W. Junk.

TIHEN, J. 1960. Two new genera of African bufonids, with remarks on the phylogeny of related
genera. Copeia 3: 225-233.

TINSLEY, R. 1973. Studies on the ecology and systematics of a new species of clawed toad, the
genus Xenopus, from western Uganda. J. Zoo/. 169: 1-27.

TINSLEY, R. 1975. The morphology and distribution of Xenopus vestitus (Anura: Pipidae) in
Central Africa. J. Zool. 175: 473-492.

TRUEB, L. 1973. Bones, frogs, and evolution. Jn: VIAL, J., ed. Evolutionary Biology of the
Anurans. Columbia: University of Missouri Press.

VERGNAUD-GRAZZINI, C. 1966. Les amphibiens du Miocéne de Beni-Mellal. Notes Serv. Géol.
Maroc 27: 43-69.

VERGNAUD-GRAZZINI, C. & WENZ, S. 1975. Les discoglossidés du Jurassique supérieur du
Montsech (Prov. de Lérida, Espagne). Annl/s Paleont. 61: 19-36.

WASSERSUG, R. 1975. The adaptive significance of the tadpole stage with comments on the
maintenance of complex life cycles in anurans. Am. Zool. 15: 405-417.

WHITING, H. 1961. Pelvic girdle in amphibian locomotion. Jn: Vertebrate Locomotion. Zool.
Soc. Lond. Symposium 5: 43-57.

80

ANNALS OF THE SOUTH AFRICAN MUSEUM

centralia 1-4
clavicle
cleithrum
coracoid
columella
dentary

distal carpals 14
frontoparietal
intermedium
ilium

ischium
maxilla

nasal
otoccipital
optic foramen
prepollex or Cl

ABBREVIATIONS
pa prearticular—articular complex
pas parasphenoid
pbp pseudobasal process
pla planum antorbitale
pmx premaxilla
pt pterygoid
pu pubis
qu quadrate
tr radiale
ru. radioulna
sc scapula
sph sphenethmoid
sq squamosal
u__ulnare
vo vomer

6. SYSTEMATIC papers must conform with the International code of zoological nomenclature
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Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (figs 8a—b).

Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

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RICHARD ESTES

RELATIONSHIPS OF THE SOUTH AFRICAN
FOSSIL FROG
EOXENOPOIDES REUNINGI (ANURA, PIPIDAE)

eS CR SE ee

aE EE: = Se aes == a

ISSN 0303-2515

‘MUSEUM

JUNE 1977 |

F THE SOUTH AFRICAN

APE TOWN

VOLUME 73 PART 3

os
A

(

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Examples (note capitalization and punctuation)

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. 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, iat masses and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4 }:

THIELE, J. 1910. Mollusca: B. Polypeeahent 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 73 ~~ Band
June 1977 Junie
Part —3 Deel

CRETACEOUS DEPOSITS NEAR BOGENFELS,
SOUTH WEST AFRICA

By

HERBERT CHRISTIAN KLINGER

Cape Town Kaapstad

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CRETACEOUS DEPOSITS NEAR BOGENFELS,
SOUTH WEST AFRICA

By
HERBERT CHRISTIAN KLINGER
South African Museum, Cape Town

(With 8 figures)

[MS. accepted 20 January 1977]

ABSTRACT

Part of what is believed to have been the Cretaceous outcrop briefly described by Haughton
in 1930 was relocated. Mining operations in the area have virtually destroyed the outcrop.
Because of its stratigraphic importance, a description and locality data are given for future
investigators. The informal term ‘Wanderfeld IV’ beds is proposed for the Cretaceous
(Cenomanian) sedimentary sequence exposed near Bogenfels. The fossils mentioned by
Merensky in 1910 from an unknown locality in the Sperrgebiet possibly indicate the presence
of Santonian or Campanian sediments. These may be correlated with offshore sediments which
have yielded Sphenoceramus aff. S. schmidti.

CONTENTS

PAGE

Introduction . : : . ee |
Description of locality and exposure : : nee
Description of stratigraphic sequence. : 2 8S
Discussion of results . ; : ; : ; so 8D
In situ vs remanié . : , : : ~. 85
Tertiary/Cretaceous sewed : : ; ay 85
Age of Cretaceous sediments . : Se EST
Nomenclatorial status of Cretaceous deposits ay
Acknowledgements . 3 : : : : ae ee ||
References . . Z ; 3 ; : : 2 OD

INTRODUCTION

The presence of onshore Cretaceous deposits in South West Africa was first
suspected by the discovery and subsequent identification of Protocardia hillana
(Sowerby), Turritella bonei Baily and Turritella (Haustator) meadi Baily by
Merensky (1910: 18) in the first decade of this century. The actual locality from
which these fossils were collected, however, remains unknown. The first unequi-
vocal evidence for the onshore presence of Upper Cretaceous marine sediments
was the fortuitous discovery by Haughton of a shelly bed consisting mainly of
Exogyra cf. columba(= Rhynchostreon cf. suborbiculatum) and a single ammonite
named Placenticeras merenskyi sp. nov. (= Proplacenticeras merenskyi) in an
area well known for abundant Tertiary fossils, previously described by BGhm
& Weissermel (1913), BGhm (1926) and Weissermel (1926). The results of the

81

Ann. S. Afr. Mus. 73 (3), 1977: 81-92, 8 figs.

82 ANNALS OF THE SOUTH AFRICAN MUSEUM

discovery were published by Haughton in 1930 (1930a: 61-63; 1930b: 361-365).
Comparison of Proplacenticeras merenskyi with placenticeratids from other areas
led Haughton to believe that the Bogenfels deposits were ‘not earlier than the
Lower Senonian’ (19306: 364). Unfortunately the locality data given were vague.
The Tertiary deposits were described as being ‘just to the east and north-east
of Wanderfeld IV, north of the old Bogenfels station, and is a small depression
running into the Langental’, and the Cretaceous deposits as being ‘at a spot a
few hundred yards to the south’ (of the Tertiary outcrops) (Haughton 19305:
362).

Subsequent to Haughton’s discovery of the shelly bed and the single
ammonite, no one has apparently been able to relocate the Cretaceous outcrop,
and doubt has arisen as to whether the Cretaceous fossils were indeed in situ, or
reworked into the Tertiary sediments, despite Haughton’s (1930b: 361) adamant
rejection of this possibility. One of the reasons why subsequent investigators
failed to relocate the outcrop is probably because mining operations have
removed virtually all the sediment overlying the bedrock, leaving hundreds of
sieved gravel heaps in their place.

Ziegler (1969: 5), having visited the area near Wanderfeld IV, and having
examined the single specimen of Proplacenticeras merenskyi which is housed in
the South African Museum, concluded that ‘in view of other palaeo-evidence
and the absence of other Ammonites in the section we have to consider it as a
reworked specimen’. On the basis of the macro-fauna a Miocene, probably ~
Burdigalian age, was proposed by Ziegler for the sediments outcropping near
Wanderfeld IV. The author himself has been quoted as considering the fossils
to be remanié (see Cooper 1974: 88) following an unsuccessful visit to the area
in 1970.

During another visit to the area in 1975 with W. G. Siesser the author was
fortunate enough to relocate part of what appears to have been the outcrop
described by Haughton. In view of the fact that this is the only known marine
onshore Cretaceous deposit on the whole west coast of Africa south of Angola,
an as accurate as possible locality map and a description of the present conditions
of the remains of the outcrop and its relation to the overlying Tertiary sediments
are provided.

DESCRIPTION OF LOCALITY AND EXPOSURE

In the sketch map (Fig. 1) compiled from an uncorrected aerial photograph
and Kaiser’s 1926 map no. 3, Granitberg, the depression running into the
Langental is shown due east of Wanderfeld IV (Fig. 2).

Subsequent to Haughton’s description of the locality, extensive mining
operations were undertaken. The conglomeratic sediments overlying the bedrock
were stripped and sieved for diamonds. The area is littered with gravel mounds;
the largest concentrations are indicated on the map. In places where the sediment
has not been disturbed, the Tertiary fossils occur as typical deflation deposits.

CRETACEOUS DEPOSITS NEAR BOGENFELS, SOUTH WEST AFRICA 83

De
Wanderfe

LEGEND
Cretaceous
outcrop

=] Tertiary &
Quaternary

Basement
<1} dolomite
a Gravel
dumps
@) Tertiary
hillock

@) Dumped rails
& sleepers

Narrow ~
gauge rail
gts

Pspo00 |

Fig. 1. Sketch map indicating locality of Tertiary and Cretaceous outcrops.
(After Kaiser 1926 and uncorrected aerial photograph.)

The surface is littered with rusty-brown turritelid gastropods, heterodont
bivalves, ostreids, bryozoans, selachian teeth and occasional specimens of the
nautiloid Aturia lotzei. An important topographical feature is a low hillock in
the western part of the depression (Fig. 3), consisting of green, sandy silts and
hard, greyish-white concretionary layers with abundant Tertiary fossils. As this
is the only major topographic feature in the depression, both the author and
Ziegler here searched in vain for the Rhynchostreon bed and incorrectly con-
cluded that the material described by Haughton was probably reworked.

The remains of the Cretaceous outcrop, at 27°23’03”S 15°24’20’E occurs
ca 70 metres due east of an intersection in the narrow-gauge railroad amidst
gravel dumps on the eastern slopes of the depression (Fig. 5). The intersection in
the narrow-gauge railroad is marked by dumped rails and sleepers, and is

84 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 2. View of depression looking north.

Eocene Section

At
A aag h
SANG OE LS

Fig. 3. View of Tertiary hillock in depression. Vehicle tracks on right foreground for scale.

CRETACEOUS DEPOSITS NEAR BOGENFELS, SOUTH WEST AFRICA 85

approximately 160 metres south of the Tertiary hillock mentioned above. The
remains of the Cretaceous exposure are represented by a few loose concretions,
ca 20 cm thick, crammed with exogyrans (Figs. 4-5) littering the surface over an
area of about 5 m?. Abundant loose exogyrans occur both on the surface and in
the surrounding gravel dumps.

An intensive search in this area for more of this material proved to be
negative.

DESCRIPTION OF STRATIGRAPHIC SEQUENCE

The stratigraphic section presented in Figure 6 is very diagrammatic and
composite, consisting of the section measured at the Cretaceous locality and also
at the Tertiary hillock as shown in Figure 3. Virtually everything is covered by
sand at the Cretaceous locality and lack of time and sandstorms made trenching
physically impossible. A few shallow pits were dug at suitable places to remove
the sand and samples were collected to gain some idea as to the stratigraphic
succession. The section at the Tertiary hillock is based on trenching undertaken
during 1970. Details of the Tertiary micropalaeontology will be given by W. G.
Siesser at a later date.

DISCUSSION OF RESULTS
In situ vs remanié

In its present condition, only trenching can provide a definite answer to the
question of whether the material is reworked or not. According to Haughton
(pers. comm.) the outcrop was still intact at the time he and A. L. du Toit had
visited the area. Both he and Du Toit were in agreement that the material was
definitely not reworked. As to the figured outcrop, the following data favour the
concept of the material being in situ. According to Haughton (19305: 362), the
Tertiary fossiliferous sediments were situated at 70 m above sea-level, and the
Cretaceous ones at 60 m. This relationship, as well as the presence of the
felspathic grit above the Rhynchostreon bed mentioned by Haughton, is also
seen in the present description of the stratigraphic sequence. Nearly all the fossils
found on the surface show some damage or other due to current sand abrasion,
which would be difficult to distinguish from damage due to reworking. The
presence of undamaged Cretaceous bivalves in the silty sand underlying the
Rhynchostreon bed, however, militates against reworking. Furthermore, the
lithologies exposed directly above and below the Rhynchostreon bed point to
low-energy milieu, and it is difficult to envisage large fragments of the shell bed
being actively eroded and transported under these conditions.

Tertiary/Cretaceous boundary

The shingle bed described by Haughton (1930b: 362) as forming the base
of the Turritella (Tertiary) beds was not exposed during trenching. The felspathic
grit bed exposed above the Rhynchostreon (Cretaceous) bed with an apparent

ANNALS OF THE SOUTH AFRICAN MUSEUM

86

"LX SWnjojnsiqsogns uoadjsoyoUudyy YA POULIN UOTEINUOD 9S00'T “p ‘317

CRETACEOUS DEPOSITS NEAR BOGENFELS, SOUTH WEST AFRICA 87

Fig. 5. View of remains of Cretaceous exposure amidst rubble heaps. Prospecting shovel on
right for scale.

dip towards the north-east could, however, be followed all along the eastern slopes
of the depression up to a short distance from the first outcrops of fossiliferous
Tertiary sediments. Whether the felspathic grit forms the base of the Tertiary or
represents termination of Cretaceous sedimentation is not known. Micropalaeon-
tological samples proved to be of no assistance in this matter. The boundary
between the Cretaceous and Tertiary systems is between beds (e) and (i) in the
section.

Age of Cretaceous sediments

Micropalaeontological samples taken directly above and below the Rhyncho-
streon bed were either barren or contained poorly preserved non-diagnostic or
unidentifiable micro-faunas. Subsequent to Haughton’s discovery of the locality,
no new faunal elements have been found here. The only indicators of age are
Proplacenticeras merenskyi, here refigured as Figure 7, and Rhynchostreon
suborbiculatum (Fig. 8A—G).

On the assumption that the group of placenticeratids, to which P. merenskyi
belongs, did not occur earlier than the Coniacian, Haughton (19305: 364) con-
cluded that it would not be possible to place the age of these deposits ‘any earlier
than the Lower Senonian’. Dingle (1973: 346) tentatively substantiated this and
postulated that these deposits could possibly represent the base of the uncon-
formity developed in south-eastern Africa described by Kennedy & Klinger
(1971). On the basis of the occurrence of Rhynchostreon cf. suborbiculatum in the
uppermost Cenomanian of Salinas, southern Angola, Cooper (1974: 87-8) sug-
gests a Late Cenomanian age for the Bogenfels deposits.

Proplacenticeras generally has a time range of Cenomanian to Coniacian

88 ANNALS OF THE SOUTH AFRICAN MUSEUM

Light brown sand and silt with macrofossils and pebbles. .
Concretionary layer with bivalves. Aturia found here.

Massively bedded light brown sandy silt with occasional
macrofossils.

Concretionary layer with bivalves and pebbles.

Light sandy silt with calcareous and siliceous concretions.
Ferruginous concretionary layer; abundant bivalves and agates.
Clayey and sandy silt; abundant articulated bivalves with

conspicuous green colouring
Concretionary layer; abundant gastropods; green coloured

Sandy silt; ferruginous concretions and abundant art. bivalves.
Light grey clayey sandy silt; massively bedded; pebbles.
Interval between felspathic grit and lowest beds in Tertiary.
Hillock; interval estimated not to exceed 1-2 metres.

3m_ coarse, felspathic grit with abundant qz and felspar
grains. Grains angular, clean. This felspathic grit layer
described by Haughton (1930a:362) can be followed all the
way northwards down the slope of the eastern side of the

depression until it finally disappears in the floor of
the depression in close proximity of the Tertiary hillock.

oO! eee eee ce ae ene 1s EOCENES: = oeaanene

Covered by sand
No pit samples possible

Covered by sand. Pit sample yielded olive-green
silt turning yellow-brownish upon weathering

Hard, calcified Rhynchostreon shell bed. Valves
both in life position and disarticulated. Abundance
suggests bioherm. Haughton reported P.merenskyi
from here.

Yellow-brown silty sand with abundant heterodont bivalves
and Rhynchostreon towards top.

Covered by sand. Pit sample yielded yellow-brownish
grit and rounded pebbles.

Blue grey and brown dolomite with typical fluted Im
weathering due to sand abrasion.

P CAMB______......... CRETACEOUS

Fig. 6. Diagrammatic section of Cretaceous and Tertiary exposure near Bogenfels.

CRETACEOUS DEPOSITS NEAR BOGENFELS, SOUTH WEST AFRICA 89

Fig. 7. SAM-10569 Proplacenticeras merenskyi (Haughton). Holotype; x1.

(Wright 1957: L390), but may occur as early as the Upper Albian (see Collignon
1963: 126). Rhynchostreon is generally restricted to the Cenomanian and Turonian
(Stenzel 1971: N1123-4). It is suspected, however, that typical forms of R.
suborbiculatum, such as these found near Bogenfels, are restricted to the Early
and Middle Cenomanian throughout the world. As Haughton states that the
P. merenskyi specimen was found in the Rhynchostreon bed, the possibility that
more than one age is represented may be excluded. The concurrent range zone
of these two species thus suggests a Cenomanian age for the RAynchostreon shell
beds near Bogenfels.

90 ANNALS OF THE SOUTH AFRICAN MUSEUM

8. A-G. Rhynchostreon suborbiculatum. Specimens SAM-—PCS5498-5503. A-—C, F-G
natural size. D-E x<1,2. H. Sphenoceramus aff. S. schmidti. SAM—PCS5504; x1.

CRETACEOUS DEPOSITS NEAR BOGENFELS, SOUTH WEST AFRICA 9]

At this stage comments on the fossils mentioned by Merensky seem oppor-
tune. The locality from which these fossils were obtained is unknown, as
Merensky did not collect them himself, but obtained them from Mr Schettler, a
representative of the Koloniale Gesellschaft. According to Haughton (1930b:
361) the Tertiary exposure near Bogenfels was shown to Schettler, but it is
unknown if the fossils were indeed collected there. According to Merensky (1910:
18), ‘News of further finds (diamonds) which were made... opposite Plum-
pudding island, arrived shortly before my departure, so that I could not visit
them. Mr. Schettler ... having just returned from this locality, showed me
samples from these.’ Reference to Plumpudding Island suggests a locality in the
vicinity of Kakaoberg or Buntfeldschuh, south of Bogenfels.

These fossils could not be traced, and Merensky’s identifications receive the
benefit of the doubt. Merensky assumed that the fossils identified by him were of
the same age as the fauna from the Umzamba Formation. If this assumption is
correct, an age of Middle Santonian to Early Campanian can be assigned to these
fossils (Klinger & Kennedy 1977). Cooper suggests a Santonian age on the basis
of the fossils identified by Merensky.

To date, no other fossils have been found to either confirm or negate the
existence of these ‘hypothetical’ Santonian/Campanian deposits. Offshore
drilling at 28°23’S 15°25’E yielded a single specimen of a juvenile inoceramid,
tentatively identified as Sphenoceramus aff. S. schmidti (Fig. 8H). The latter seems
to indicate a Middle Santonian to Late Campanian age for the offshore exposures.
Being of the same age, the offshore deposits could possibly be the equivalents of
the onshore deposits which allegedly yielded the fauna identified by Merensky.

Nomenclatorial status of Cretaceous deposits

As the outcrop at Wanderfeld IV is not, at present, a mapable unit, it is
suggested that it should retain informal status as the “‘Wanderfeld IV beds’ until
similar lithologies are found onshore or offshore.

ACKNOWLEDGEMENTS

The author wishes to express his gratitude to Dr C. Stocken, Mr R. Little
and Mr D. Minney of Consolidated Diamond Mines of South West Africa for
their generous assistance without which field work would have been impossible.
Dr W. G. Siesser of the Marine Geoscience Unit of the University of Cape Town
organized the 1975 excursion and his invitation to join him in this venture is
gratefully acknowledged. Thanks are due to Dr S. H. Haughton for his comments
on the locality and the outcrop. Dr E. G. Kauffman (Washington) kindly
identified the exogyrans and the inoceramid; the latter was donated by Mr
R. Joynt of the Marine Diamond Corporation. Dr H. dela R. Winter of SOEKOR
gave permission for parts of Ziegler’s unpublished report to be quoted, for which
the author is grateful. Mr C. Hunter and Mr V. Branco kindly assisted with the
illustrations.

92 ANNALS OF THE SOUTH AFRICAN MUSEUM

REFERENCES

Boum, J. 1926. Uber Tertiare Versteinerungen von den Bogenfelser Diamantfeldern. In:
Kaiser, E., ed. Die Diamantenwiiste Siidwestafrikas 2. Berlin: Dietrich Reimer (Ernst
Vohsen).

Boum, J. & WEISSERMEL, W. 1913. Uber Tertidre Versteinerungen von den Bogenfelser Diamant-
feldern. Beitr. geol. Erforsch. deutsch. Schutzgebiete 5: 1-58.

CoLLiGNon, M. 1963. Atlas des fossiles caracteristiques de Madagascar (Ammonites). Fasc. X.
Albien. Service Geologique Tananarive.

Cooper, M. R. 1974. The Cretaceous stratigraphy of South Central Africa. Ann. S. Afr. Mus.
66: 81-107.

DINGLE, R. V. 1973. The geology of the continental shelf between Luderitz and Cape Town
(South Africa) with special reference to Tertiary strata. J. geol. Soc. Lond. 129: 337-363.

HAUGHTON, S. H. 1930a. Note on the occurrence of Upper Cretaceous marine Beds in South
West Africa. Trans. geol. Soc. S. Afr. 33: 61-63.

HAUGHTON, S. H. 19305. On the occurrence of Upper Cretaceous marine fossils near Bogenfels,
South West Africa. Trans. R. Soc. S. Afr. 18: 361-365.

KAISER, E., ed. 1926. Die Diamantenwiiste Siidwestafrikas 1. Berlin: Dietrich Reimer (Ernst
Vohsen).

KENNEDY, W. J. & KLINGER, H. C. 1971. A major intra-cretaceous unconformity in eastern
South Africa. J. geol. Soc. Lond. 127: 183-186.

KLINGER, H. C. & KENNEDY, W. J. 1977. Upper Cretaceous ammonites from a borehole near
Richards Bay, South Africa. Ann. S. Afr. Mus. 72: 69-107.

MERENSKY, H. 1910. The diamond deposits of Lideritz-land, German South West Africa.
Trans. geol. Soc. S. Afr. 12: 13-23.

STENZEL, H. B. 1971. In: Moore, R. C., ed. Treatise on Invertebrate Paleontology. Part N.
Mollusca 6. Bivalvia. Geological Society of America & University of Kansas Press.

WEISSERMEL, W. 1926. Neues tiber Tabulate, Hydrozoen und einer Hexakoralle aus dem
Tertiar der Bogenfelser Diamantfelder. Jn: Kaiser, E., ed. Die Diamantenwiiste Siid-
westafrikas 2. Dietrich Reimer (Ernst Vohsen).

WriGuHT, C. W. 1957. In: Moore, R. C., ed. Treatise on invertebrate Paleontology. Part L.
Mollusca 4. Cephalopoda. Geological Society of America & University of Kansas Press.

ZIEGLER, W. H. 1969. Cenozoic Geology and Morphology of the coast of the Western Cape
Province and southern Namib desert, South Africa and South West Africa. Unpublished
Report, ESSO Exploration South Africa Incorporated.

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

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.
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An author’s name when cited must follow the name of the taxon without intervening
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species or subspecies is transferred from its original genus. The name of a subsequent user of
a scientific name must be separated from the scientific name by a colon.

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

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

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

Note punctuation in the above example:
comma separates author’s name and year
semicolon separates more than one reference by the same author
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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.

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by initials or full names
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Punctuation should be loose, omitting all not strictly necessary

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Specific name must not stand alone, but be preceded by the generic name or its abbreviation
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Name of new genus or species is not to be included in the title: it should be included in the
abstract,.counter to Recommendation 23 of the Code, to meet the requirements of
Biological Abstracts.

HERBERT CHRISTIAN KLINGER

CRETACEOUS DEPOSITS NEAR BOGENFELS,
SOUTH WEST AFRICA

/OLUME 73 PART 4 JUNE 1977 : ISSN 0303-2515

( OF THE SOUTH AFRICAN |
MUSEUM

CAPE TOWN

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

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

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Examples (note capitalization and punctuation)

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

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

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

Koun, A. J. 1900a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309-320.

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

THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270.

(continued inside back cover)

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

Volume . 73 ~~ ~+# Band
June 1977 Junie
Part 4 Deel

DEEP-SEA AMPHIPODS FROM WEST OF
tare POINT, sOUTH AFRICA

By

CHARLES L. GRIFFITHS

Cape Town Kaapstad

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DEEP-SEA AMPHIPODS FROM WEST OF CAPE POINT,
SOUTH AFRICA

By
CHARLES L. GRIFFITHS

Institute of Oceanography, University of Cape Town

(With 6 figures)

[MS. accepted 26 January 1977]

ABSTRACT

Two new species, Evonyx scutatus sp. nov. and Epimeria concordia sp. nov., are described
from this small collection, while of the remaining five species four are recorded for only the
second time and are new to southern African waters.

CONTENTS
PAGE
Introduction . F ; : s 93
Description of material . a” OF
Acknowledgements. : . 104
References . ; : ; . 2104
INTRODUCTION

Although the amphipods of shallower waters around South Africa are
relatively well documented, very few records exist from depths exceeding 1 000 m.
The first significant series of such samples was that collected by the S.S. Pieter
Faure between 1879 and 1907, from the vicinity of the Cape of Good Hope, and
described by K. H. Barnard (1916, 1925). Since that time only occasional, widely
scattered samples of deep-sea amphipods have been reported from the region,
notable amongst these being the collections of the Galathea expedition (J. L.
Barnard 1961) and the R.V. Vema cruises (J. L. Barnard 19625).

The present material is derived from twelve samples collected by Dr F. H.
Talbot of the South African Museum on board the R.S. Africana IJ, during
August and December 1959. An Agassiz trawl of approximately 1 cm mesh was
used and the area sampled was adjacent to, but somewhat deeper than, that
explored by the S.S. Pieter Faure. Due to the large size of mesh used only a few
relatively large amphipods were recovered, but these are of considerable interest
as they include two species new to science and four others new to the region.
All the specimens have been deposited in the South African Museum, Cape
Town.

93

Ann. S. Afr. Mus. 73 (4), 1977: 93-104, 6 figs.

94 ANNALS OF THE SOUTH AFRICAN MUSEUM

DESCRIPTION OF MATERIAL
Suborder GAMMARIDEA
Family Dexaminidae
Lepechinella (?) sucia J. L. Barnard, 1961
Fig. 1
Lepechinella sucia J. L. Barnard, 1961: 99, fig. 69.

Records
SAM-A13653, 33°49'S 16°30’E, 2 700 m, 27 August 1959, 3 males, 8 females.

Remarks

The degree of intraspecific variation amongst members of this exclusively
abyssal genus is poorly understood, since many species are known from only a
few individuals. The present material shows a close affinity to the unique male
of L. sucia recorded from the Tasman Sea by J. L. Barnard (1961). The present
specimens show more general setation than the Tasmanian individual, but lack
its distinct lateral rows of setae on first and second. pleonal epimera. The
resemblance as regards other significant features, such as shapes of the coxae and
pereonal teeth, is, however, so close that separation of the two forms seems
unjustified, at least until intraspecific stability of setation patterns is better:
understood. To facilitate comparison figures of a typical South African male are
provided.

Distribution
Tasman Sea, (?) South Africa.

Family Ischyroceridae
Bathyphotis tridentata Stephensen, 1944
Fig. 2
Bathyphotis tridentata Stephensen, 1944: 26, figs 17, 18.

Records
SAM A-13656, 34°49’S 16°30’E, 2 700 m, 27 August 1959, a single male.

Remarks

There can be no doubt that the present specimen is synonymous with that
described by Stephensen, since there are only minor quantitative differences in
the characteristic forms of gnathopod 2, pereiopod 3 and pleon segment 4.

There are, however, some points of difference which are a considerable
taxonomic significance. Stephensen noted a paucity of spines on maxilla | of his

DEEP-SEA AMPHIPODS FROM WEST OF CAPE POINT, SOUTH AFRICA 95

.
=

Fig. 1. Lepechinella (?) sucia J. L. Barnard, 1961
Male, 13 mm. A. Lateral aspect. B. Accessory flagellum. C. Mandibular palp. D. Coxa 1. E. Uropod 3. F. Telson.

‘uospaL “A ‘¢ podoig ‘qd ‘[ eyrxey ‘OD ‘s[qipueyy ‘gq ‘jodse [e10j}e 7] “Vv
“ph6l “ussusyda}s oyojuapiay sjoyddyjog ‘7 ‘S1.q

ANNALS OF THE SOUTH AFRICAN MUSEUM

96

DEEP-SEA AMPHIPODS FROM WEST OF CAPE POINT, SOUTH AFRICA 97

specimen (4 on outer plate, 3 on palp), a feature which he considered generically
significant. Careful dissection of the present male shows a more usual figure of
ten spines on the outer plate and five on the palp (some of the spines shown by
Stephensen were broken, indicating that the maxilla may have suffered some
damage, hence his low spine count).

Stephensen placed his material close to Eurystheus (= Gammaropsis), but
the species has subsequently been removed to Ischyroceridae (J. L. Barnard
1962a) and the structure of uropod 3 has since come to be regarded as ‘. . . with
elongate peduncle, rami short .. . outer ramus hooked apically’ (J. L. Barnard
1973). In his original description Stephensen makes no mention of a hooked
outer ramus, although such a feature could possibly be inferred from his figure 18.
Rather, he described the rami as ‘... narrow, acute, equal in length and
breadth’ —a description consistent with present observation, although the rami
show a minute immersed apical spine when viewed under high-power
magnification.

From the above evidence it would appear that the genus Bathyphotis is not
as discreet, nor as typically ‘ischyrocerid’, as has come to be supposed, although
the excavate coxa 4 remains unique amongst both Corophiidae and Ischyroceri-
dae. The placement of the genus remains open to question and a decision should
perhaps await a definitive revision of the entire group. In particular, the impor-
tance of shortening of rami of uropod 3 and the significance of this factor relative
to uncination of the outer ramus await clarification. An extensive discussion of
the utilization of these and other features is given by J. L. Barnard (1962a),
whose Eurystheus ventosa, for example, is closely allied to Bathyphotis (especially
considering the revised description of maxilla 1 herein). E. ventosa has recently
been made the type of a new genus Ventojassa J. L. Barnard, 1970, and trans-
ferred to Ischyroceridae. However, both this genus, Bathyphotis and Microjassa
have third uropods tending to merge with the condition found in Corophiidae.

Family Lysianassidae
Eurythenes gryllus (Lichtenstein, 1822)
Eurythenes gryllus: J. L. Barnard, 1961: 35, figs 5-7.

Records

SAM-A13653, 34°42’S 16°54’E, 3 200 m, 8 December 1959, a single female,
30 mm.

Remarks

Closely resembles the female figured by J. L. Barnard (1961). May be dis-
tinguished from E. obesus by the larger article 2 of pereiopods 3-5 and shorter
dactyl of the same limbs.

Distribution
Atlantic and North Pacific.

98 ANNALS OF THE SOUTH AFRICAN MUSEUM

Euonyx scutatus sp. nov.
Fig. 3
Description (of female, 16 mm)

Head shorter than pereon segment 1, ocular lobes acute, eyes absent;
flagellum of antenna | of one long and eighteen short articles, accessory flagellum
of nine articles; antenna 2 half as long again as 1, flagellum of thirty-seven
articles; epistome not projecting anteriorly, only slightly sinuous; mandibular
molar a large plate with raised margins, incisor simple, palp 3-articulate; palp
of maxilla | bi-articulate, outer plate with ten strong serrate spines, inner plate
bearing three plumose setae; outer plate of maxilliped excavate medio-distally.

Pereon dorsally carinate, segments 5—7 with progressively more pronounced
posterior teeth; coxa | triangular, gnathopod | chelate, articles 5 and 6 subequal
in length; palm of gnathopod 2 straight, transverse; coxa 5 developed into a
conical hump, other coxae normal; article 4 of pereiopods 3-5 strongly expanded
and distally produced.

Pleon segments 1-3 carinate, bearing postero-dorsal teeth, pleon segment 4
with a conical posterior hump; first pleonal epimeron smoothly rounded, 2 and
3 more quadrate; rami of uropod 1 lanceolate, 2 with outer ramus 80 per cent
length of inner; uropod 3 with inner ramus apically truncated, outer ramus
lanceolate, slightly the longer; telson 80 per cent cleft.

Holotype
' SAM-A13652, female, 16 mm; unique.

Type locality
34°37’'S 17°03’E, 2 900 m, 8 December 1959.

Relationships

Although some other species of Euonyx (E. chelatus, E. conicurus) show a
dorsal tooth on pleon segment 4, no others have the extensive series of pereonal
and pleonal teeth found in E. scutatus sp. nov. This species is also distinguished
by its triangular coxa | and shield-like coxa 5 as well as by details of the structure
of gnathopod 2.

Lepidepecreoides nubifer J. L. Barnard, 1971
Fig. 4
Lepidepecreoides nubifer J. L. Barnard, 1971: 41, figs 26-27.
Records

SAM-A10547, 34°36’S 17°00’E, 2 740 m, 10 December 1959.

Remarks

This is only the second record of this species and the first of a female. The
specimen closely resembles the smaller male described by J. L. Barnard (1971),

DEEP-SEA AMPHIPODS FROM WEST OF CAPE POINT, SOUTH AFRICA 99

Fig. 3. Euonyx scutatus sp. nov.

Female, 16 mm. A. Lateral aspect with epistome enlarged. B. Mandible. C. Maxilla 1.
D. Maxilliped. E-F. Gnathopods 1, 2. G. Uropod 3. H. Telson.

100 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 4. Lepidepecreoides nubifer J. L. Barnard, 1971.

Female, 17 mm. A. Head. B. Antenna 1. C. Mandible. D. Maxilla 1. E. Mazxilliped.
F-G. Gnathopods 1, 2. H. Pereiopod 3. I. Dorsal profile of pereon segment 7 and pleon.
J. Uropod 3. K. Telson.

DEEP-SEA AMPHIPODS FROM WEST OF CAPE POINT, SOUTH AFRICA 101

except for its more tubercular dorsal profile and the longer processes of article
2 of pereiopod 3. Variability of dorsal profile may be a factor of size or sex, but
at present cannot be regarded as taxonomically significant. Variations in the
lengths of the processes of article 2 of pereiopod 3 are documented for the type
species of the genus, L. xenopus K. H. Barnard, although in this case the processes
were longer in smaller individuals.

Distribution :
Oregon 2 860 m, South Africa 2 740 m.

Family Paramphithoidae
Epimeria concordia sp. nov.

Fig. 5
Description (of female, 30 mm)

Rostrum elongate, reaching tip of article 3 of antenna 1, head with distinct
ocular bulge; pereon segments dorsally smooth, except for small posterior hump
on segment 7; article 6 of gnathopod | widening slightly, posterior margin with
six equally spaced slender spines, palm straight, almost transverse, defined by a
single slender spine, dactyl serrate; gnathopod 2 similar to | but palm defined
by two spines; coxa 4 with long anteroventrally curved cusp and oblique ridge;
coxa 5 strongly produced laterally, forming a large triangular ‘wing’ when
viewed from above; coxa 6 with a much smaller lateral process; coxa 7 coniform;
article 2 of pereiopod 5 proximally dilated.

Pleon segments 1-3 each with a large upright mediodorsal tooth, segment 4
with a quadrate notch basally and small erect tooth posteriorly; pleonal epimera
with accessory tooth on posterior margin, postero-distally acute, second epimeron
with an oblique ridge; uropods lanceolate, projecting equally; telson distinctly
emarginate apically.

Holotype
SAM-A13651, female, 30 mm, unique.

Type locality
34°36’S 17°00’E, depth 2 740 m, 10 December 1959.

Relationships

This species is remarkable for the enlarged lateral projection of coxa 5,
wnich readily distinguishes it from other species occurring in southern Africa.
In other respects it is similar to the closely related group comprising E. glaucosa
J. L. Barnard, E. cora J. L. Barnard, E. subcarinata Nagata and E. pacifica
Gurjanova. None of these forms, however, exhibit the combination of elongate
rostrum, large pleonal teeth and accessory teeth on the pleonal epimera, as found
in E. concordia sp. nov.

ANNALS OF THE SOUTH AFRICAN MUSEUM

102

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“AOU “ds vIpsoquod Didaudy *°¢ “S1q

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103

DEEP-SEA AMPHIPODS FROM WEST OF CAPE POINT, SOUTH AFRICA

104 ANNALS OF THE SOUTH AFRICAN MUSEUM

Suborder CAPRELLIDEA
Family Caprellidae
Caprella ungulina Mayer, 1903
Fig. 6
Caprella ungulina Mayer, 1903: 127, pl. 5 (fig. 36), pl. 8 (figs 30-31).

Records

33°50’S 17°21’E, 1 100 m, 25 August 1959, numerous males and juveniles
attached to appendages of the giant stone crab Neolithoides asperrimus K. H.
Barnard.

Remarks

One of the few caprellids occurring below 1 000 m, readily identified by the
strong spines on the palms of pereiopods 5—7. This species has not been recorded
since its original description in 1903, hence Mayer’s somewhat rudimentary
figures are supplemented here.

Distribution

West coast of North and South America; South Africa (Mayer does not
state whether his material was found in association with stone crabs, as was the
case with the present specimens).

ACKNOWLEDGEMENTS

The author is indebted to the South African Museum for the loan of material
upon which this work was based and for the provision of library facilities.
Financial support was provided by the South African Council for Scientific and
Industrial Research. The manuscript was kindly typed by Mrs S. Hardman.

REFERENCES

BARNARD, J. L. 1961. Gammaridean Amphipoda from depths of 400 to 6 000 meters. Galathea
Rep. 5: 23-128.

BARNARD, J. L. 1962a. Benthic marine Amphipoda of southern California: Families Aoridae,
Photidae, Ischyroceridae, Corophiidae, Podoceridae. Pacif. Nat. 3: 1-72.

BARNARD, J. L. 19625. South Atlantic abyssal amphipods collected by R.V. Vema. In: Abyssal
Crustacea. Vema Res. Ser. 1: 1-78.

BARNARD, J. L. 1970. Sublittoral Gammaridea (Amphipoda) of the Hawaiian Islands.
Smithson. Contr. Zool. 34: 1-286.

BARNARD, J. L. 1971. Gammaridean Amphipoda from a Deep-Sea Transect off Oregon.
Smithson. Contr. Zool. 61: 1-86.

BARNARD, J. L. 1973. Revision of Corophiidae and related families (Amphipoda). Smithson.
Contr. Zool. 151: 1-27.

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. 1925. Contributions to the crustacean fauna of South Africa. 8. Further
additions to the list of Amphipoda. Ann. S. Afr. Mus. 20: 319-380.

Mayer, P. 1903. Die Caprellidae der Siboga Expedition. Siboga Exped. 34: 1-158.

STEPHENSEN, K. 1944. Crustacea Malacostraca, VIII: (Amphipoda IV). Danish Ingolf-Exped.
3: 1-51.

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

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

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

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

Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15A
Nucula (Leda) eng Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: re pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (figs 8a—b).

Nucula largillierti Philippi, 1861:
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 peeiaee of prefixed surnames in all fee 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

(c) Scientific names, but not their vernacular derivatives
e.g. Therocephalia, but therocephalian

Punctuation should be loose, omitting all not strictly necessary

Reference to the author should be expressed in the third person

Roman numerals should be converted to arabic, except when forming part of the title of a
book or article, such as
‘Revision of the Crustacea. Part VIII. The Amphipoda.’

Specific name must not stand alone, but be preceded by the generic name or its abbreviation
to initial capital letter, provided the same generic name is used consecutively.

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

DEEP-SEA AMPHIPODS FROM
WEST OF CAPE POINT,
SOUTH AFRICA

—_ = > ~~ Ye

VOLUME 73 PART 5 — UNE 1977 ISSN 0303-2515

.
|

| ANN ALS
!

OF THE SOUTH AFRICAN —
ee 2 irs

07.8 cd ee RODY4n

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

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

‘Smith (1969) describes .. .’

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Examples (note capitalization and punctuation)

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

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

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

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

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

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 73 Band
June 1977 Junie
Part 5 Deel

THE SOUTH AFRICAN MUSEUM’S
MEIRING NAUDE CRUISES
PARI ”3
HYDROIDA

By

N. A. H. MILLARD

Cape Town Kaapstad

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THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES
PART 3

HYDROIDA

By
N. A. H. MILLARD

South African Museum, Cape Town

(With 10 figures and 1 table)

[MS. accepted 2 February 1977]

ABSTRACT

The paper describes a collection of hydroids from off the east coast of South Africa, most
of them from depths of over 500 m. In all there are 34 species, of which 8 are new records for
the country. Among the latter are 1 new genus— Uniscyphus—and 3 new species — Cladocarpus
natalensis, Uniscyphus fragilis and Zygophylax inconstans.

CONTENTS

PAGE
Introduction . , : Bal
IMistof species. . : = 106
Systematic account. . 108
Discussion . , , arn eet)
Acknowledgements. - = £30
References. . , : 4 7130

INTRODUCTION

The hydroids described in this paper were dredged off the coast of Natal
during two cruises of the R.V. Meiring Naude undertaken by the Marine Biology
Department of the South African Museum during May 1975 and May 1976.
The Station Data for these two cruises were given in an earlier number of this
journal (Louw 1977) and will not be repeated here.

The depths of the samples varied from 40 to 1 200 m; most of them were
over 500 m, so that these samples were from well over the edge of the continental
Shelf at 200-400 m, an area which is poorly known and where little collecting
has been done. |

105

Ann. S. Afr. Mus. 73 (5), 1977: 105-131, 10 figs, 1 table.

106 ANNALS OF THE SOUTH AERICAN MUSEUM

LIST OF SPECIES

* New records for South Africa
+ Discussed further in the following pages

Station South African
number Museum number
Family Bougainvilliidae
*+Garveia crassa (Stechow, 1923) . A ; : : -” SM 58 SAM-H2875
SM 66 SAM-H2876

Family Campanulinidae

*tEgmundella ?superba Stechow, 1921 . : ; } . SM 38 SAM-H1967
Medeeria rotunda (Quoy & Gaimard, 1827) . ‘ . SM 23 SAM-H1963
SM 86 SAM-H2808

tOpercularellasp.- 32 Fe . SM 38 SAM-H1969
SM 103 SAM-H2854

*+t Stegolaria geniculata (Allman, 1888) . : r . SM 38 SAM-H1968

Family Haleciidae

Halecium tenellum Hincks, 1861... a . SM 92 SAM-H2812
SM 103 SAM-H2858

Family Lafoeidae

Acryptolaria conferta (Allman, 1877) . : : A - -OSMet5 SAM-H1952
SM 23 —
SM 86 SAM-—H2807
SM 94 SAM-H2847
SM 103 SAM-—H2852
SM 107 SAM-H2856
Acryptolaria rectangularis (Jarvis, 1922) . i : , SMS —
SM 16 SAM-H1953
SM 23 SAM-H1957
SM 86 SAM-H2805
SM 94 SAM-H2848

SM 107 —
Filellum serratum (Clarke, 1879)... Page « 2.) SMES SAM-H1978

SM 43 —
+Lafoea dumosa (Fleming, 1820) . : ; : é . SM 23 SAM-H1958

SM 38 SAM-H1965
SM 67 SAM-H2801
SM 86 SAM-H2871
SM 92 SAM-H2811
SM 99 SAM-H2849
SM 103 SAM-H2851

Zygophylax africana Stechow, 1923 . ; ; 3 - SMEz3 SAM-H1973
SM 86 SAM-H2806

SM 92 —
*tZygophylax brownei Billard, 1924 ; : : : . SM 23 SAM-H1974
SM 86 SAM-H2874
*tZygophylax inconstans sp. Nov. . : : : : . SM 23 SAM-H1975
SM 43 SAM-H1977
Zygophylax sibogae Billard, 1918 : : : : = oSMy238 SAM-H1956

SM 31 SAM-H1964
SM 52 SAM-H2800
SM 86 SAM-—H2804
SM 92 SAM-H2846

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 107

Station South African
number Museum number

Family Campanulariidae

Campanularia hincksu Alder; 1856- . .. . «| .~¢ SM 23 SAM-H1961
Wipnoacravieri (Billard, 1904)-".. » . .« .. « ~SM.20 SAM-H2859
SM 79 SAM-H2913
SM 80 —
SM 95 SAM-H2926
Chyna paniensis (Vanhofien, 1910)~...°. 2 . .. sl | 6SM 16 SAM-H1954

Family Sertulariidae

Sertularella leiocarpa (Allman, 1888) . ; : : 2 SIMETS SAM-H1951
SM 38 SAM-H1979
SM 86 SAM-—H2809

SM 103 —
*+ Symplectoscyphus amphoriferus (Allman, 1877) ; , SM 38 SAM-H1981
SM 86 SAM-H2810
Symplectoscyphus arboriformis (Markt.-Turn., 1890) . SM 43 SAM-H1980
Symplectoscyphus paulensis Stechow, 1923 ; : . SM 100 SAM-H2850
SM 107 SAM-H2855
*+ Uniscyphus fragilis g. nov., Sp. nov. . ; ; F . SM 43 SAM-H1982

Family Plumulariidae

+ Antennella quadriaurita Ritchie, 1909 . ; : : a (SNES SAM-H1962
SM 38 SAM-H1983

SM 43 SAM-H1966

SM 52 SAM-H2877

SM 83 SAM-H2878

SM 86 SAM-H2879

SM 103 SAM-—H2880

Cladocarpus distomus Clarke, 1907. ie teat Pek SI Oe SAM-H2866
SM 86 SAM-—H2865

SM 94 SAM-H2864

+ Cladocarpus dofleini (Stechow, 1911) . : , : . ' SM 86 SAM-—H2860
Cladocarpus millardae Vervoort, 1966 : : 4 . SM 86 SAM-—H2803
*+ Cladocarpus natalensis sp. nov. . ; : . : am lv 23 SAM-H1972
SM 86 SAM-H2861

+ Cladocarpus sinuosus Vervoort, 1966 . : : P SME 23 SAM-H1970

SM 86 SAM-H2863
SM 103 SAM-H2862

Halopteris glutinosa (Lamouroux, 1816) . 5 : . SM 86 SAM-H2869
Halopteris polymorpha (Billard, 1913) : : : -- (SM.23 SAM-H1959

SM 86 SAM-—H2870
Kirchenpaueria triangulata (Totton, 1930) . : : 2 SIME D2 SAM-H2799

SM 71 SAM-H2802
SM 86 SAM-—H2863

Nemertesia ramosa Lamouroux, 1816. R : : eS NL2S SAM-H1960
SM 103 SAM-H2853
Plumularia mossambica Millard, 1975. : . SM 86 SAM-H2867

Thecocarpus flexuosus flexuosus (Lamouroux, 18 1 6) S20 mA. AES SAM-H2857

108 ANNALS OF THE SOUTH AFRICAN MUSEUM

SYSTEMATIC ACCOUNT
Family Bougainvilliidae
Garveia crassa (Stechow, 1923)
Fig. 1A—-C
Bimeria crassa Stechow, 19236: 103. Stechow, 1925: 414, fig. 4.

Description

Colonies growing luxuriantly over the tubes of the polychaet worm, Loimia
sp., and almost completely obscuring them. Stem strongly fascicled and about
1 mm thick at base, branching freely and quite irregularly, with many of the
larger branches reuniting to form a complex meshwork, reaching a maximum
height of 40 mm. The thicker parts of the stem commonly containing large spaces
between the bundles of tubes, these spaces packed with mud and Foraminifera.
Terminal branches unfascicled or lightly fascicled, bearing hydranth pedicels
which are usually narrowest at origin and wider distally. Perisarc roughly corru-
gated and folded throughout, continued over the bases of the hydrothecae to
form pseudohydrothecae, but leaving the tentacles free. Hydranth with about
nine tentacles.

Gonophores shortly stalked, borne rather sparsely on stem and hydranth
pedicels, oval, covered with a thin envelope of perisarc, the largest 0,33 mm long
and 0,17 mm wide, in the form of fixed sporosacs, though not mature enough to
determine sex.

Nematocysts of at least two kinds:

(i) Desmonemes; 3,6 x 2,4 — 4,2 x 3,2 um.
(ii) Microbasic euryteles; 5,4 x 2,7 — 6,0 x 3,3 um.

Remarks

This species was described by Stechow from 741 m off the coast of Somali-
land. It is a new record for South Africa, where it occurs in roughly the same
depth (720-850 m). Stechow did not mention the substratum and he presumably
had only a detached stem available to him. It would be interesting to know
whether the species always occurs on polychaet tubes.

In accordance with modern practice the genus Bimeria is restricted to those
species where the bases of the tentacles are clothed with tubes of perisarc. This
species must thus be transferred to the genus Garveia Wright, 1859, in which the
pseudohydrotheca terminates below the tentacle bases.

Family Campanulinidae
Egmundella superba Stechow, 1921
Fig. 1D-G

Egmundella superba Stechow, 1921: 226. Stechow, 1923a: 126, fig. R. Vervoort, 1966: 110,
fig. 10.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 109

Fig. 1. Garveia crassa (Stechow). A. Part of a fascicled stem bearing hydranths. B. Four
hydranths. C. Gonophore.

Egmundella superba Stechow. D. Hydrotheca and pedicel. E. Hydrotheca. F. Hydrorhiza
with nematophores and origin of pedicel. G. Nematophore.

Scale in mm/10.

110 ANNALS OF THE SOUTH AFRICAN MUSEUM

Description |

A colony of seven solitary hydrothecae and many sterile pedicels arising
from a hydrorhiza creeping on sponge spicules.

Pedicel long, with two to four distinct annulations close to base, otherwise
smooth except for occasional regeneration nodes. Hydrotheca not distinctly
demarcated from pedicel, top-shaped and widest at margin, depth two and a half
to three and a third times maximum diameter, without diaphragm or annular
thickening. Operculum deep, with about eight irregularly folded valves, which
are not demarcated from thecal margin. Hydranths completely absent.

Nematothecae abundant, arising from hydrorhiza, of irregular length,
tubular but slightly swollen at distal end.

Measurements (mm)

Total length . : : . 3,21-6,20
Pedicel, diameter . . 0,08-0,09
Hydrotheca, approximate depth . ., .0;80-1,2e
maximum diameter . . 0,32-0,38
Nematotheca, length . . ~0.05-0,42
maximum diameter . : ; e002
Remarks

Of all the known species of Egmundella this material most closely resembles |
E. superba. The hydrotheca is slightly broader than that of the holotype
(redescribed by Vervoort 1966), but is otherwise similar, and the pedicel is
similarly annulated at the base only. The nematothecae differ in their variable
length and in their profuse growth which may completely cover the hydrorhiza
like a bristly mat.

The type locality and only previous record of E. superba is St Thomas, West
Indies (depth not given). It is a new record from South Africa.

Opercularella sp.
Fig. 2

Description

Several branched stems reaching 22 mm in height and growing on sponge
spicules. Stem branching sympodially, fascicled, giving off alternate hydrothecae
from an axial tube, straight or very slightly geniculate, the two rows of hydro-
thecae in one plane. Branches rather irregular, occasionally subaiternate and
arising below every third and fourth hydrotheca, similar to stem but unfascicled
or lightly fascicled only, the two rows more or less in one plane. Stem and
branches either unsegmented or with a faint and very oblique node immediately
above the origin of each hydrotheca. Some solitary hydrothecae arising separately
from hydrorhiza.

Hydrotheca pedicellate. Pedicel shorter than hydrotheca and not clearly
demarcated from it (boundary only recognizable by attachment of hydranth base

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 111

Fig. 2. Opercularella sp. Various hydrothecae and opercula, a solitary hydrotheca at bottom
right. Scale in mm/10.

ti2 ANNALS OF THE SOUTH AFRICAN MUSEUM

to hydrotheca), forming a very acute angle with stem and often pressed against
it at base, sometimes with one or two indistinct corrugations. Hydrotheca
tubular, asymmetrical and curved away from stem, with adcauline wall more
convex than abcauline. Solitary hydrothecae less curved. No diaphragm visible.
Operculum of four or five fragile converging segments which are not sharply
demarcated from thecal margin.

Gonothecae absent.

Measurements (mm)

Pedicel, length : : . 0,07-0,26

Hydrotheca, length abcauline . . 0,35-0,54

maximum diameter . : i . 0,11-0,16
Remarks

This material is unlike any other described species. It is perhaps closest to
‘?Opercularella spec. no. 2’ of Vervoort (1966: 108), but differs from it in the
curved hydrothecae and the smaller number of opercular segments. The oper-
culum is very delicate and crumples easily; it can only be seen clearly by slicing
off the top of the hydrotheca and viewing from above. Since the material is not
very well preserved and is infertile the writer prefers not to describe it as a new
species.

Stegolaria geniculata (Allman, 1888)
Fig. 3

Cryptolaria geniculata Allman, 1888: 41, pl. 20 (figs 1, la, 15).

?Cryptolaria operculata Nutting, 1905: 947, pl. 3 (fig. 4), pl. 10 (figs 12-14). Ritchie, 1910: 9.
Stegopoma operculatum: Billard, 1941: 16, fig. 1.

Stegolaria geniculata: Vervoort, 1946: 299, figs 2-3. Edwards, 1973: 593.

Stegolaria operculata: Edwards, 1973: 594.

Description

Several branching fan-shaped stems reaching a maximum height of 40 mm
and several smaller ones, growing on a gorgonian skeleton, sponge spicules and
a worm tube.

Stem strongly fascicled, giving rise to alternate hydrothecae and roughly
alternate or subalternate branches from an axial tube. Branches similar to stem,
fascicled almost to the end. The two rows of branches and hydrothecae in one
plane.

Hydrotheca tubular, curved smoothly outwards, adnate to stem or branch
for half to two-thirds height and in the thicker parts of the stem with the adnate
part immersed among the peripheral tubes. Axillary hydrotheca with abcauline
wali adnate to branch. Base of hydrotheca without diaphragm or perisarcal
thickening, but usually demarcated by an indentation of the abcauline wall. Distal
part of hydrotheca very delicate and usually damaged. Operculum of the
Stegopoma type, consisting of two pleated valves seated in the embayments
between two gable-like processes of the margin.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 113

iol Bene Sees een (eer Set hee |

Fig. 3. Stegolaria geniculata (Allman). A. Part of stem from distal end. B-E. Parts of stem
with gonothecae. F. Distal ends of two hydrothecae to show opercula. Scale in mm/10.

114 ANNALS OF THE SOUTH AFRICAN MUSEUM

Gonotheca sac-shaped, completely adnate to branch. Aperture facing away
from branch and sometimes raised slightly above it, subterminal, circular, with
an operculum of fragile converging segments. Containing up to five gonophores
(probably male) one above the other.

Measurements (mm)

Hydrotheca, length abcauline (approx.) . . 0,96—-1,62
length adcauline, adnate part . . 0,72-0,96
length adcauline, free part. . . 0,36-0,93
diameter at mouth . . 0,19-0,40

Gonotheca, length ' . ~*~ 1112502
maximum diameter . : . 0,26-0,46
diameter of aperture. ‘ . O10

Remarks

The author can see little to distinguish Stegolaria operculata from S. genicu-
lata other than the markedly geniculate stem of the latter. In the present material
the degree of geniculation is variable and in general intermediate between the
two, the tips of some branches being almost as geniculate as those illustrated by
Allman and Vervoort for S. geniculata and other branches being completely
straight as in S. operculata. She has therefore united the two species. The gono-
thecae are exactly like those described by Vervoort (1946).

The genus Stegolaria has been retained in view of the peculiar gonothecae;
since these contain fixed sporosacs the genus cannot be united with Modeeria.
The status of Stegopoma awaits further information on reproduction.

Distribution

Fiji (type locality), Hawaii, East Indies (Celebes and Kei Island), Malay
Archipelago. A deep-water species ranging from 253 to 910 m. A new record
from South Africa.

Family Lafoeidae

Lafoea dumosa (Fleming, 1820)
Remarks

The opinion of Cornelius (1975) is accepted that Lafoea fruticosa (M. Sars)
is a synonym for L. dumosa (Fleming). This cosmopolitan species is well known
from deeper waters of the South African coast.

Zygophylax brownei Billard, 1924
Fig. 4

Lafoea pinnata: Browne, 1907: 25.
Zygophylax pinnata: Billard, 1923: 14, fig. 1A.
Zygophylax brownezi Billard, 1924: 64. Leloup, 1940: 11, pl. 1 (fig. 7). Patriti, 1970: 28, fig. 30.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 115

Description

Two colonies with stems reaching a maximum height of 61 mm, stiff,
fascicled, bearing alternate hydrothecae and subalternate branches in one plane.
Branches usually arising immediately below every third and fourth hydrotheca,
bearing alternate hydrothecae, sometimes rebranching in a similar manner to the
stem. Where the branching follows this regular subalternate scheme, both hydro-
thecae and branches arise from a single axial tube, but this arrangement is some-
times obscured by extra branches and hydrothecae from peripheral tubes.
These possibly represent auto-epizootic colonies but it is no longer possible to
distinguish the hydrorhizae.

Hydrotheca and pedicel together forming a bilaterally symmetrical tubular
figure which is more convex on the adcauline side. Pedicel with no node separating
it from the apophysis but demarcated by an indentation on adcauline side,
separated internally from hydrotheca by a well-developed diaphragm. Hydro-
theca everted at margin.

Nematothecae scarce, occurring singly on stem apophyses, usually only one
or two to a stem; tubular, with everted margin, variable in length.

Gonothecae (present in Station SM 23) quite separate from one another, not
conjoined in a coppinia but clustered thickly round certain parts of the stem;
deep-oval, not compressed, with two apertures (rarely three), each on the end of
a recurved tubular neck; containing planula larvae; with no special accumula-
tions of nematothecae.

Measurements (mm) Station SM 23 Station SM 86
Pedicel, length adcauline . . 0,06—0,10 0,05—0,12
Hydrotheca, length adcauline . . 0,34-0,41 0,31-0,47

diameter at margin . 2) FO, 160.99 0,14-0,18
Gonotheca, length . . 0,08—-1,32 _
maximum diameter (below necks) . 0,31-0,38

Remarks

Z. brownei is closely related to Z. biarmata Billard, 1905, and is in fact
included with it by Broch (1918). It is, however, retained as a separate species
by Leloup (1940) on the basis of the sparser distribution of nematothecae. This
is the first record of Z. brownei from the southern hemisphere and the first
description of gonothecae; the latter are similar in general shape to those of
Z. biarmata (as described by Saemundsson (1912) under the name of Lictorella
levinseni) but are proportionally longer and more slender, and are without the
numerous nematothecae described by Broch (1918).

Both these species have a north Atlantic distribution. Records of Z. biarmata
from other parts of the world (Jaderholm 1919, from Japan; Jarvis 1922, from
east Africa) were all sterile and thus still need confirmation.

116 ANNALS OF THE SOUTH AFRICAN MUSEUM

Distribution

Eastern North Atlantic from France to Morocco, 20-752 m. Type locality:
Bay of Biscay. A new record from South Africa.

B
A
B

C AN key
| (GP

Fig. 4. Zygophylax brownei Billard. A. Part of stem. B. Hydrothecae. C. Gonothecae, that
on extreme left in side view, the two on extreme right each with three openings. Scale in mm/10.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES dole

ZLygophylax inconstans sp. nov.
Fig. 5
Material

Holotype: SAM-—H1975. Station SM 23: 27°44,4’S 32°42,8’E, 400-450 m.
Other material: SAM—H1977. Station SM 43: 28°45,5’S 32°24,5’E, 360-420 m.

Description of holotype

Colony growing on a polyzoan and consisting of slender, straggling stems
often reattaching to one another and to the polyzoan and producing a tangled
irregular growth one or two centimetres in height. Stem unfascicled or lightly
fascicled, unbranched or branching irregularly and in any plane, unsegmented,
geniculate, bearing alternate hydrothecae on short apophyses which arise at the
‘elbows’. Peripheral tubes arising at origins of branches.

Hydrotheca and pedicel not sharply demarcated externally, together forming
a deep-campanulate figure which may be radially symmetrical or slightly
bilaterally symmetrical with the adcauline surface more convex than the
abcauline. Margin of hydrotheca everted. Diaphragm distinct, usually oblique.

Nematothecae numerous, one to three on each apophysis (usually two), and
scattered on peripheral tubes of stem and on hydrorhiza; tubular, two-
chambered, with short basal chamber and long distal chamber, with everted
margin.

Coppiniae present surrounding thicker parts of stem, consisting of a mass of
conjoined gonothecae, but without modified hydrothecae or nematothecae.
Gonotheca saccular and of irregular shape, with a single aperture with a flared
margin on the summit of a short tubular neck.

Measurements (mm)

Pedicel, length adcauline : . 0,03-0,06
Hydrotheca, length adcauline . . 0,22-0,30
diameter at margin. . 0,10-0,12
Gonotheca, length . reaching . 5 a One
maximum diameter . reaching. 0,52
Remarks

The ramifications of these colonies are so intimately associated with the
polyzoan that it is almost impossible to separate the two. The hydrorhiza runs
along the upright stem of the host, separating from it and attaching to other
parts at intervals; it produces erect stems, solitary hydrothecae and numerous
nematothecae.

There are two species of Zygophylax which share certain characters with this
material, namely a closed coppinia, a gonotheca with a single terminal aperture,
small deep-campanulate hydrothecae, and two nematothecae to each hydrotheca-
bearing apophysis. These are Z. armata (Ritchie, 1907) and Z. profunda Quelch,

118 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 5. Zygophylax inconstans sp. nov. from the holotype, SAM—H1975. A. Part of stem.
B-C. Hydrothecae. D. Axillary hydrotheca, and origin of peripheral tubes. E. Coppinia.
Scale in mm/10.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 119

1885. Both of these have a much stiffer colony and a more heavily fascicled stem,
and in both the coppinia is provided with numerous long branching nemato-
thecae. In Z. armata the gonothecae are more regular and hexagonal in section.
The schizoholotype of Z. profunda from Cape Verde is a mounted slide (B.M.
no. 85.7.21.1) of infertile fragments in very poor condition. The hydrothecae are
shorter and more strongly curved than in the present material (Fig. 6A). The
fertile material from Madeira assigned to Z. profunda by Totton (1930) (slide
B.M. no. 19.8.15.2) has hydrothecae more similar in shape to the present material
though very slightly smaller (Fig. 6B—C). The coppinia is identical apart from
the presence of branching nematothecae. It is mainly this last character that thus
separates Z. profunda from Z. inconstans.

Family Sertulariidae
Symplectoscyphus amphoriferus (Allman, 1877)
Fig. 7A—D

Sertularella amphorifera Allman, 1877: 22, pl. 15 (figs 8-10). Nutting, 1904: 88, pl. 20 (figs 1-2).
Billard, 1906: 183.
Symplectoscyphus ?amphoriferus: Millard, 1967: 182, fig. 4E—-F.

Description

Several unfascicled stems reaching a maximum height of 21 mm, some of
them branching alternately. Stem and branches geniculate in distal parts.
Branches given off at a wide angle (over 80°) and with a dichotomous effect due
to a more strongly marked geniculation at the origin of each branch. Nodes
indistinct.

Hydrotheca deep and slender, adnate for about one-third adcauline length,
curved outwards, adcauline free part straight or slightly concave. Margin with
three teeth, one adcauline and two latero-abcauline. No internal teeth.

One gonotheca present, pear-shaped, with eleven raised transverse ridges
and a slender terminal neck.

Measurements (mm)

imemode lencth . . . «| -s OU6O-O98
Hydrotheca, length abcauline . ats te U6 "U.45
length adcauline, adnate part . . 0,20-0,23
length adcauline, free part. ~ | 10;33-0:40
diameter at margin . ; . 0,12-0,14
Gonotheca, length ; a lkeo
maximum diameter . oe O74
Remarks

This material is very similar to that recorded from the southern Indian Ocean
(Millard 1967). The presence of pseudodichotomous branches and a larger pro-

120 ANNALS OF THE SOUTH AFRICAN MUSEUM |

Fig. 6. Zygophylax profunda Quelch. A. Hydrothecae from the schizoholotype, BM 85.7.21.1.
B. Hydrothecae, and C, coppinia, from Totton’s material, BM 19.8.15.2. Scale in mm/10.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 1Dt

Fig. 7. Symplectoscyphus amphoriferus (Allman). A-B. Hydrothecae. C. Part of stem.
D. Gonotheca.

Uniscyphus fragilis sp. nov. from the holotype, SAM-—H1982. E. Hydrotheca and pedicel.
F. Hydrotheca. G. Distal end of hydrotheca with regenerated margins and operculum.

Scale in mm/10.

122 ANNALS OF THE SOUTH AFRICAN MUSEUM

portion of the hydrotheca adnate to the stem is closer to the type material and
supports the identification.

Distribution

Except for one record from the southern Indian Ocean, this species is known
only from the northern Atlantic from the West Indies to south-east of Iceland.
All the records are from deep water and range from 185 to 1 256 m. Type
locality: Double-headed Shot Key. A new record from South Africa.

Uniscyphus gen. nov.
Diagnosis
Colony stolonial, with pedicellate hydrothecae arising direct from a creeping
hydrorhiza. Hydrotheca cylindrical, with three marginal teeth and an operculum
of three valves seated in the bays between the teeth and meeting in the centre as
a pyramid.
Type species: Uniscyphus fragilis sp. nov.

Uniscyphus fragilis sp. nov.

Fig. 7E-G
Material

Holotype: SAM-H1982. Station SM 43: 28°45,5’S 32°24,5’E, 360-420 m. Five
hydrothecae, three mounted on a slide.

Description

Hydrorhiza creeping. Hydrotheca solitary, pedicellate. Pedicel long, at least
three times length of hydrotheca, slender, not annulated, arising at right angles
to hydrorhiza. Hydrotheca terminal, cylindrical, not annulated, with three well-
developed, equally spaced marginal teeth. No internal teeth. Operculum of three
valves. Gonothecae absent.

Measurements (mm)

Pedicel length : ; . 2,4-3,9

Hydrotheca, depth to tips of teeth . . 0,70-0,75
diameter at margin . . 0,25-0,29
diameten/@epiiy = 0) lene e554 Onc

Remarks

There is no existing genus to contain this delicate species. In its stolonial
form it resembles Calamphora, which, however, has four marginal teeth and an
annulated hvdrotheca. It seems to bear the same relationship to Parascyphus that
Calamphora bears to Sertularella. The hydranths are not well preserved, and,
although it is difficult to be certain, no abcauline blind pouch could be seen.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 123

Family Plumulariidae
Antennella quadriaurita Ritchie, 1909
Fig. 8

Antenella quadriaurita Ritchie, 1909: 92, fig. 9. Leloup, 1932: 162, pl. 16 (fig. 2).
Antennella quadriaurita: Stechow, 1919: 113. Millard, 1966: 492.

Antenella quadriaurita, forma africana Broch, 1914: 26.

Antenella africana: Stechow, 1925: 492, fig. 11.

Antennella africana: Millard, 1975: 331, fig. 107A—E.

Discussion

The seven samples of Antennella, of which only the last two were fertile,
have long, slender athecate internodes bearing two to four nematothecae each.
This suggested an affinity with A. quadriaurita rather than with the South African
species A. africana (both are species with two pairs of lateral nematothecae), and
since there was already a doubt as to the separate entity of these two species a
reappraisal of the South African material was undertaken. Also available for
comparison was a new sample from Tristan da Cunha (SAM-H1949), and
another from Nightingale Island (SAM-—H1991), both collected in 1971.

It has previously been claimed (Millard 1975) that the only character
separating these two species is the presence of, usually, one nematotheca on each
athecate internode in A. africana and two or three in A. quadriaurita. Counts of
nematothecae in the available samples (Table 1) show no geographical relation-
ship. For example, the sample from Tristan has one nematotheca on 82 per cent
of the internodes, as against Ritchie’s type of A. quadriaurita from Gough Island,
and material from the Vema Seamount (South Atlantic), where most internodes
have two or three nematothecae. On the west and south coasts of South Africa
most internodes have one nematotheca, yet in the sample from Port Elizabeth
all internodes have two nematothecae, and in the seven samples of the present
collection from Natal most have two or three.

There does, however, appear to be a relationship between the number of
nematothecae and the length of the internode. This is demonstrated in Figure 8,
whence it is apparent that, although there is much variation within a sample, in
general longer internodes have more nematothecae. This diagram also indicates
that the seven deep-water samples from Natal (Numbers 14-15, 17-21) are most
closely related to the type material from Gough Island (Number 16), from which
they are most distant geographically.

Within a single colony, the longer athecate internodes bearing several
nematothecae tend to occur near the base of a hydrocladium and shorter ones
with one nematotheca near the distal end.

From these considerations it is concluded that A. quadriaurita and A. africana
cannot be retained as separate species, and the latter is sunk in the former.

The distribution of the composite species is:

Central and South Atlantic Ocean, including Havanna (Stechow 1919), the

Tristan da Cunha group (Ritchie 1909, and this work), tropical west Africa

NUMBER OF NEMATOTHECAE

124 ANNALS OF THE SOUTH AFRICAN MUSEUM

0,7 0,8 0,9 1,0
INTERNODE LENGTH (mm)

Fig. 8. The average length, with range, of the intermediate athecate internodes related to the
number of nematothecae which they bear in different samples of Antennella.

False Bay

Table Bay

Agulhas Bank (34°30’S 20°56’E)
West of Cape Peninsula
Lambert’s Bay

Agulhas Bank (34°35’S 21°23’E)
Mossei Bay

Nightingale Island

Saldanha Bay

Off Ltideritz Bay

Vema Seamount

Te ee a 2

fom fee

Tristan da Cunha

. Port Elizabeth

Off Natal (SM 43)

. Off Natal (SM 23)

Gough Island (from Ritchie 1909)
Off Natal (SM 86)

. Off Natal (SM 83)

Off Natal (SM 103)

. Off Natal (SM 38)
. Off Natal (SM 52)

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 125

(Broch 1914), Vema Seamount (Millard 1966), and South Africa (Millard
1975);
Indian Ocean, including India (Leloup 1932) and South Africa (Millard 1975);
Pacific Ocean, including New Zealand (Ralph 1961).
The type locality is Gough Island (Ritchie 1909).

TABLE 1

The percentage number of nematothecae per athecate internode for various samples of
Antennella. The samples are arranged geographically, starting with the Atlantic stations and
west coast, and ending with the Natal coast. The last seven samples are those from this collection.

Number of nematothecae Depth
per internode, per cent n (m)
1 2 3 4

Gough Is. (from Ritchie 1909) . : : 3 — 40 5 5 20 183
Tristan . : , : : ; ; mY ERD 16 2 — 308 s
Nightingale Is... 5 : : : , 2 780 20 _ — 35 ?
Vema Seamount . : : : : 2 ees) 74 pu — 90 42-50
Liideritz Bay. : : : ; ‘ : = 1 269 30 1 -- 226 35
Lambert’s Bay. : : : : : a eo 8 — — 101 20
Saldanha Bay 2 : ; ‘ : : sy) a4 25 1 _ 118 35
Table Bay. 5 : : : i 98 D -- _ 33 9
West of Cape Peninsula : : 5 ; = 95 5 — - 19 79
False Bay... ; : ; = oo 1 — — Sit 0-27
Agulhas Bank: 34°30’ S 20°56’ E . : 0 989 11 -- — 28 3
Agulhas Bank: 34°35’S 21°23’E : ; yes 0) 20 — — 30 68
Mossel Bay . : ; 3 P : 4) 68 Siti -— — 38 10-20
Port Elizabeth . : : : ‘ : : — 100 _— — St 9
Natal, SM 43 : : : ; i 2 95 3 — 113 360-420
Natal,SM 103 ; ; ; . f -— 22 67 1 9 680
Natal, SM 38 : : ; , : _ 29 64 a 14 775-825
Natal, SM 83 : ; : : ‘ : é _ 1] 89 _ 28 600-810
Natal, SM 86 : : 4 ; ; oo 23 Th — 48 550
Natal, SM 23 5 : ; ; : io tee 51 47 2 100 400-450

Natal, SM 52 oe , ; , A es = 80 20 15 720

Cladocarpus dofleini (Stechow, 1911)
Fig. 9D-F

Dinotheca dofleini: Stechow, 1925: 508, figs 49-52. Vervoort, 1966: 162, figs 63-64.
Cladocarpus dofleini: Millard, 1975: 421, fig. 130G.

Description

Two fascicled stems bearing phylactocarps, the longer 90 mm in height.
Structure and dimensions very similar to those of Vervoort (1966), to which the
following points may be added.

The number of cauline nematothecae between two successive hydrocladia
varies from two in the distal region of the stem to six in the proximal part. The
hydrocladial internodes have a shorter distal region than those illustrated by

126 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 9. Cladocarpus sinuosus Vervoort. A. Hydrotheca with an abcauline intrathecal septum,
and B, one without. C. Hydrotheca of var. edentatus Vervootrt.

Cladocarpus dofleini (Stechow). D. Phylactocarp, and E, one of its nematothecae. F. Hydro-
theca.

Scale in mm/10.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES he

Stechow (1925) and are more like those of Vervoort, but the ‘spine’ on the
hydrotheca is long, like that of Stechow. The distal end of the internode is
occasionally cut off as a short athecate internode. The phylactocarps are similar
to those described by Stechow, but have no gonothecae.

Measurements (mm)

Hydrocladium, internodelength . ; : . 0,87-0,98

Hydrotheca, distance from adcauline edge to spine . . 0,42-0,55

Seaimcleteatimianenin, «%. ge 4h Mel non My 9 2020-0,27
Remarks

This rare species is known from the African coast from the equator to the
Agulhas Bank, and occurs only in depths over 425 m.

Cladocarpus natalensis sp. nov.

Fig. 10
Material
Holotype: SAM—H2861. Station SM 86: 27°59,5’S 32°40,8’E, 550 m. Five stems,
fertile.

Other material: SAM-—H1972. Station SM 23: 27°44,4’S 32°42,8’E, 400-40 m.5
Five stems, infertile.

Description of holotype

The tallest stem 50 mm in height and bearing about seventy alternate hydro-
cladia. Hydrorhiza mat-like. Stem fascicled, unbranched, bearing the hydrocladia
from an axial tube which is exposed on the anterior surface. Two cauline nemato-
thecae between the origins of any two consecutive hydrocladia, of which one is
axillary. Five or six septa present between two consecutive hydrocladia in the
older part of the stem, but septa absent in the distal part. Segmentation obscure.

Hydrocladia 4-6 mm in length and bearing up to ten hydrothecae on anterior
surface, consisting of sigmoidally curved thecate internodes separated by straight
or slightly oblique nodes. Each internode with numerous septa (12-16), one
hydrotheca and three nematothecae (one median inferior and one pair laterals).
Distal part of internode short and terminating just above thecal margin.

Hydrotheca sigmoidally curved; abcauline wall very strongly convex
immediately above base, strongly concave above this and with the maximum
concavity at about half height, then widening to margin; base carried forwards
with curvature and taking with it the adthecal ends of three or four internodal
septa; the basal convexity capped by a hollow perisarcal horn of variable length
and direction; adcauline wall strongly curved near base, the curvature decreasing
smoothly to margin. A curved adcauline intrathecal septum present in basal
region and close above thecal floor. Margin with one inturned median abcauline
tooth, with the rest of the edge irregularly serrated and often lower at the
adcauline side, forming an angle of 50-80° with the distal end of the internode.

128 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 10. Cladocarpus natalensis sp. nov. A—D from the holotype, SAM-—H2861, and E-G from

SAM-H1972. A-B. Hydrothecae. C. Phylactocarp, and D, one of its nematothecae.

E-F. Hydrothecae. G. Part of stem showing cauline nematothecae and origins of two hydro-
cladia. Scale in mm/10.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 129

Median inferior nematotheca free from hydrotheca and seated well below
its base, with two apertures, one terminal and one on upper surface near base.
Lateral nematotheca overtopping thecal margin, bifurcated, with two terminal
apertures, one more elevated than the other, and one on mesial surface.

Phylactocarps borne one on first internode of hydrocladium and rarely one
on second internode as well, curved forwards and forming a double row on front
of stem. Phylactocarp bearing a double row of long nematothecae, with many
septa both in the nematothecae and in the main axis. Nematotheca with four
apertures, one terminal, one subterminal, one on the end of a tubular process
near base, and one on the side of this tubular process. Gonothecae absent except
for one empty one; this is oval and with a terminal aperture.

Measurements (mm) Holotype
SAM-—H2861 SAM—H1972
Hydrocladium internode length . 0,51-0,74 0,80—0,96
Hydrotheca, depth, abcauline base to
tip of median tooth : ‘ . 0,18-0,35 0,29-0,58
depth, horn to adcauline edge . . 0,34-0,52 0,43-0,75
diameter at margin. : ; b 01720371 0,23-0,29

Variation and remarks

In this species the degree of curvature of the hydrotheca and the length of
the horn vary markedly from stem to stem, and to a certain degree within the
same stem. In SAM-H1972 the horn may vary from hardly recognizable (as in
C. inflatus Vervoort) to about twice the width of the base, and is usually directed
obliquely downwards. In the holotype the horn is in general longer and the
hydrotheca more curved, so that in extreme cases the horn points obliquely
upwards parallel with the axis of the distal half of the hydrotheca and reaching
well beyond the distal end of the internode. The extreme variations in the two
samples are illustrated in Figure 10.

Other variations include one branching hydrocladium which has developed
as a stem, a branching nematotheca on the phylactocarp (Fig. 10C), and the
presence of minute teeth round the edge of the hydropore in some hydrothecae.
The internodal septa may be as many as 20 in the second sample.

In the curvature of the hydrotheca and the development of the horn this
species is intermediate between C. unicornus Millard, where the internode and its
septa are not involved in the curvature, and C. dofleini (Stechow), where the
curvature is extreme and practically all the septa are involved. C. natalensis also
differs from C. dofleini in the greater number of internodal septa, in the shorter
distal part of the internode and in the longer and septate nematothecae on the
phylactocarp.

Cladocarpus sinuosus Vervoort, 1966
Fig. 9A-C
Cladocarpus sinuosus Vervoort, 1966: 155, figs 55-57. Millard, 1975: 428, fig. 132E-H.

130 ANNALS OF THE SOUTH AFRICAN MUSEUM

Description

These colonies provide further information on the variability of this interest-
ing species.

One colony (Station SM 86) consists of about fifty stems reaching a maxi-
mum height of 38 mm. The hydrocladia have thick perisarc with up to ten inter-
nodal septa and the hydrothecae are more strongly bent than in previous
descriptions. In some of the hydrothecae an abcauline intrathecal septum occurs
at a slightly lower level than the adcauline one, reminiscent of C. /eloupi Millard;
hydrothecae both with and without the abcauline septum may occur on the same
stem.

Phylactocarps occur in this sample, bearing many oval gonothecae on their
inner surfaces between the origins of the nematothecae. The gonothecae have
broad subterminal apertures.

Another colony (Station SM 103) consists of a single infertile stem of 10 mm
with only a few hydrocladia remaining. It clearly belongs to Vervoort’s var.
edentatus, although a very short abcauline marginal tooth does occur in some
of the distal hydrothecae. An interesting feature of this colony is the presence
of a short longitudinal septum against the concavity of the abcauline thecal wall
of some hydrothecae, reminiscent of C. paries Millard.

DISCUSSION

It is not intended to discuss the geographic distribution of the species from
this area in any detail at present, since a forthcoming paper will deal with the
distribution of the whole of the southern African hydroid fauna utilizing many
more records. It may, however, be mentioned in passing that the deep-water
hydroids from off the Natal coast appear to be of an extremely mixed nature,
including endemic, cosmopolitan, tropical and temperate species with no particu-
lar bias to any one component and no clear pointers to the origin of the fauna.

ACKNOWLEDGEMENTS

Financial assistance from SANCOR (The South African National Council
for Oceanographic Research) towards the cost of ship’s time is gratefully
acknowledged, and also the willing and helpful co-operation of all members of
the crew of the Meiring Naude.

I should also like to express thanks to the British Museum (Natural History)
for the loan of slide material for comparison.

REFERENCES

ALLMAN, G. J. 1877. Report on the Hydroida collected during the exploration of the Gulf
Stream by L. F. de Pourtalés, assistant United States coast survey. Mem. Mus. comp.
Zool. Hary. 5: 1-66.

ALLMAN, G. J. 1888. Report on the Hydroida dredged by H.M.S. Challenger during the years
1873-76. Part II. The Tubularinae, Corymorphinae, Campanularinae, Sertularinae and
Thalamophora. Rep. Voy. Challenger 1875-76, Zoology 23 (70): 1-90.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES isi

BILLARD, A. 1906. Hydroides. Expéd. scient. Travailleur-Talisman 8: 153-244.

BILLARD, A. 1923. Note sur quelques Hydroides des cétes de France. Bull. Soc. zool. Fr. 48:
13-20.

BILLARD, A. 1924. Note critique sur divers genres et espéces d’Hydroides avec la description
de trois espéces nouvelles. Revue suisse Zool. 31: 53-74.

BILLARD, A. 1941. Note sur une espéce d’Hydroide peu connue: Stegopoma operculatum
(Nutting). Bull. Soc. zool. Fr. 66: 16-17.

Brocu, H. 1914. Hydrozoa benthonica. Beitr. Kennt. Meeresfauna Westafr. 1: 19-50.

Brocu, H. 1918. Hydroida II. Dan. Ingolf-Exped. 5 (7): 1-205.

Browne, E. T. 1907. The hydroids collected by the ‘Huxley’ from the north side of the Bay
of Biscay in August, 1906. J. mar. biol. Ass. U.K. 8: 15-36.

CoRNELIUS, P. F. S. 1975. A revision of the species of Lafoeidae and Haleciidae (Coelenterata:
Hydroida) recorded from Britain and nearby seas. Bull. Br. Mus. nat. Hist. (Zool.) 28:
375-426.

Epwarps, C. 1973. The medusa Modeeria rotunda and its hydroid Stegopoma fastigiatum, with
a review of Stegopoma and Stegolaria. J. mar. biol. Ass. U.K. 53: 573-600.

JADERHOLM, E. 1919. Zur Kenntnis der Hydroidenfauna Japans. Ark. Zool. 12 (9): 1-34.

JARVIS, F. E. 1922. The hydroids from the Chagos, Seychelles and other islands and from the
coasts of British East Africa and Zanzibar. Trans. Linn. Soc. Lond. (Zool.) 18: 331-360.

Letoup, E. 1932. Une collection d’*hydropolypes appartenant |’Indian Museum de Calcutta.
Rec. Indian Mus. 34: 131-170.

LeLoup, E. 1940. Hydropolypes provenant des Croisiéres du Prince Albert I®? de Monaco.
Résult. Camp. scient. Prince Albert I 104: 1-38.

Louw, E. 1977. The South African Museum’s Meiring Naude cruises. Part I. Station data 1975,
1976. Ann. S. Afr. Mus. 72 (8): 147-159.

MILLARD, N. A. H. 1966. Hydroids of the Vema Seamount. Ann. S. Afr. Mus. 48: 489-496.

MILLARD, N. A. H. 1967. Hydroids from the south-west Indian Ocean. Ann. S. Afr. Mus. 50:
169-194,

MILLARD, N. A. H. 1975. Monograph on the Hydroida of southern Africa. Ann. S. Afr. Mus.
68: 1-513.

NutTING, C. C. 1904. American hydroids. Part II. The Sertulariidae. Spec. Bull. U.S. natn.
Mus. 4 (2): 1-151.

NUuTTING, C. C. 1905. Hydroids of the Hawaiian islands collected by the steamer ‘Albatross’
in 1902. Bull. U.S. Fish Comm. 23: 931-959.

Patriti, G. 1970. Catalogue des Cnidaires et Ctenaires des cotes Atlantiques Marocaines.
Hydraires. Tray. Inst. scient. chérif: (Zool.) 35: 11-60.

RALPH, P. M. 1961. New Zealand thecate hydroids. Part IV. The family Plumulariidae. Trans.
ix, soe. IN.Z. (Zool.) 1: 19-74.

Ritcuigz, J. 1909. Supplementary report on the hydroids of the Scottish National Antarctic
Expedition. Trans. R. Soc. Edinb. 47: 65-101.

Ritcuig, J. 1910. The hydroids of the Indian Museum. Rec. Indian Mus. 5: 1-30.

SAEMUNDSSON, B. 1912. Bidrag til Kundskaben om de islandske Hydroider. II. Vidensk. Meddr
dansk naturh. Foren. 63: 67-107.

STECHOW, E. 1919. Zur Kenntnis der Hydroidenfauna des Mittelmeeres, Amerikas und anderer
Gebiete. Zool. Jb. (System. Abt.) 42: 1-172.

STECHOW, E. 1921. Uber Hydroiden der Deutschen Tiefsee-Expedition, nebst Bemerkungen
uber einige andre Formen. Zool. Anz. 53: 223-236.

STECHOW, E. 1923a. Zur Kenntnis der Hydroidenfauna des Mittelmeeres, Amerikas und
anderer Gebiete. II. Teil. Zool. Jb. (System. Abt.) 47: 29-270.

STECHOW, E. 19236. Uber Hydroiden der Deutschen Tiefsee-Expedition, nebst Bemerkungen
uber einige andre Formen. Zool. Anz. 56: 97-119.

STECHOw, E. 1925. Hydroiden der deutschen Tiefsee-Expedition. Wiss. Ergebn. dt. Tiefsee-
Exped. ‘Valdivia’ 17: 383-546.

ToTToN, A. K. 1930. Coelenterata. Part V. Hydroida. Nat. Hist. Rep. Br. Antarct. Terra Nova
Exped. 5: 131—252.

VERVOORT, W. 1946. Exotic hydroids in the collections of the Rijksmuseum van Natuurlijke
Historie and the Zoological Museum at Amsterdam. Zool. Meded. Leiden 26: 287-351.

VERVOORT, W. 1966. Bathyal and abyssal hydroids. Galathea Rep. 8: 97-174.

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

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

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

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

Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (figs 8a—b).

Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

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

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

In describing new species, one specimen must be designated as the holotype; other speci-
mens mentioned in the original description are to be designated paratypes; additional material
not regarded as paratypes should be listed separately. The complete data (registration number,
depository, description of specimen, locality, collector, date) of the holotype and paratypes
must be recorded, e.g.:

Holotype
SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach,
Port Elizabeth (33°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 Giies of prefixed surnames in all peeeeees. whch used in the text, if not Cane
by initials or full names
e.g. Du Toit but A.L.du Toit; Von Huene but F. von Huene

(c) Scientific names, but not their vernacular derivatives
e.g. Therocephalia, but therocephalian

Punctuation should be loose, omitting all not strictly necessary

Reference to the author should be expressed in the third person

Roman numerals should be converted to arabic, except when forming part of the title of a
book or article, such as
‘Revision of the Crustacea. Part VIII. The Amphipoda.’

Specific name must not stand alone, but be preceded by the generic name or its abbreviation
to initial capital letter, provided the same generic name is used consecutively.

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.

N. A. H. MILLARD

THE SOUTH AFRICAN MUSEUM’S
MEIRING NAUDE CRUISES

PART 3

HYDROIDA

VOLUME 73 PART 6 JUNE 1977 3 ISSN 0303-2515

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
(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

3. MANUSCRIPT, to be submitted in triplicate, should be typewritten and neat, double spaced
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Major headings of the paper are centred capitals; first subheadings are shouldered small
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Footnotes should be avoided unless they are short and essential.

Only generic and specific names should be underlined to indicate italics; all other marking
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4. ILLUSTRATIONS should be reducible to a size not exceeding 12 x 18 cm (19 cm including
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should be given in the legend; if the latter, then the final reduction or enlargement should be
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All illustrations, whether line drawings or photographs, should be termed figures (plates
are not printed; half-tones will appear in their proper place in the text) and numbered in a
single series. Items of composite figures should be designated by capital letters; lettering of
figures is not set in type and should be in lower-case letters.

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 = of publication given in text, e.g.:

‘Smith (1969) describes .

‘Smith (1969: 36, fig. 16) eek es.

‘As described (Smith 1969a, 19695; ke ies
‘As described (Haughton & Broom 1927).

‘As described (Haughton et al. 1927)...’

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

(b) Full references at the end of the paper, arranged alphabetically by names, chronologically
within each name, with suffixes a, b, etc. to the year for more than one paper by the same
author in that year, e.g. Smith (1969a, 19695) and not Smith (1969, 1969a).

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

For journal article give title of article, title of journal in italics (abbreviated according to the World list o
scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses, volume number, part
number (only if independently paged) in parentheses, pagination (first and last pages of article).

Examples (note capitalization and punctuation)

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

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. én. 74: 627-634.

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

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

THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270.

(continued inside back cover)

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

Volume 73 ~~ Band
June 1977 Junie
Part 6 Deel

Yh (Sey,
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THE SOUTH AFRICAN MUSEUM’S
METRING NAUDE CRUISES
PART 4
ECHINODERMS

By

AILSA M. CLARK

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material
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THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES
PART 4
ECHINODERMS

By
AILsA._M. CLARK
British Museum (Natural History), London

[MS. accepted 15 February 1977]

ABSTRACT

This paper lists the echinoderm species recently collected in deep water off Natal and is
annotated to indicate the eighteen new records for the South African area and the resultant
extensions of range. Notes are also given to distinguish the additional species from those keyed
and described in a recent paper.

CONTENTS

PAGE
Introduction : : : p , 133
List of the species collected. 4, 1
Systematic section. : YO “36
Acknowledgements . . : ; 146
References . ; b : : : 146

INTRODUCTION

The material upon which this report was based was collected by the Meiring
Naude during 1975 and 1976 between 26°50’S and 29°13’S off Natal at depths
mostly between 500 and | 300 metres. Further details of the Station List will be
found in the first paper of this series (Louw 1977). One holothurian was inadver-
tently included (pressed against an echinothuriid). Otherwise the collection con-
sists mainly of ophiuroids, with smaller numbers of asteroids, echinoids and
crinoids.

Having just completed what was thought to be a comprehensive survey of
the echinoderm fauna of southern Africa south of Capricorn, except for holo-
thurians (Clark & Courtman-Stock 1976), it was disillusioning to find that no
less than 10 species of ophiuroids, 5 of asteroids, 2 echinoids and the 1 holothurian
are new to the fauna of this area. Most, if not all, of these new records are
extensions of range from the Indo-West Pacific, which was to be expected from
our previous knowledge of the distribution of the echinoderms of south-east
Africa. Unfortunately, positive identifications could not be given for all the
additional species because of inadequacy of material, notably for four of the
starfish species. Hopefully this situation will soon be resolved by further
collections.

iss

Ann. S. Afr. Mus. 73 (6), 1977: 133-147.

134 ANNALS OF THE SOUTH AFRICAN MUSEUM

Apart from the records of species new to southern Africa, this material also
provides minor extensions of range, both horizontal and vertical, for species
already cited in the main faunal study.

In the list of species collected that follows, new records for southern Africa
(south of the Tropic of Capricorn) are marked *. The latitude/longitude degree
grid references and extensions of depth range are given to supplement those in
Clark & Courtman-Stock (1976).

LIST OF THE SPECIES COLLECTED

Class CRINOIDEA

Democrinus chuni (Déderlein, 1907)

Sts SM 16, 53, 61, 75, 78, 86, 94, 103 and 109, 384-1 300 metres, 46 specimens.
Grid additions: 28/32/vd, 26/33/vd (vd = very deep, 500+ metres).
Crotalometra magnicirra (Bell, 1905)

St. SM 86, 550 metres, 2 specimens. Grid addition: 27/32/vd.

Antedonid sp. indet.

Sts SM 38, 66, 86 and 107, 550-1 000 (71 200) metres, 4 poor specimens.

Subclass ASTEROIDEA

Astropecten leptus H. L. Clark, 1926

St. SM 83, 810-600 metres, 1 specimen. Grid addition: 28/32/vd; depth range extended from
375 metres.

Persephonaster roulei euryplax Mortensen, 1933

St. SM 71, 1050 metres, 1 specimen. Grid addition: 27/33/vd; depth range extended from.
410 metres.

Astropectinid sp. juv. ?P. roulei euryplax or Psilaster acuminatus Sladen, 1889

St. SM 66, 780-720 metres, 1 specimen. (R = 22 mm).

*Cheiraster triplacanthus Fisher, 1913

St. SM 71, 1 050 metres, 3 specimens. Grid reference: 27/33/vd.

Previously known from the Moluccas.

* Pseudarchaster sp. aff. P. myobrachius Fisher, 1906

St. 72, 1 050 metres, 1 specimen.

P. myobrachius is known from the Hawaiian Islands.

Calliaster acanthodes H. L. Clark, 1923

SAM-—A22780, Tugela Bank, 1/4/1974, coll. O.R.I., 1 specimen. (Not from Meiring Naude
collection.)

* Henricia sp. ?H. microplax Fisher, 1917

St. SM 86, 550 metres, | specimen.

H. microplax is known from the Philippine Islands.

* Solaster sp. aff. S. paxillatus Sladen, 1889 and S. tropicus Fisher, 1913

Sts SM 44 and 66, 720-768 (?780) metres, 2 specimens.

S. paxillatus is known from south-east Japan to Alaska and S. tropicus from the Moluccas.
Hymenaster sp. indet.

St. SM 107, 1 200-1 000 metres, 3 poor specimens.

Pteraster or Diplopteraster sp. indet.

St. SM 66, 780-720 metres, 1 poor specimen.

* Zoroaster Sp.

St. SM 107, 1 200-1 000 metres, | poor specimen.

Eight species of Zoroaster have been recorded from the Indian Ocean.

Subclass OPHIUROIDEA

Ophioscolex dentatus forma spiniger Mortensen, 1933
Sts SM 16 and 23, 384 (?376)—400 (2450) metres, 5 specimens.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 5

Amphilimna ? sp.

St. SM 53, 720 metres, 1 poor specimen.

Ophiacantha baccata Mortensen, 1933

Sts SM 23 and 86, 450 (?400)—-550 metres, 23 specimens. Depth range extended from 440 metres.
Ophiacantha striolata Mortensen, 1933

St. SM 23, 450-400 metres, 1 poor specimen. Grid addition: 27/32/d (d = 100-499 metres).
*Ophiolimna ?perfida (Koehler, 1904)

St. SM 77, 780 metres, 1 poor specimen. Grid reference 27/32/vd.

O. perfida is known from Indonesia to southern Japan.

*Ophiophthalmus relictus (Koehler, 1904)

Sts SM 43(?), 60(?), 66(?), 74 and 107, 420 (2360) — 1 000 (?1 200) metres, 26 specimens, 17 of
them small and not positively identifiable. Grid references 28/32/vd, 27/32/vd.

Previously known from the Gulf of Aden, Timor and southern Japan.

*Ophioplinthaca papillosa H. L. Clark, 1939

St. SM 107, 1 200-1 000 metres, 2 specimens. Grid reference: 28/32/vd.

Previously known from the Gulf of Aden and the Maldive area.

*Ophioplinthaca rudis (Koehler, 1897)

Sts SM 58, 66, 71, 74, 90 and 107, 780 (2720) — 1 050 (?1 200) metres, 72 specimens. Grid
references: 28/32/vd, 27/32/vd, 27/33/vd.

Previously known from the Zanzibar area to Timor and southern Japan.

Ophiothamnus remotus forma cordatus Mortensen, 1933

Sts SM 16, 23, 60, 78, 86 and 103, 384 (2376) — 800 (?810) metres, 94 specimens. Grid additions:
28/32/vd, 27/32/vd; depth range extended from 457 metres.

Ophiotreta durbanensis (Mortensen, 1933)

Sts SM 23 and 86, 450 (2400) — 550 metres, 5 specimens. Grid addition: 27/32/vd; depth range
extended from 450 metres.

*Ophiotreta matura (Koehler, 1904)

Sts SM 22, 77(?) and 107, 700 (2492) — 1 000 (?1 200) metres, 3 specimens. Grid references:
28/32/vd, 27/32/vd.

Ophiotreta sp. aff. O. matura (Koehler)

St. SM 92, 720-650 metres, 1 small specimen.

Ophiotreta sp. indet.

Sts SM 66 and 67, 680-780 metres, 3 poor specimens.

Amphiura albella Mortensen, 1933

St. SM 67, 700-680 metres, 7 specimens. Grid addition: 27/32/vd; depth range extended from
412 metres.

Amphiura grandisquama natalensis Mortensen, 1933

Sts SM 16, 38, 60, 86 and 103, 384 (2376) — 800 (2825) metres, 33 specimens. Grid additions:
28/32/vd, 27/32/d, vd.

*Amphiura sp. aff. A. atlantica Ljungman, 1867

Sts SM 53 and 60, 720-800 (?810) metres, 13 small specimens.

Amphipholis squamata (Delle Chiaje, 1828)

Sts SM 78, 86, 94 and 103, 550-750 metres, 11 small specimens. Grid additions: 28/32/vd,
27/32/vd; depth range in South Africa extended from 172 metres.

Amphilepis scutata Mortensen, 1933

Sts SM 53 and 60, 720-800 (?810) metres, 4 specimens. Grid additions: 27/32/vd, 26/33/vd;
depth range extended from 410 metres.

*Histampica duplicata (Lyman, 1875)

Sts SM 66, 77 and 103, 680-780 metres, 5 specimens. Grid references: 28/32/vd, 27/32/vd.
Previously known from the tropical West Atlantic and East and West Pacific.

Ophiothrix aristulata Lyman, 1879

Sts SM 16(?) and 23, 384 (?376) — 400 (2450) metres, 3 small specimens. Grid addition: 27/32/d.
* Anophiura simplex H. L. Clark, 1939

St. SM 53, 720 metres, 1 specimen. Grid reference: 26/33/vd.

Previously known from south of Arabia.

*Aspidophiura corone Hertz, 1927

Sts SM 31, 53, 60 and 78, 720-800 (?810) metres, 56 specimens. Grid references: 28/32/vd,
27/32/vd, 26/33/vd.

Previously known from off equatorial east Africa.

136 ANNALS OF THE SOUTH AFRICAN MUSEUM

Astrophiura permira Sladen, 1879 .
Sts SM 16, 23, 60, 86, 94, 103 and 109, 384 (2376) — 1 300 metres, 74 specimens. Grid additions:
28/32/vd, 27/32/vd; depth range extended from 376 metres.

* Homalophiura schmidtotti (Hertz, 1927) comb. n.

St. SM 38, 775-825 metres, 1 specimen. Grid reference: 28/32/vd.
Previously known from off equatorial east Africa and Indonesia.

Ophiura flagellata (Lyman, 1878)

St. SM 107, 1 200-1 000 metres, 8 specimens. Grid addition: 28/32/vd.
Ophiurid sp. juv. ? Homalophiura sp.

St. SM 109, 1 300 metres, 1 small specimen.

Ophiopallas paradoxa Koehler, 1904

St. SM 16, 376-384 metres, 1 small specimen.

Class ECHINOIDEA

Araeosoma paucispinum H. L. Clark. 1924

St. SM 15, 454-280 metres, 3 specimens.

Phormosoma bursarium A. Agassiz, 1881

Sts SM 22, 38, 53 and 107, 700 (2492) — 1 000 (?1 200) metres, 7 specimens. Grid additions:
28/32/vd, 26/33/vd; depth range extended from 840 metres.

Temnopleurus reevesi (Gray, 1855)

St. SM 23, 450-400 metres, 1 small specimen. Grid addition: 27/32/d; depth range extended
from 102 metres.

*Echinocyamus scaber forma subconicus Mortensen, 1948

Sts SM 15, 23, 60, 69, 78, 86 and 103, 450 (?280) — 800 (?810) metres, 30 specimens. Grid
references: 28/32/vd, 27/32/d, vd.

Forma subconicus previously known from the Kei Islands, Indonesia but E. scaber from off
equatorial east Africa to south-east Australia and the Hawaiian Islands.

*Gymnopatagus magnus A. Agassiz & H. L. Clark, 1907

St. SM 107, 1 200-1 000 metres, upper side only of 1 specimen. Grid reference: 28/32/vd.
Previously known from the Andaman Islands to Indonesia and Japan.

Class HOLOTHURIOIDEA

*Orphnurgus glaber Walsh, 1891
St. SM 107, 1 200-1 000 metres, 1 specimen (squashed against an echinothuriid in preservation).
Previously known from the Bay of Bengal to Japan and the Hawaiian Islands.

SYSTEMATIC SECTION

Class CRINOIDEA
Family Thalassometridae

Crotalometra magnicirra (Bell)

Antedon magnicirra Bell, 1905: 141, pl. 4.
Crotalometra magnicirra: Gislén, 1938: 17-18. A. H. Clark, 1950: 97-100. A. M. Clark, 1974:
427-429. A. M. Clark & Courtman-Stock, 1976: 18.

St. SM 86, 27°59,5’S 32°40,8’E, 550 metres, 2 specimens.

One specimen is small. The other has arm length 70+ mm, probably c.
100 mm. The high rugose centrodorsal is 4,6 mm high, the breadth (obscured by
the cirri) approximately the same.

The specimen is unusual in having 6 out of the 7 second division (IIBr)
series present with only 2 ossicles rather than the 4 characteristic of the genus
Crotalometra. Only a single such short IIBr series was found in all 16 specimens

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 137

of the type series studied in 1974, though Gislén (1938) found 4 out of 10 such
series in one of his specimens.

The mature cirri number c. XXVII, with c. vii immature ones, arranged in
10 columns of usually 3 cirri, separated into 5 pairs of columns by narrow
triangular bare radial areas, as in the type series. Two intact peripheral cirri
have 68 and 66 segments, the maximum otherwise recorded for this species being
64; also the seventh segment is the transition one, rather than the ninth or tenth.
The closely appressed division series are finely rugose laterally, where the
syntypes are smooth.

Finally the first pinnule (which is P, in this case since the I[Br series have
only two ossicles) has 29+ c. 2 segments and measures just over 14 mm, com-
pared with up to 27 segments and c. 12 mm length in Pp of the syntype described
in 1974; otherwise the shape is similar.

Despite these small differences, there does not appear to be sufficient
justification for regarding this sample as specifically distinct from Crotalometra
magnicirra, especially considering the geographical approximation, the nearest
previous record for the species being c. 29,5°S 31,5°E.

Of the two other species of Crotalometra, C. rustica, known from the Bay of
Bengal to Indonesia, and C. sentifera, from the Maldive area, both have the
brachials beyond the arm bases more or less markedly produced and spinose at
their distal edges.

Antedonidae sp(p). indet.

St. SM 38, 28°21,9’S 32°34,6’E, 775-825 metres, 1 broken specimen.

St. SM 66, 27°17,5’S 32°54,1’E, 780-720 metres, 1 small specimen.

St. SM 86, 27°59,5’S 32°40,8’E, 550 metres, 1 small specimen.

St. SM 107, 28°37,8'S 32°38,4’E, 1 200-1 000 metres, | small specimen.

The best of these specimens, from station SM 107, at least seems to have
some affinity with Tonrometra of the subfamily Bathymetrinae except that P, is
smaller than P, or P3, not larger. The centrodorsal is low conical and there is
some tendency for arrangement of the sockets in vertical rows. The cirrus seg-
ments number up to 25, the distal ones short and keeled dorsally. The division
series and brachials have spinose distal edges.

Class STELLEROIDEA
Subclass ASTEROIDEA
Family Astropectinidae

Astropecten leptus H. L. Clark

Astropecten leptus H. L. Clark, 1926: 6-8, pl. 1 (figs 3-4). Mortensen, 1933: 234, pl. 9 (figs 3-4).
Madsen, 1950: 169. Cherbonnier & Nataf, 1973: 1268-1272, fig. 8A—F, pl. 12. A. M. Clark
& Courtman-Stock, 1976: 51.

St. SM 83, 28°00,5’S 32°46,4’E, 810-600 metres, | specimen.
R/r (the major to minor radius) = 40/8 mm = 5,0/1, compared with
59/11 mm = 5,3/1 in the holotype of A. /eptus from off Durban in 287-348

138 ANNALS OF THE SOUTH AFRICAN MUSEUM

metres. This specimen differs in having no enlarged superomarginal spines, the
occurrence of which is variable in most species of Astropecten. It also has 3
actinal plates each side of the interradius compared with 2 in the holotype, though
Madsen found up to 4 and Cherbonnier and Nataf up to 5 in their specimens
from West Africa.

Family Benthopectinidae

Cheiraster triplacanthus Fisher

Cheiraster triplacanthus Fisher, 1913: 206. Fisher, 1919: 205-208, pl. 48 (figs 3-4), pl. 55

(figs 1, la).

St. SM 71, 27°21,3'S 33°03;9'E, W050 metres, 3 specimens:

In the asteroid key of Clark & Courtman-Stock (1976: 33) this species runs
down to Luidiaster hirsutus from which it differs in the greater extent of the
superomarginal plates on the upper side and the triple distal superomarginal
spines.

This record represents an extension of range from Celebes in the Moluccas
in | 280 metres, only the holotype being known hitherto. .

The largest specimen has R/r 42/9 mm = 4,7/1, compared with 49/12 mm
= 4,1/1 in the holotype. The two or three up and inwardly directed slightly
curved superomarginal spines on each plate near the arm tips, characteristic of
the species, are very distinctive; also the subambulacral spines are not at all »
enlarged. The bilobed papularia each have c. 35 pores, whereas Fisher estimated
c. 80 in the holotype.

Family Goniasteridae
Pseudarchaster sp. aff. P. myobrachius Fisher, 1906

See: Fisher 1906: 1037.

St. SM 725 27-17,8 5 33 0455 EB: f O50 metres, Ispecimenr

R/r = 44/16 mm = 2,8/1. This ratio is intermediate between those cited
in Clark & Courtman-Stock (1976: 35) of 3/1 or more for P. tessellatus Sladen,
1889 and 2,0-2,7/1 for the shorter-armed P. brachyactis H. L. Clark, 1923,
already known from southern Africa but not on the south-east side. Recent
studies by Halpern (unpublished thesis on the family Goniasteridae) indicate
that the range of variation in the proportions of Psewdarchaster species is much
greater than has previously been supposed. However, there are also a number of
morphological differences between this specimen and P. tessellatus.

These include the smaller number of marginal plates, 26 in each series
compared with c. 35 in tessellatus of similar R, the capitate rather than tapering
form of the relatively short armament of the actinal plates and the absence of
enlarged spinelets on these plates, as well as the shortness of the spines on the
interradial inferomarginals. These two last characters agree with P. myobrachius
Fisher, 1906, in which R/r of the holotype is 2,6/1 and there are 23 supero-

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 139

marginals in each series at R 34 mm. However, that species has been recorded
only from the Hawaiian Islands (in 780-1 240 metres).

Family Echinasteridae
Henricia sp. ? H. microplax Fisher, 1917

See: Fisher 1919: 437.

Si. SM 86, 27°59,5'S 32°40,8'E, 550 metres, 1 specimen.

Among the three species of Henricia from southern Africa cited in the
asteroid key of Clark & Courtman-Stock (1976: 42), this specimen differs from
H. abyssalis in the more solid opaque spinelets and from H. retecta and H. ornata
in the more compact skeleton and the multiple series of spinelets on the adambu-
lacral and actinal plates.

It agrees with H. microplax Fisher, from the Philippines, in the fine and very
compact skeleton with only single papulae in the individual meshes, the rela-
tively conspicuous inferomarginal plates (at least in the proximal half of the arm)
with two or three series of intermarginal plates separating them near the arm
base from the smaller superomarginal series and the multiple series of subambu-
lacral spines. Proximally there are no papulae below the superomarginals but in
the distal half of the arm where the plate arrangement is more irregular the
papulae approximate to the adambulacrals.

Family Solasteridae

Solaster sp. aff. S. paxillatus Sladen, 1889 and S. tropicus Fisher, 1913

Solaster sp., possibly S. paxillatus Sladen or S. tropicus Fisher, A. M. Clark & Courtman-
Stock, 1976: 40 (footnote).

See: Sladen 1889: 452: Fisher 1919: 444.

St. SM 44, 29°40,6’S 32°32,5’E, 722-768 metres, 1 specimen.

St. SM 66, 27°17,5’S 32°54,1’E, 780-720 metres, 1 small specimen.

In comparison with the multiradiate South African solasterid Crossaster
penicillatus Sladen, 1889, the skeleton of these two eight-armed specimens is
much finer and more compact. Their affinities lie with S. paxillatus from south-
east Japan and S. tropicus from the Moluccas, which are closely related. The
holotypes of both have nine arms, but Fisher (1911) found additional North
Pacific specimens of S. paxillatus to have eight to ten arms.

Subclass OPHIUROIDEA
Family Ophiacanthidae
Ophiolimna sp. ? O. perfida (Koehler, 1904)
See: Koehler 1922: 64.
St. SM 77, 27°31,6’S 32°50,0’E, 780 metres, | specimen.

In the ophiuroid key of Clark & Courtman-Stock (1976: 121) this specimen
cuts across dichotomy no. 68, having the apical oral papilla (or pair of papillae)

140 ANNALS OF THE SOUTH AFRICAN MUSEUM

somewhat blunted and a very broad fourth oral papilla present each side,
coupled with very large lateral arm plates. The distal oral papilla, the relatively
broad, rounded single tentacle scale and the coarse granules on the oral plates
serve to distinguish it from the other South African Ophiacanthidae. It agrees
with Ophiolimna perfida Koehler from the Philippines and Indonesia in the
coarse granules on the oral plates, the operculiform distal oral papilla overlying
the second oral tentacle scale, the very long and quite smooth arm spines and
the single large rounded tentacle scale. The only difference noted is that the disc
spines are more elongate than usual in O. perfida, the height sometimes exceeding
twice the breadth (compare Koehler 1922, pl. 92 (fig. 6)).

Ophiophthalmus relictus (Koehler)

Ophiacantha relicta Koehler, 1904: 106-107, pl. 17 (figs 4-6).
Ophiophthalmus relictus: Koehler, 1922: 124-127, pl. 9 (figs 1-4), pl. 95 (fig. 3). H. L. Clark,
1939: 54-55.

St. SM 43, 28°45,5’S 32°24,5’E, 420-360 metres, 10 specimens.

St. SM 60, 27°09,6’S 32°58,2’E, 800-810 metres, 6 small specimens ( ?re/ictus).

St. SM 66, 27°17,5’S 32°54,1’E, 780-720 metres, | small specimen ( ?re/ictus).

St. SM 74, 27°38,6’S 32°52,6’E, 860 metres, | specimen.

St. SM 107, 28°37,8’S 32°38,4’E, 1 200-1 000 metres, 8 specimens.

This species cuts across dichotomy no. 69 in the 1976 key, having moderately
large but well-separated radial shields; this character, together with the granules °
bordering the dorsal arm plates, serves to distinguish it from the other ophiacan-
thids of southern Africa.

The disc scaling is just distinct between the low spaced rugose granules. In
the smaller specimens the granules tend to be limited to the edges of the scales.
The bare radial shields are separated by about their own breadth. The dorsal arm
plates are rhombic or slightly bell-shaped, usually with bead-like granules along
their distal edges. The arm spines are only finely rugose and relatively short,
only the uppermost of the six on the first two free segments exceeding the segment
in length.

Ophioplinthaca papillosa H. L. Clark
Ophioplinthaca papillosa H. L. Clark, 1939: 49-51, figs 10-11.

St. SM 107, 28°37,8’S 32°38,4’E, 1 200-1 000 metres, 2 specimens.

This species cuts across dichotomy no. 75 in the 1976 key, having five arms
but the distalmost oral papilla papilliform. Apart from the number of arms, it
differs from Ophioplinthaca sexradia in having very long arm spines.

The holotype of O. papillosa was collected at the same John Murray
Expedition station in the Gulf of Aden at 1 270 metres as a specimen of O. rudis
(Koehler) and off Natal the two still appear to be sympatric. Although both
species have the interradially creased disc and large bare radial shields charac-
teristic of the genus, O. rudis has the radial shields separated and the disc spine-
lets (if present) slender, whereas in O. papillosa the radial shields are more or less

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 141

broadly contiguous and the armament is in the form of short stumps with thorny
crowns, also the oral papillae of O. rudis are much more numerous and irregular
than the three each side of O. papillosa.

Ophioplinthaca rudis (Koehler)

Ophiomitra rudis Koehler, 1897: 358, pl. 9 (figs 74-75).
Ophioplinthaca rudis; Koehler, 1922: 142-147, pl. 24 (figs 1-6), pl. 96 (fig. 1). H. L. Clark,
1939: 46-47.

St. SM 58, 27°09,7’S 33°01,0’E, 850-790 metres, 2 specimens.

St. SM 66, 27°17,5’S 32°54,1’E, 780-720 metres, | specimen.

Seo 71, 27° 21,3°S 33°03,9 E, 1 050 metres, 32 specimens.

St. SM 74, 27°38,6’S 32°52,6’E, 860 metres, 85 specimens.

St. SM 90, 28°09,8’S 32°47,4’E, 940 metres, 3 specimens.

St. SM 107, 28°37,8’S 32°38,4’E, 1 200-1 000 metres, 54 specimens.

This species falls with difficulty into the Ophiacanthidae as defined in
dichotomy no. 68 of the 1976 key, since the oral papillae are numerous and
irregular, though the large lateral arm plates and the long arm spines should
indicate this family. Also the radial shields, though large and bare, are not
contiguous, thus blurring the division of no. 69. Other comments on the distinc-
tion of O. rudis are given under O. papillosa above.

Ophiotreta matura (Koehler)

Ophiacantha matura Koehler, 1904: 112-113, pl. 23 (figs 2-4).
Ophiotreta matura: Koehler, 1922: 76-81, pls 12-14, pl. 15 (figs 1-3). H.L. Clark, 1939:
53-54. A. M. Clark & Courtman-Stock, 1976: 1, 121 (footnotes).

St. SM 22, 27°45,0’S 32°44,8’E, 700-492 metres, 1 specimen.

St. SM 77, 27°31,6’S 32°50,0’E, 780 metres, 1 specimen.

St. SM 107, 28°37,8’S 32°38,4’E, 1 200-1 000 metres, | specimen.

This species too has multiple and irregular oral papillae but the large lateral
arm plates ally it with this family. It is distinguished from Ophiotreta durbanensis
in a footnote on p. 121 of the 1976 key.

Ophiotreta sp. aff. O. matura (Koehler)

See: Koehler 1922: 76.

St. SM 92, 28°14,5’S 32°40,6’E, 720-650 metres, 1 small specimen.

Whereas preserved specimens of Ophiotreta matura usually retain (or show)
a dark colouration of the arm spines, this specimen is white throughout and the
spines are transparent. They are also much more thorny than in O. matura and
even the tentacle scales are thorny. The long disc spinelets form a dense coat;
they end in several thorns. The lowest arm spine becomes hooked on the distal
segments, as also in O. matura.

142 ANNALS OF THE SOUTH AFRICAN MUSEUM
Family Amphiuridae
Amphiura sp. aff. A. atlantica Ljungman, 1867

See: Mortensen 1933: 351.

St. SM 53, 26°51, 1'S°33"12,5/E; 720 metres; 2 specimens:

St. SM 60, 27°09,6’S 32°58,2’E, 800-810 metres, 11 small specimens.

The largest specimen, from station 53, has d.d. (disc diameter) 4,5—5,0 mm,
the radial shields are completely separate, equal in length to about half the disc
radius, the disc scaling is rather deficient ventrally, the oral plates have an almost
superficial flange bearing the first oral tentacle scale, the distal oral papilla is
spiniform, the consecutive dorsal arm plates are narrowly separated by the lateral
arm plates, there are five arm spines proximally, the lower ones tending to be
slightly truncated and even a little bihamulate with a suggestion of a distal hook
and there is one small tentacle scale. The reduced ventral scaling, the modified
arm spines and the small tentacle scale agree with Amphiura atlantica, to which
species it runs down in the 1976 key; also the smaller specimens (d.d. up to only
2 mm) do have the radial shields just contiguous and lack tentacle scales
altogether, which may be true of A. atlantica. However, the flanged oral plates
are quite distinct, approximating to those of Amphilepis, which is ranged in a
separate subfamily of Amphiuridae. They are shared by Amphiura pycnostoma
H. L. Clark, 1911, from south-east Japan but that species has the distal oral
papilla papilliform, the successive dorsal arm plates contiguous and only three
arm spines. The holotype of A. pycnostoma has one small tentacle scale.

Family Ophiactidae
Histampica duplicata (Lyman)

Ampniura duplicata Lyman, 1875: 19, fig. 87, pl. 5 (fig. 78). Lyman, 1882: 136, pl. 17 (figs 10-12).
Amphiactis duplicata: Koehler, 1922: 204-205, pl. 63 (figs 1-4).
Histampica duplicata: A. M. Clark, 1970: 73.

St. SM 66, 27°17,5’S 32°54,1’E, 780-720 metres, 2 specimens.
St. SM 77, 27°31,6’S 32°50,0’E, 780 metres, | specimen.
St. SM 103, 28°31,7’S 32°34,0’E, 680 metres, 2 specimens.

This species, currently included in the Ophiactidae, may not run down to that
family in dichotomy no. 60 of the 1976 key since its apical oral papilla is not
markedly broadened and is in series with several lateral oral papillae each side.
Difficulty may also arise at no. 68 since the oral structure may be confused with
that of some ophiacanthids. The very coarse, well-defined, naked disc scales,
separate radial shields, smooth and not excessively long arm spines, two tentacle
scales and usual subdivision transversely of the proximalmost ventral arm plate
into two, should serve to distinguish Histampica duplicata.

It is likely that this species will prove to be cosmopolitan.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 143

Family Ophiuridae
Anophiura simplex H. L. Clark
Anophiura simplex H. L. Clark, 1939: 119-121, figs 55-56.

Se oll 53, 26°51,1’S 33°12,5'E, 720 metres, 1 specimen.

In the 1976 key* this species cuts across dichotomy no. 102, since only the
first three pairs of pores are present on each arm but the pores of the first segment
have more than one scale. The unique holotype of A. simplex, from the south
Arabian coast, had lost its oral papillae, which are intact on this specimen and
are close-fitting and rectangular, like those of Anophiura planissima H. L. Clark,
1939, from the Gulf of Aden. The absence of arm combs distal to the radial
shields, the thin disc with a single large interradial plate between the neighbouring
radial shields and the reduction of the tentacles after the basal pores should
serve to distinguish this species from the others known from southern Africa.

The Natal specimen has d.d. 6,0 mm and a.l. (arm length) 10+1 or 2 mm,
so that a.l./d.d. probably does not exceed 2/1, confirming H. L. Clark’s estimate
that the arms of A. planissima are more attenuated at a ratio of c. 2,5/1 than
those of A. simplex. The matt, rather than shiny, surface texture and the prolonged
lateral angles of the proximal ventral arm plates also agree with A. simplex
rather than A. planissima.

Aspidophiura corone Hertz
Aspidophiura corone Hertz, 1927: 79-80, pl. 7 (figs 1-2).

St. SM 31, 28°04,5’S 32°42,8’E, 740 metres, | small specimen.

Sie yies3, 20 St,1’S 33°12,5’E, 720 metres, 16 specimens.

St. SM 60, 27°09,6’S 32°58,2’E, 800-810 metres, 35 specimens.

Si7 5 76, 27 3156'S 32°50,0’E, 750 metres, 4 specimens.

Like Anophiura simplex, this species has reduced tentacle pores and single
large interradial disc scales between the radial shields, but it differs in having well-
developed spiniform arm combs distal to the radial shields and the rosette of
primary disc plates very conspicuous, fully contiguous and with a distinct boss
on the central plate.

Astrophiura permira Sladen

Astrophiura permira Sladen, 1879: 401-415, pl. 20. Hertz, 1927: 83-85, pl. 7 (figs 4-5). Morten-
sen, 1933: 394-396, figs 90-91. A. M. Clark & Courtman-Stock, 1976: 188.

St. SM 16, 27°33,0’S 32°44,6’E, 384 metres, 17 specimens.

St. SM 23, 27°44,4’S 32°42,8’E, 450-400 metres, 10 specimens.
St. SM 60, 27°09,6’S 32°58,2’E, 800-810 metres, 22 specimens.
St. SM 86, 27°59,5’S 32°40,8’E, 550 metres, 5 specimens.

* Note an error in dichotomy 93 of the 1976 key (Clark & Courtman-Stock 1976: 124)
in which the second alternative should lead to no. 101, not 100.

144 ANNALS OF THE SOUTH AFRICAN MUSEUM

St. SM 94, 28°16,3’S 32°38,8’E, 670 metres, 2 specimens.

St. SM 103, 28°31,7’S 32°34,0’E, 680 metres, 17 specimens.

St. SM 109, 28°41,0’S 32°36,8’E, 1 300 metres, 1 specimen.

The arms of this species are almost invariably broken but a very few of these
are intact. A very young complete specimen with the true disc diameter (less the
marginal fringe) 1,1 mm and the expanded disc diameter 1,7 mm, has a free arm
of five segments (the first segment becoming incorporated into the disc) measuring
1,7 mm in length. A larger specimen with expanded disc diameter 4,5 mm has a
complete arm of eight free segments measuring 2,5 mm to the edge of the marginal
fringe. The terminal ossicle is slightly tapering but otherwise almost cylindrical.
At this size, there are five pairs of podia within the disc. A detached complete
free arm of sixteen segments measures 6,5 mm. The maximum known expanded
disc diameter in this species is 14 mm.

Homalophiura schmidtotti (Hertz) comb. n.
Ophiuroglypha schmidt-otti Hertz, 1927: 91-93, fig. 5, pl. 7 (figs 11-12).

St. SM 38, 28°21,9’S 32°34,6’E, 775-825 metres, 1 specimen.

Mortensen (1933) reduced Ophiuroglypha Hertz to the rank of a subgenus
of Ophiura but restored it in 1936 (Mortensen 1936: 316) when dealing with the
type species, Ophioglypha lymani Ljungman, 1870. The main distinction of
Ophiuroglypha was a modification of the middle one of the three arm spines on
the distal segments into an outwardly turned hook. Comparison of O. schmidtotti
with O. lymani shows marked differences. O. /ymani has relatively longer,
markedly carinate arms, with the successive dorsal arm plates quite broadly
contiguous, the arms straight-sided, a much smoother disc, discrete apical oral
papillae and tentacle pores extending to about the twelfth arm segment.
Conversely, there is a close resemblance between O. schmidtotti and Ophioglypha
inornata Lyman, 1878, the type-species of Homalophiura. Both have the second
oral tentacle pore opening completely outside the oral slit (in Ophiuroglypha
lymani the pore runs into the oral slit), only four or five proximal arm segments
with tentacle pores, the arm combs more or less reduced with short bead-like
papillae, the disc and arm plates markedly thickened and the oral papillae and
tentacle scales rectangular and very close-fitting. The main difference is the
greater number of disc scales and the smaller size of the primaries, which are
separated by only single series of relatively large scales in H. inornata, an Atlantic
species. H. schmidtotti is very similar to Homalophiura glypta H. L. Clark, 1939,
from the Maldive area, especially in the apparently sunken middle parts of the
larger disc scales, but these are relatively smaller and have a very shiny texture in
HA. glypta.

In the 1976 key, this species runs down to no. 112 but differs from Ophiura
(Ophiuroglypha) irrorata and costata in having the radial shields asymmetrically
contiguous mid-radially, all the oral papillae rectangular and very close-fitting
and the arms markedly moniliform.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 145

The present specimen has d.d. 8,0 mm; the arms are all broken within 10 mm
but were probably not more than as long again, so that a.l./d.d. was probably
2,0—2,5/1.

Class ECHINOIDEA
Family Fibulariidae

Echinocyamus scaber forma subconicus Mortensen
Echinocyamus scaber forma subconicus Mortensen, 1948: 188, pl. 46 (figs 34-36).

St. SM 15, 28°31,0’S 32°45,6’E, 454-280 metres, 1 specimen.

St. SM 23, 27°44,4’S 32°42,8’E, 450-400 metres, 14 specimens.

St. SM 60, 27°09,6’S 32°58,2’E, 800-810 metres, 1 specimen.

Sigs 69, 27°12,2'S 32°56,0’E, 660 metres, .15 specimens.

Seo 7s, 27 31,6'S 32°50,0’E, 750 metres, 2 specimens.

Si sM 86, 27°59,5'S 32°40,8’E, 550 metres, 6 specimens.

St. SM 103, 28°31,7’S 32°34,0’E, 680 metres, 4 specimens.

Several of these specimens were alive when collected and are complete with
spines. Their colour in spirit is green. Although the periproct is usually bare of
spinelets, one specimen has c. 12 periproctal spines and another about 6. This
throws some doubt on the weight of this character which is one of those supposed
by Mortensen to distinguish the subgenus Mortonia. Only Echinocyamus australis
(Desmoulins), from the Hawaiian and neighbouring Pacific Islands, and E. poly-
porus Mortensen, from New Zealand, were included by Mortensen in Mortonia.

The test is usually low rounded subconical, much as in the specimen from
the Kei Islands illustrated by Mortensen. Length/breadth/height of one specimen
is 7,75/6,80/3,75 mm, 1|/ht = 2,1/1. In this specimen there are three pore pairs
on each side of each petal.

The largest specimen, from station SM 86, is flatter than most, 1|/br/ht
8,3/7,5/3,0 mm, |/ht = 2,8/1. Its periproct is naked, with relatively few plates.
There are four pore pairs on each side of each petal. All the ocular pores on the
apical system, except for the posterior one, are enlarged to the same size as the
genital pores, as in Echinocyamus grandiporus Mortensen from the West Indies
and Azores. This last character is not shared by the other specimens.

Echinocyamus scaber runs down to dichotomy no. 43 in the Clark &
Courtman-Stock key (1976: 211). It is distinguishable from Echinocyamus elegans
by the presence of glassy tubercles scattered over the upper side, by the much
shorter petals, E. e/egans having eight or nine pore pairs each side at this size,
and by the slightly conical shape.

Family Brissidae

Gymnopatagus magnus A. Agassiz & H. L. Clark

Gymnopatagus magnus A. Agassiz & H. L. Clark, 1907: 133. Mortensen, 1951: 447-449, figs
219, 220a, pl. 26 (figs 7-9), pl. 27 (figs 8-10), pl. 61 (figs 19-26).

St. SM 107, 28°37,8’S 32°38,4’E, 1 200-1 000 metres, upper side of one
specimen.

146 ANNALS OF THE SOUTH AFRICAN MUSEUM

In the 1976 key, this species runs down to dichotomy no. 53. It is dis-
tinguished from Spatagobrissus mirabilis by the well-developed frontal notch in
the test and from Brissopsis lyrifera capensis by the enlarged aboral primary
spines and the shape of the peripetalous fasciole, which is not concave between
the petals.

Class HOLOTHURIOIDEA
Family Deimatidae

Orphnurgus glaber Walsh

Orphnurgus asper var. glaber Walsh, 1891: 198.
Orphnurgus glaber: Hansen, 1975: 39-46, figs 12-13, pl. 8 (figs 5-7).

St. SM 107, 28°37,8’S 32°38,4’E, 1 200-1 000 metres, 1 squashed specimen.

This specimen is in poor condition, completely flattened and bearing the
imprint of the echinothuriid against which it was pressed. There are twenty
tentacles. Hansen stresses the variability of the spicules in this species. Here they
are mostly in the form of rods with branching or ornamented ends, many
resembling Hansen’s figures 13.30, 41 and 43 but others are shorter and stout
with elaborately spiny ends, like some of the spicules of Orphnurgus asper Théel,
from the West Indies.

ACKNOWLEDGEMENTS

I am indebted to the authorities of the South African Museum and particu-
larly to Dr N. A. H. Millard for the opportunity of studying this very interesting
collection.

REFERENCES

Aaassiz, A. & CLARK, H. L. 1907. Preliminary report on the Echini collected by the U.S. Fish
Commission steamer ‘Albatross’. Bull. Mus. comp. Zool. Hary. 51: 107-139.

BELL, F. J. 1905. The Echinoderma found off the coast of South Africa. 4. Crinoidea. Mar.
Invest. S. Afr. 4: 139-142.

CHERBONNIER, G. & NATAF, G. 1973. Astropecten des cétes occidentales d’Afrique. Bull. Mus.
natn. Hist. nat. Paris (3) Zool. 120: 1232-1300.

CiarK, A. H. 1950. A monograph of the existing crinoids. 1 (4c). Bull. U.S. natn. Mus. 82
(4c): 1-383.

CiarK, A. M. 1970. Notes on the family Amphiuridae. Bull. Br. Mus. (nat. Hist.) Zool. 19:
1-81.

CLARK, A. M. 1974. Notes on some echinoderms from southern Africa. Bull. Br. Mus. (nat.
Hist.) Zool. 26: 421-487.

CLARK, A. M. & CouRTMAN-STOCK, J. 1976. The echinoderms of southern Africa. London:
British Museum (Nat. Hist.).

_ CxarK, H. L. 1911. North Pacific Ophiurans in the collection of the United States National
Museum. Bull. U.S. natn. Mus. 75: 1-302.

CLARK, H. L. 1923. The Echinoderm fauna of South Africa. Ann. S. Afr. Mus. 13: 221-435.

CLARK, H. L. 1926. Echinoderms from the South African Fisheries and Marine Biological
Survey. 2. Sea-stars (Asteroidea). Rep. Fish. mar. biol. Surv. Un. S. Afr. No. 4. Spec. Rep.
No. 7: 1-34.

CLARK, H. L. 1939. Ophiuroidea. Scient. Rep. John Murray Exped. 4 (2): 29-136.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 147

FisHER, W. K. 1906. The starfishes of the Hawaiian Islands. Bull. U.S. Fish. Commn 1903:
987-1130.

FISHER, W. K. 1911. Asteroidea of the North Pacific and adjacent waters. 1. Bull. U.S. natn.
Mus. 76 (1): 1-419.

FisHER, W. K. 1913. New starfishes from the Philippine Islands, Celebes and the Moluccas.
Proc. U.S. natn. Mus. 46: 201-224.

FISHER, W. K. 1919. Starfishes of the Philippine Seas and adjacent waters. Bull. U.S. natn. Mus.
100 (3): 1-546.

GISLEN, T. 1938. Crinoids of South Africa. K. svenska Vetensk Akad. Handi. (3) 17 (2): 1-22.

HALPERN, J. A. 1970. A monographic revision of the Goniasterid sea stars of the North Atlantic.
Unpublished thesis, University of Miami.

HANSEN, B. 1975. Systematics and biology of the deep-sea holothurians. 1. Elasipoda. Galathea
Rep. 13: 1-262.

Hertz, M. 1927. Die Ophiuroiden der deutschen Tiefsee-Expedition. 1. Wiss. Ergebn. dt.
Tiefsee-Exped. Valdivia 22: 59-122.

KOEHLER, R. 1897. Echinodermes recueillis par l’Investigator dans |’Ocean Indien. 1. Les
ophiures de mer profonde. Annls Sci. nat. (8) 4: 277-372.

KOEHLER, R. 1904. Ophiures de l’Expedition du Siboga. 1. Ophiures de mer profonde. Siboga
Exped. 45a: 1-167.

KOEHLER, R. 1922. Ophiurans of the Philippine Seas and adjacent waters. Bull. U.S. natn. Mus.
100 (3): 1-486.

LJUNGMAN, A. 1867. Ophiuroidea viventia huc usque cognita enumerat. Ofvers. K. Vetensk-
Akad. Férh. Stockh. 23: 303-336.

LJUNGMAN, A. 1870. Om tvanne nya arter Ophiurider. Ofvers. K. VetenskAkad. Férh. Stockh.
27: 471-475.

Louw, E. 1977. The South African Museum’s Meiring Naude cruises. Part 1. Station data 1975,
1976. Ann. S. Afr. Mus. 72: 147-159.

LyMaAN, T. 1875. Ophiuridae and Astrophytidae of the Hassler Expedition. J//. Cat. Mus. comp.
Zool. Harv. 8: \—34.

LyMAN, T. 1882. Ophiuroidea. Rep. scient. Results Voy. Challenger, Zool. 5: 1-386.

Mapsen, F. J. 1950. The Echinoderms collected by the Atlantide Expedition, 1945-46. 1.
Asteroidea. Atlantide Rep. 1: 167-222.

MORTENSEN, T. 1933. Echinoderms of South Africa (Asteroidea and Ophiuroidea). Vidensk.
Medadr dansk. naturh. Foren. 93: 215-400.

MORTENSEN, T. 1936. Echinoidea and Ophiuroidea. ‘Discovery’ Rep. 12: 199-348.

MorTENSEN, T. 1948. A monograph of the Echinoidea. 4 (2) Clypeastroida. Copenhagen: C. A.
Reitzel.

MORTENSEN, T. 1951. A monograph of the Echinoidea. 5 (2) Spatangoida 2. Copenhagen: C. A.
Reitzel.

SLADEN, W. P. 1879. On the structure of Astrophiura, a new and aberrant genus of Echino-
dermata. Ann. Mag. nat. Hist. (5) 4: 401-415.

SLADEN, W. P. 1889. Asteroidea. Rep. scient. Results Voy. Challenger, Zool. 30: 1-935.

WALSH, J. H. T. 1891. List of deep-sea holothurians collected during seasons 1887-1891, with
descriptions of new species. Natural History notes from H.M. Jnvestigator. J. Asiatic Soc.
Bengal. 60: 197-204.

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

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

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

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

Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: mee pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (figs 8a—b).

Nucula largillierti Philippi, 1861
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.:

Holotyp
SAM-A13535 i 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

(c) Scientific names, but not their vernacular derivatives
e.g. Therocephalia, but therocephalian

Punctuation should be loose, omitting all not strictly necessary

Reference to the author should be expressed in the third person

Roman numerals should be converted to arabic, except when forming part of the title of a
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‘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.

AILSA M. CLARK

THE SOUTH AFRICAN MUSEUM’S
MEIRING NAUDE CRUISES
PART 4

ECHINODERMS

VOLUME 73 PART 7 AUGUST 1977 3 ISSN 0303-2515

iw THE SOUTH AFRICAN
MUSEUM

OF.0 od es ROAR

INSTRUCTIONS TO AUTHORS

1. MATERIAL should be original and not published elsewhere, in whole or in part.
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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. én. 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, ese masses and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4): 1-51.

THIELE, J. 1910. Mollusca: B. Polyplacontions 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 73 ~~ # Band
August 1977 Augustus
Part oy Deel

CRETACEOUS FAUNAS FROM ZULULAND
AND NATAL, SOUTH. AFRICA
THE AMMONITE FAMILY TETRAGONITIDAE
MYATT, [900

By

WILLIAM JAMES KENNEDY
&
HERBERT CHRISTIAN KLINGER

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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Printed in South Africa by In Suid-Afrika gedruk deur
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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

CRETACEOUS FAUNAS FROM ZULULAND AND NATAL,
SOUTH AFRICA
THE AMMONITE FAMILY TETRAGONITIDAE HYATT, 1900

By

WILLIAM JAMES KENNEDY
Geological Collections, University Museum, Oxford

&

HERBERT CHRISTIAN KLINGER
South African Museum, Cape Town

(With 27 figures)

[MS. accepted 29 March 1977]

ABSTRACT

Members of the Tetragonitidae typically form a minor portion of the ammonite fauna of
the South African Cretaceous, being important only in the Upper Campanian and Lower
Maastrichtian. The following species are described: Tetragonites? heterosulcatus Anthula,
Tetragonites subtimotheanus Wiedmann, Tetragonites superstes Van Hoepen, Tetragonites cf.
epigonum (Kossmat), Saghalinites cala (Forbes), Saghalinites nuperus (Van Hoepen), Pseudo-
phyllites indra (Forbes), Pseudophyllites teres (Van Hoepen) and Pseudophyllites latus (Marshall).
The majority of all these forms are widely distributed circum-indic species, but T. ? heterosulcatus
has been previously recorded only from Daghestan, the Caucasus, Bulgaria and Tunisia. The
material allows a revision of the poorly known T. superstes, S. nuperus and P. teres, whilst
T. virgulatus van Hoepen is confirmed as a synonym of P. indra. Ontogenetic series of
T. subtimotheanus are illustrated, as is the ontogeny and variation in S. cala. This latter material
shows a range of ventral ridges and striations associated with the ventral lobe and siphuncle of
unknown function.

CONTENTS
PAGE
Introduction : 2 : ' : ; ‘ . 149
Location of specimens. a E cl? 150
Field localities . : : : : ey ev
Dimensions of specimens ‘ : : ESO
Suture terminology . ‘ , : : : < “AST
Systematic palaeontology . ; : ; ; - ahstl
Stratigraphy ‘ : ; ‘ ; : 40, MES
Acknowledgements . : 4 : ; ; a» 2h93
References . : : . : : : ; av 195
INTRODUCTION

The Tetragonitidae are a small group of ammonites, conservative in external
morphology, but showing a progressive, sexlobate suture line with a formula
ELU,U;=S U,Is. The group evolved from the Gaudryceratidae during the
Aptian via Eogaudryceras (Eotetragonites); they range from the Upper Aptian

149

Ann. S. Afr. Mus. 73 (7), 1977: 149-197, 27 figs.

150 ANNALS OF THE SOUTH AFRICAN MUSEUM

to the Lower Maastrichtian, and have a wide distribution, being locally common
in areas as far apart as Alaska, New Zealand, Japan and Antarctica. They are,
however, rare in the Boreal region; only a few specimens are known from Europe
and Asia north of the Alpine fold belts, and none are recorded from the Western
Interior of the United States. In South Africa, species are known from the Lower/
Middle Albian to Maastrichtian, and are relatively common in the Upper
Campanian and Lower Maastrichtian.

The following species are described below:

Tetragonites? heterosulcatus Anthula

Tetragonites subtimotheanus subtimotheanus Wiedmann

Tetragonites superstes Van Hoepen

Tetragonites cf. epigonum Kossmat

Saghalinites cala (Forbes)

Saghalinites nuperus (Van Hoepen)

Pseudophyllites indra (Forbes)

Pseudophyllites teres (Van Hoepen)

Pseudophyllites latus (Marshall)

LOCATION OF SPECIMENS

The following abbreviations are used to indicate the source of material:

BMNH British Museum (Natural History), London
MHNG Muséum d’Histoire Naturelle, Geneva
EMP Ecole des Mines, Paris

MHNP Muséum d’Histoire Naturelle, Paris

SAS South African Geological Survey, Pretoria
™ Transvaal Museum, Pretoria

DM Durban Museum

UPE University of Pretoria

SAM South African Museum, Cape Town

NMB National Museum, Bloemfontein.

FIELD LOCALITIES

Outline details of field localities referred to in this paper are given by
Kennedy & Klinger (1975); full descriptions of sections are deposited in the
library of the Palaeontology Department of the British Museum (Natural
History) and in the Palaeontology departments of the South African Museum
and the South African Geological Survey.

DIMENSIONS OF SPECIMENS

All dimensions given below are in millimetres:

D = diameter, Wb = whorl breadth, Wh = whorl height, U = umbilicus.
Figures in parentheses are dimensions as a percentage of the total diameter.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 151

SUTURE TERMINOLOGY

The suture terminology of Wedekind (1916, see Kullman & Wiedmann
1970) is followed in the present work:

I; = internal lobe with septal lobe, U = umbilical lobe, L = lateral lobe,
iE — external lobe.

SYSTEMATIC PALAEONTOLOGY

Phylum MOLLUSCA

Class CEPHALOPODA Cuvier, 1797
Subclass AMMONOIDEA Zittel, 1884
Order LYTOCERATIDA Hyatt, 1899
Superfamily |TETRAGONITACEAE Hyatt, 1900
Family Tetragonitidae Hyatt, 1900
Genus Tetragonites Kossmat, 1895

Type species
Ammonites timotheanus Pictet (1848: 295, pl. 2 (fig. 6), pl. 3 (fig. 1)) by
original designation.

Diagnosis

Moderately evolute tetragonitids characterized by a round to subrectangular
whorl section and oblique prorsiradiate constrictions which are typically flexed
over the venter and have a distinct concave ventral sinus. Suture formula
ELU,U;=S Uj,lIs. The external lobe (E) is deeper than the first lateral lobe (L);
there is an irregularly bifid or trifid termination to the first lateral saddle (E/L)
and a nearly symmetrical lateral lobe (L).

Discussion

Tetragonites can be separated from Saghalinites Wright & Matsumoto, 1954
on the basis of the more evolute coiling, lower expansion rate and much simpler
suture of that genus. Pseudophyllites Kossmat, 1895 is much more inflated and
involute, has a much higher expansion rate, a typically rounded, frequently com-
pressed whorl section, no constrictions and a reticulate ornament, whilst the
suture is far more highly subdivided.

The genus Epigoniceras Spath, 1925 with Tetragonites epigonus Kossmat,
1895 as type species was proposed for Lower Turonian to Maastrichtian tetra-
gonitids which differed from earlier Tetragonites sensu stricto in having a
retracted suspensive lobe with a much larger number of auxiliary elements.
Epigoniceras has been accepted by Usher (1952), Wright & Matsumoto (1954),
Wright (1957) and Collignon (1956, 1965a, 19656, 1966, 1969), but Howarth
(1958: 9) has. successfully proved that straight, recurved and even upcurved
suspensive lobes are to be found throughout the time range of the group, so that
Epigoniceras does not bear separation from Tetragonites even on a stratigraphic
basis. Wiedmann (1973: 609) has introduced the genus Carinites, with Tetragonites

152 ANNALS OF THE SOUTH AFRICAN MUSEUM

spathi Fabre (1940: 214, pl. 6 (fig. 1), text-fig. 26) as type species for tetragonitids
with a subrectangular whorl section, persistent, irregularly spaced constrictions
projected on the flanks and converging to a marked ventral peak, and a per-
sistent, if feeble ventral keel. Wiedmann regarded the keel as a feature unknown
in any other tetragonitid species, but similar ‘keels’ of this type are quite wide-
spread in the material here present, occurring for instance in Tetragonites
superstes Van Hoepen (1921: 10, pl. 2 (figs 17—20)). A distinct keel also occurs in
Tetragonites epigonus Kossmat (e.g. Usher 1952, pl. 2 (fig. 7)) and such features
are widely developed in Saghalinites cala (Forbes) (p. 168). This feature is not
a floored keel, rather it is a raised siphonal area associated with the siphuncle.
Rather similar keels are found in many ammonite groups, and appear to be
variable features of specific value only, suggesting that Carinites may possibly
be placed in the synonymy of Tetragonites.

Collignon (1956: 98) listed twenty-eight species and varieties of Tetragonites
and Epigoniceras in his review of the genus, and since his work, Aptian to
Cenomanian species have been described and discussed by Wiedmann (1962a,
19626, 1973), Wiedmann & Dieni (1968) and Murphy (1967a, 1967b) so that
over thirty names are in current usage and there are a number of other named
forms, chiefly based on nuclei, which may be synonyms of better known species,
or which can only be regarded as nomen dubia.

The authors find it difficult to believe that this host of species, many with
overlapping or identical geographic and stratigraphic ranges, is indeed a true ©
picture of the evolution of the genus, but the current material is inadequate for
a satisfactory critical appraisal of the list.

Occurrence

Tetragonites first appears in the Upper Aptian, and ranges to the Maastrich-
tian. The genus has an almost world-wide distribution, with records from
Antarctica, South Africa, Mozambique, New Zealand, Madagascar, Japan,
Sakhalin, Algeria, Tunisia, the western Mediterranean, Spain, southern France,
Switzerland, England, northern France, a variety of localities in central Europe,
and Sinai. In North America there are records from Texas and northern Mexico,
California, Oregon, British Columbia, and Alaska.

Tetragonites? heterosulcatus Anthula, 1899
Figs 1A—F, 2A—B

Lytoceras (Tetragonites) heterosulcatus Anthula, 1899: 99-100, pl. 7 (fig. 4), non pl. 7 (fig. 5).
Pervinquiére, 1907: 73.

Tetragonites heterosulcatus Drushchits, 1956: 102, pl. 7 (figs 27-28), text-fig. 44; 1960: 260,
text-fig. 68. Murphy, 1967a: 32, text-fig. 14. Dimitrova, 1967: 31, pl. 11 (figs 1-1a),
text-fig. 13.

Type
Lectotype herein designated, the original of Anthula’s (1899, pl. 7 (fig. 4))
from the Akusha Shales (Aptian) of Daghestan.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 153

Material

Three specimens, BMNH C78827—C78829, from Bed J of the Mzinene
Formation, locality 171, Mlambongwenya Spruit, northern Zululand (Albian
II-III).

Dimensions

D Wb Wh Wb/Wh U
Lectotype 24,0 13,0(54) 9,0(37) 1,4 8,0(33)
(from Anthula)
BMNH C78827 49,0 24,8(51) 20,8(42) 1,2 10,9(22)
BMNH C78828 — 23,0(—) 19,0(—) 1,2 _
Description

The coiling is fairly evolute, about 40 per cent of the previous whorl being
covered. The whorls expand at a moderate rate, are depressed, and have their
greatest thickness below mid-flank. The umbilicus is of moderate size, 22 per
cent of the diameter, and fairly deep. The umbilical wall is vertical, the umbilical
shoulder abruptly rounded, the flanks flattened and subparallel, the ventrolateral
shoulders abruptly rounded, the venter broad and flattened.

The surface of the shell (Figs 1 A—C, 2B) bears fine growth striae and strong,
flexuous prorsiradiate constrictions, fourteen or fifteen on the outer whorl, each
preceded by a fine, low, rounded collar. The constrictions are narrow and quite
deeply incised; they arise at the umbilical seam, pass straight up the umbilical
wall, sweep forwards across the ventrolateral shoulder and are strongly prorsi-
radiate and markedly convex across the flanks. They flex backwards across the
upper flank and ventrolateral shoulder to form a broad ventral sinus, which
deepens as size increases.

The internal mould (Figs 1D-F, 2A) is smooth save for the constrictions,
which are broader and deeper than on the test.

The suture, rather poorly exposed in the present material (see Drushchits
1956: 102, text-fig. 442), includes a large, asymmetrically bifid first lateral saddle
(E/L), a smaller, bifid second lateral saddle (L/U,), separated by a large bifid
lateral lobe (L). There are three auxiliary saddles on the suspensive lobe (Us),
the first large and subtrifid, whilst there is a long first, and an incipient second
saddle on the internal suture. The septal lobe is horseshoe shaped, and of
moderate size (Fig. 1A).

Discussion

The depressed, rectangular whorl section and remarkable constrictions,
becoming closely spaced and strongly recurved in adults, together with the deep
ventral sinus find an exact match in the large specimens of Tetragonites hetero-
sulcatus figured by Drushchits (1960) and Dimitrova (1967), whilst small speci-

154 ANNALS OF THE SOUTH AFRICAN MUSEUM

D E F

Fig. 1. Tetragonites ? heterosulcatus Anthula. A-C. BMNH C78827. D-F. BMNH C78828.
Both from Bed J of the Mzinene Formation, Albian II-III at locality 171, Mlambongwenya
Spruit, northern Zululand. x 1.

mens figured by Drushchits (1956, 1960) link these with Anthula’s juvenile :
holotype. No other Tetragonites species shows this combination of characters, q
whilst the superficially similar Eogaudryceras (Eotetragonites) such as E. duvalia-
num (d’Orbigny) (e.g. Murphy 1967a, pl. | (figs 2-3)), E. umbilicostriatus Collig- |
non (1963, pl. 248 (fig. 1060)), E. plurisulcatus Breistroffer (= Tetragonites duvali
Anthula (non d’Orbigny) 1899: 99, pl. 7 (fig. 3a—b)), E. wintunius (Anderson)

CRETACEOUS FAUNAS FROM SOUTH AFRICA 155

A B

Fig. 2. Tetragonites ? heterosulcatus Anthula. A. BMNH C78829. B. BMNH 78828. Both
from Bed J of the Mzinene Formation, Albian II-III, at locality 171, Mlambongwenya Spruit,
northern Zululand. x 1.

(1938, pl. 16 (figs 2—-5)). E. shoupi Murphy (19676: 22, pl. 3 (figs 7-9)) and E. jalla-
bertianus (Pictet) (1848: 302, pl. 4 (fig. 2a—b)) have straight or flexuous constric-
tions, never develop a strong ventral sinus, and have markedly different relative
proportions.

The generic affinity of the species is enigmatic. Murphy (1967a, 19676) in
the most recent revision of this species placed it in Tetragonites. Wiedmann (1973)
has subsequently referred the species to Eotetragonites because it lacks an
umbilical lobe; this divergence reflects the difficulties sometimes encountered in
placing forms in the two genera where differing criteria—sutures vs. form of
constriction—are used. In consequence the authors have questioned reference
of this species to Tetragonites until a full review of the Tetragonites/Eotetragonites
plexus is available.

Occurrence

Upper Aptian of Daghestan, the Caucasus and Bulgaria, Aptian of Tunisia,
and Lower/Middle Albian of Zululand.

156 ANNALS OF THE SOUTH AFRICAN MUSEUM

Tetragonites subtimotheanus subtimotheanus Wiedmann, 1962
Figs 3A—C, 4A—-F, 5A-H, 6A-G

Ammonites timotheanus Stoliczka, 1865: 146, pl. 73 (figs 3-4, 6) non 5S.

Lytoceras (Tetragonites) timotheanus Kossmat, 1895: 133, pl. 17 (figs 11, 13). Collignon, 1928:
18, pl. 1 (fig. 18).

Lytoceras timotheanum Anderson, 1902: pl. 7 (figs 145-148).

Tetragonites timotheanus Crick, 1907: 172, pl. 10 (fig. 15-15a), non pl. 13 (fig. 5—Sa)
(= Desmoceras latidorsatum (Michelin)). |

Tetragonites aff. T. timotheanus Imlay, 1960: 100, pl. 12 (figs 24-28).

Tetragonites subtimotheanus Wiedmann, 1962a: 131, 172. Collignon, 1963: 22, pl. 249
(fig. 1071). Pars. Murphy, 1967a: 62, pl. 5 (figs 11, 13), non pl. 6 (figs 5, 8) (= T. blaisoni
Collignon), text-figs 34-35.

Tetragonites rectangularis alaskaensis Murphy, 1967a: 46, pl. 6 (figs 9-10, 14), pl. 7 (figs 2, 9),
text-figs 22-23.

?Tetragonites madagascariensis Murphy, 1967a: 68.

Tetragonites subtimotheanus subtimotheanus Wiedmann, 1973: 592, pl. 1 (fig. 5?), pl. 2 (fig. 2),
pl. 3 (figs 1-5), pl. 7 (fig. 8?), text-fig. 2.

Type
The holotype is the original specimen figured by Kossmat (1895) as pl. 17

(fig. 15), from the lower part of the Utatur Group of southern India. Designated
by Wiedmann (1962a: 171).

Material

Fourteen specimens from the Mzinene Formation of the Mzinene River and
Skoenberg regions: BMNH C78834 from locality 70 (Middle Albian), NMB
D367a—b from Bed 7 at locality 51, SAS H207/8/9 from Bed 8 or 9 at locality 51,
and BMNH C78835 from Bed 9 or 10 at the same locality (Upper Albian),
BMNH C78831 from locality 61 (Albian VI), and C78832—C78833 (Lower-—
Middle Cenomanian) from locality 62, SAS 689, SAS 1034 and BMNH C18142
are probably from the Lower to Middle Cenomanian of the same locality, whilst
a further specimen, SAS 10869, is probably from the Upper Albian of the
Mzinene River. UPE 270-271 and BMNH C78836 are from locality 64 (Albian
V). A juvenile, BMNH C78837, best referred to as T. sp. juv. cf. subtimotheanus,
comes from locality 106 (Albian V).

Dimensions

D Wb Wh Wb/Wh U
Holotype
(after Kossmat 1895) 35,0 19,0(54) 14,0(40) 35 10,0(29)

Indian specimens
(after Murphy 1967a)

MHNG 9
Odium,
larger specimen Sie 31,1(60) = 21,5(41) 1,45 14,6(28)

157

CRETACEOUS FAUNAS FROM SOUTH AFRICA

“eIJOYOMZ] Wakes oy} UO

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Il uviqyy ‘uoNnewi0., sudUIZ| 94) Woy

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‘DeQa/D HNWA ‘UUeUIpaIAA suubaysouilsgns snubaylouljqns sajluospaja[, *¢ “SIA

158

ANNALS OF THE SOUTH AFRICAN MUSEUM

D Wb Wh Wb/Wh UF
MHNG 9
Odium,
smaller specimen | 32,4 18,5(57) 13,1(40) 1,41 10,1(31)
MHNG 9
Odium,
smaller specimen 2 20,3 11,3(56) 8,3(41) 1,36 6,4(32)
Indian specimens
(after Kossmat) a 80,0 45,0(56) 41,0(51) 1,09 16,0(20)
b DEO 14,5(54) 10,5(39) 1,38 8,0(30)
BMNH C18142 24,1 13,5(£56) 10,4(43) 1,29 7,0(29)
BMNH C78832 — 18,8(—) 14,2(—) 1.32 —
BMNH C78833 — 13,2(—) 10,1(—) 1,30 —
BMNH C78834 at S055 23,7(47) 23,1(46) 1,02 13,9(27)
at 68,5 38,5(56) 32,8(48) ey 19/5(8)
at 84,0 45,2(54) 37,5(44) 1,20 PES(2 5)
SAS A1034 29,4 S55) 12,2(41) ean 92GB)
SAS A689 19,5 9,9(51) 1,237) 137 G2ZG1,7)
SAS 1869 64,3 36,2(56) 27,3(42) [32 15,1(23)
UPREQTI 47,5 26,0(55) 22,5(47) Li5 10,0(21) }
Description

The coiling is involute, over two-thirds of the previous whorl being covered.
The whorls are trapezoidal, very depressed when young (whorl breadth to whorl
height ratio is up to 1,37), becoming somewhat rounded and less depressed when
adult (breadth: height ratios of 1,2 to 1,09) and expanding fairly slowly. The
greatest breadth is at the umbilical shoulder. The umbilicus is deep, and of
moderate size, 28-32 per cent of diameter in the middle growth stages, but
becoming proportionally smaller—25 per cent of diameter or less—in larger
specimens. The umbilical wall is flat and subvertical,. the umbilical shoulder
abruptly rounded. The flanks are distinctly flattened and convergent, the ventro-
lateral shoulder abruptly rounded, and the venter broad and flattened.

The shell surface is finely striate. These striae pass straight across the
umbilical wall, running normal to the umbilical seam. They sweep forwards
across the umbilical shoulder, and are strongly prorsiradiate over the flank and
markedly concave. They flex backwards over the ventrolateral shoulder and form
a pronounced sinus over the siphonal area. As size increases, the flexure of the
striae increases and they become very concave on both flanks and venter.

There are five or six collar-like ribs and associated constrictions per whorl
in smaller specimens; adults bear seven to eight, with some crowding on the body
chamber. Collars are faint on the flanks, but strengthen considerably on the

CRETACEOUS FAUNAS FROM SOUTH AFRICA 159

shoulder and venter. They are rounded, with a gentle adapical and steep adaper-
tural slope. The constrictions parallel growth striae; they arise at the umbilical
seam, are deep across umbilical and ventrolateral shoulders, but relatively weak
on flank and venter.

The internal mould is smooth, save for the constrictions, which are far more
prominent than on the test. They arise at the umbilical seam, pass straight up the
umbilical wall and forwards across the umbilical shoulder, where they are
strongly marked. They are concave and strongly prorsiradiate on the flanks and

D E F

Fig. 4. Tetragonites subtimotheanus subtimotheanus Wiedmann. A-—C. BMNH C78834, from

the Mzinene Formation, Albian II, on the farm Izwehelia, north north-east of Hluhluwe,

Zululand. D-F. BMNH C78836, from bed 4 of the Mzinene Formation, Albian V, at locality

71 on the southern tributary of the Munywana Creek, Zululand. This specimen shows a well-
preserved septal lobe. x 1.

160 ANNALS OF THE SOUTH AFRICAN MUSEUM

flex backwards over the shoulder with a striking ventral sinus. The constrictions
become broader and more strongly flexed as size increases.

The adult aperture shows features very similar to the collars associated with
constrictions, being marked by a thickened lip.

None of the present specimens shows the external suture, but the septal faces

G H

Fig. 5. Tetragonites subtimotheanus subtimotheanus Wiedmann. A—C. SAS A10869, from the
Upper Albian Mzinene Formation on the Mzinene River, Zululand. D-F. SAS A1034.
G-H. BMNH C18142 figured by Crick (1907, pl. 10 (fig. 15-15a)). All from the Lower or
Middle Cenomanian Mzinene Formation, Skoenberg, on the Mzinene River, Zululand. x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 161]

of a number of specimens show two large internal saddles on either side of the
dorsal lobe and a prominent septal lobe (Fig. 4F). Some specimens, including
that figured by Crick (1907, pl. 10 (fig. 15-15a)) show, in addition, a tiny third
internal saddle.

Discussion

A series of Tetragonites ranging from Middle Albian to Middle Cenomanian
are linked together by their involute coiling, depressed trapezoidal whorls and
markedly flexuous constrictions, collar ribs and growth striae. Although variable
in their relative proportions and the degree of incision and flexure of constric-
tions, reference to a single species seems acceptable. The specimens match closely
with Tetragonites subtimotheanus subtimotheanus as figured by Stoliczka (1865,
pl. 73 (figs 3-3a, 44a)), Kossmat (1895, pl. 7 (fig. 13-13a)) and Wiedmann
m7: 592, pl. | (fig. 57), pl. 2 (fig. 2), pl. 3 (figs 1-5), pl. 7 (fig. 8 ?)). The latter
author (Wiedmann 1973: 594) discusses fully how TJ. subtimotheanus subtimothea-
nus differs from related forms. T. timotheanus (Pictet 1848: 295, pl. 2 (fig. 6),
pl. 3 (fig. 1)) has a similar juvenile shell, remaining evolute throughout, and loses
its constrictions early in ontogeny. Tetragonites rectangularis Wiedmann (1962a:
178, pl. 14 (fig. 3), text-fig. 39) has a broad, rectangular whorl section at first,
losing its distinct ventro-lateral shoulders at a diameter of around 30 mm, whilst
the constrictions are straighter, less crowded and decline earlier. Tetragonites
kitchini Krenkel (1910: 226, pl. 22 (fig. 8)) has a subrectangular whorl section

Bie: E F G

Fig. 6. Tetragonites subtimotheanus subtimotheanus Wiedmann. A-C, G. BMNH (78832.
D-F. BMNH (C78833. Both from the Lower or Middle Cenomanian Mzinene Forma-
tion, Skoenberg, on the Mzinene River, Zululand. x 1.

162 ANNALS OF THE SOUTH AFRICAN MUSEUM

which persists to greater diameters, and generally five constrictions per whorl.

Tetragonites subtimotheanus subspecies maclearni Wiedmann (1973: 595,
pl. 4 (fig. 1), pl. 5 (figs 1-3), text-fig. 3) has been distinguished from the typical
form on the basis of the development of an oval, rather than broadly rounded
whorl section in adults (compare Wiedmann 1973, text-figs 2a—e, 3a—b).

Occurrence

This species is known from the upper Lower Albian to Lower Cenomanian
of the Queen Charlotte Islands, British Columbia and the Chitina Valley, Alaska,
the Upper Albian of Oregon, the Upper Albian and Lower Cenomanian of
Madagascar and southern India, and the Middle Albian to Lower (and Middle?)
Cenomanian of Zululand.

Tetragonites superstes Van Hoepen, 1921
Figs 7A—D, H—J, 8, 12A—C

Tetragonites superstes Van Hoepen, 1921: 10, pl. 2 (figs 17-20), text-fig. 6. Spath, 1922: 119,
pl. 6 (fig. 6).

Tetragonites cf. epigonum Spath, 19216: 42.

Epigoniceras superstes Collignon, 1956: 87, pl. 11 (fig. 3a—b); 1969: 14, pl. 517 (fig. 2034).

Type

The holotype is TM 564, the original of Van Hoepen (1921, pl. 2 (figs 17—18))
from the Umzamba Formation (Late Santonian to Early Campanian) at locality
1, the mouth of the Umzamba River, Transkei (Pondoland).

Material

Seven specimens, TM 525, TM 564-566, BMNH C19416, SAM-K7029,
SAM-K7096, all from the Umzamba Formation (Late Santonian to Early
Campanian), locality 1, at the mouth of the Umzamba River, Transkei
(Pondoland).

Dimensions
D Wb Wh Wb/ Wh U

Holotype TM 564 40,0 18,2(45) 17,3(43) 1,05 10,4(26)
Paratype TM 566 32,4 15,5(48) 14,2(44) 1,09 8,3(26)
From Collignon 1956: 87

MNHP 2098 44.0 21,0(48) 19,0(43) ia 13,0(30)
MNHP 2099 44.0 21,0(48) 18,0(41) 1,1 13,0(30)
MNHP 2100 52,0 22,0(42) 22,0(42) 1,0 14,0(27)
Description

The shell is small and involute with a moderately high expansion rate. The
whorl section is depressed (whorl breadth: height ratio 1,05 to 1,15) with the

CRETACEOUS FAUNAS FROM SOUTH AFRICA 163

greatest breadth just below mid-flank. The umbilicus is about 26 per cent of the
diameter, rather deep, with a subvertical wall of moderate height. The umbilical
shoulder is abruptly rounded, the flanks gently rounded, merging with broadly
rounded shoulders and venter. The whorl section is thus between subrectangular
and elliptical. The surface of the test bears very fine prorsiradiate growth striae,
whilst the internal mould is smooth. Some individuals show faint spiral ridges,
one over the siphonal area and a pair on each shoulder (Figs 7D, H, 12C).

Constrictions are faint on the mould, and virtually invisible when the shell
is preserved. They appear to be absent on juveniles, but up to four per half-
whorl are present during the later growth stages, as in TM 566. The mature
aperture, present on a number of specimens (Fig. 7A, C) is also constricted.
These apertural constrictions are deep and quite wide, originating at the umbilical
seam. They pass normally across the inner part of the umbilical wall and are
strongly prorsiradiate across the flanks, flexing backwards across the ventro-
lateral shoulders and passing across the venter with a shallow, concave ventral
sulcus.

E F G H | J

Fig. 7. A-D, H-J. Tetragonites superstes Van Hoepen. A-B. BMNH C19416 (original of
Spath 1922, pl. 6 (fig. 6)). C-—D. Holotype, TM 564. H-J. TM 566, a paratype. E-G. Tetra-
gonites cf. epigonus Kossmat. An unregistered specimen in the collections of the Durban
Museum. All specimens are from the Umzamba Formation, of late Santonian to early
Campanian age, at locality 1, the mouth of the Umzamba River, Pondoland. x 1.

164 ANNALS OF THE SOUTH AFRICAN MUSEUM

The suture line (Fig. 8) has a large, asymmetric trifid first lateral saddle
(E/L), a smaller trifid second lateral saddle (L/U,), and a suspensive lobe with
a large trifid first auxiliary saddle. The first lateral lobe (L) is large and irregularly
subdivided. The first of the four auxiliary lobes is trifid.

Discussion

Tetragonites superstes most closely resembles Tetragonites popetensis Yabe
(1903: 48, pl. 7 (figs 4a—b, 6)), a Campanian to Maastrichtian species known
from Japan and California. In Yabe’s species the position of the constrictions
is rather clearly marked on the shell surface by rounded ribs,which are markedly
flexed. The shell is also more evolute (U = 31% at a diameter of 34 mm in the
type).

Differing proportions and number and style of constrictions readily separate
Tetragonites superstes from contemporary species such as Tetragonites glabrus
Jimbo and its variety problematica Matsumoto (1942: 672, figs 1-1b, 2a—2b),
Tetragonites garuda (Forbes) (1846: 102, pl. 7 (fig. la-c)), Tetragonites mitrai-
kyense Collignon (1956: 86, pl. 11 (fig. 2-2b)), Tetragonites beantalyensis Collig-
non (1956: 83, pl. 10 (fig. 1-1b)) and the various species revised by Henderson
(1970). .

Occurrence

All present material comes from the Umzamba Formation of Transkei
(Pondoland), and cannot be dated more accurately than Late Santonian to Early
Campanian. The species is also known in Madagascar where it occurs from
Lower to Middle Campanian.

U, L E

Fig. 8. External suture of Tetragonites superstes Van Hoepen. TM 564. x 10.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 165

Tetragonites cf. epigonus Kossmat, 1895
Figs 7E-G, 9A-—C

Compare:

Ammonites timotheanus Stoliczka, 1865: 146, pl. 73 (fig. 5 only).

? Desmoceras pyrenaicum De Grossouvre, 1894: 168, pl. 25 (fig. 2), non pl. 37
(fig. 9), text-fig. 73.

Lytoceras (Tetragonites) epigonus Kossmat, 1895: 135, pl. 17 (figs 4a—c, 5a—b, 10).
Kilian & Reboul, 1909: 14. |

?Tetragonites timotheanus Whiteaves, 1903: 329.

Tetragonites cf. epigonus Yabe, 1903: 49, pl. 7 (fig. 3).

non Lytoceras (Tetragonites) epigonum Boule, Lemoine & Thévenin, 1906: 13,
pl. 3 (fig. 1-la). This specimen is an Albian desmoceratid. Marshall, 1926:
149, pl. 21 (fig. 10), pl. 29 (figs 6-7) = Tetragonites marshalli Collignon.
Anderson, 1958: 187, pl. 65 (figs 4-5), pl. 67 (fig. 3-3a) = Tetragonites pope-
tensis Yabe.

Lytoceras (Tetragonites) epigonum Pervinquiére, 1907: 76, pl. 3 (figs 27—28), text-
figs 15-16. Basse, 1928: 461, pl. 30 (figs 1-3). Collignon, 1931: 14, pl. 2
(figs 5-7), pl. 3 (fig. 4).

Tetragonites epigonus Paulcke, 1906: 174.

?Tetragonites sp. indet. Spath, 19216: 42, pl. 7 (fig. 3).

? Epigoniceras epigonum Spath, 1925: 29, pl. 1 (fig. 2a—b).

A. B C

Fig. 9. Tetragonites cf. epigonus Kossmat. BMNH C78838, from the St Lucia Formation,
Campanian IV-V, at locality 73, on the lower reaches of the Mzinene River, Zululand. x 1.

166 ANNALS OF THE SOUTH AFRICAN MUSEUM

? Lytoceras (Tetragonites) aff. epigonus Basse, 1939: 45.

Epigoniceras epigonum Matsumoto, 1942: 671. Usher, 1952: 55, pl. 2 (figs 6-7),
pl. 3 (fig. 1), pl. 31 (fig. 13). Collignon, 1956: 85; 1965a: 8, pl. 417 Gagai 23)
1966: 3, pl. 456 (fig. 1855).

Tetragonites jurianus angolanus Haas, 1952: 12-15, figs 21, 23-25 only.

Tetragonites cf. epigonus Howarth, 1958: 9, pl. 1 (fig. 12a—b).

Tetragonites epigonus Matsumoto, 1959: 153, text-fig. 75.

Type
Lectotype (herein designated) is the original of the larger of Kossmat’s
(1895, pl. 17 (fig. 4a—c)) figured specimens.

Material

Two specimens; a dubious juvenile in the collections of the Durban Museum,
figured by Spath (19214, pl. 7 (fig. 3)) from the Umzamba Formation of Late
Santonian or Early Campanian age, locality |, the mouth of the Umzamba River,
Transkei (Pondoland), and a further specimen, BMNH C78838 from the St Lucia
Formation at locality 73 on the lower Mzinene River (Coniacian IV—-V).

Dimensions
D Wb Wh Wb/Wh U

BMNH C78838. Maximum diameter is 71 mm. Proportions at:

65,0 — 31,3(48) — 14,5(22)

S15 28,5(55) 26,0(50) 1,09 12,0(23)
From Kossmat (1895: 135)

59,0 27,0(46) 26,0(44) 1,04 15,0(25)

2320 11,0(44) 10,0(43) al 7,0(30)
Description

The coiling is involute, the whorls expanding at a moderate rate. The whorl
section is slightly depressed, with the greatest breadth close to the umbilical
shoulder. The umbilicus is small (22 % of the diameter) and relatively deep, with
a high, subvertical wall. The umbilical shoulder is abruptly rounded whilst the
sides are somewhat flattened and convergent, the ventro-lateral shoulders distinct
and the rather wide venter broadly rounded.

The shell surface is corroded, and no trace of ornament remains. Traces of
seven constrictions are, however, visible on the outer whorl, each associated with
a low, rounded rib. The constrictions arise at the umbilical seam, pass straight
up the umbilical wall, sweep forwards across the umbilical shoulder and are
markedly prorsiradiate and slightly concave across the flanks; they flex distinctly
backwards over the ventro-lateral shoulders, and there is a broad, shallow ventral
sinus. The sutures are not exposed.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 167

Discussion

The juvenile specimen noted by Spath (1921b; 1922, pl. 4 (fig. la-c)) is
difficult to place with certainty, but the lack of ornament, relative proportions,
umbilical wall and flank shape all match closely with Kossmat’s (1895, pl. 17
(fig. Sa—b)) smaller specimen. The authors’ other specimen, although poorly
preserved, has proportions, whorl section and constrictions, all of which suggest
reference to Kossmat’s species.

Occurrence

Tetragonites epigonus is a long-ranging form known from the Turonian to
the Campanian. Its geographic distribution includes Antarctica, Angola, Mada-
gascar, Japan, southern India, British Columbia, south Patagonia, north Africa
and southern France. In South Africa the Pondoland occurrence can be dated
no more firmly than Late Santonian to Early Campanian. The Zululand example
is firmly dated as Coniacian IV-V.

Genus Saghalinites Wright & Matsumoto, 1954
Type species
Ammonites cala (Forbes, 1846: 104, pl. 8 (fig. 4a—c)) by original designation
of Wright & Matsumoto (1954: 110).

Diagnosis

Very evolute tetragonitids with a low expansion rate, the whorls being
slightly depressed. The whorl section is typically rounded when young, becoming
distinctly octagonal during later growth stages in most species. The shell surface
is smooth, or bears only growth striae. Constrictions are consistently present,
and may be weak to strong, straight or sinuous. The suture is relatively simple,
with trifid major saddles, and a retracted suspensive lobe.

Discussion

The name Saghalinites was introduced by Shimizu in 1934 as a nomen
nudum; validation extends from the work of Wright & Matsumoto in 1954.
Originally proposed as a subgenus of Epigoniceras Spath, 1925, the work of
Howarth (1958), Matsumoto (1959) and Wiedmann (1962a, 19626, 1973) sug-
gests that Epigoniceras does not bear separation from Tetragonites sensu Stricto,
as noted elsewhere (p. 151). Howarth and Matsumoto have treated Saghalinites
as a subgenus of Tetragonites, whilst Birkelund (1965) and Wiedmann (1962a),
amongst others, have given it full generic status. Wiedmann (1973: 589) has
subsequently suggested that Saghalinites is no more than a synonym of Tetra-
gonites. In the authors’ views, however, the features of the type species and other
well-known Santonian to Maastrichtian species referred to the genus indicate a
distinct monophyletic offshoot from contemporaneous Tetragonites which merits
generic separation; the group appears as distinctive as the bulk of the genera of
Tetragonitaceae.

168 ANNALS OF THE SOUTH AFRICAN MUSEUM

Saghalinites is readily separable from Pseudophyllites Kossmat, 1895, in that
that genus is inflated, very involute, and typically has a rounded whorl, higher
than wide, and is ornamented by longitudinal and transverse striae, whilst the
suture line is highly subdivided and there are no constrictions.

Tetragonites Kossmat, 1895, is also typically more inflated, with a higher
expansion rate, and more involute coiling.

The origin of Saghalinites clearly lies in Tetragonites, from which it evolved
in the Late Santonian (or possibly the Coniacian, according to Collignon (1956:
82)).

The following species and varieties have been referred to the genus:
1. Saghalinites cala (Forbes) (1846: 104, pl. 8 (fig. 4a-c)). Campanian to
Maastrichtian.

2. Saghalinites nuperus (Van Hoepen) (1921: 13, text-fig. 8, pl. 3 (figs 3-4)).
Santonian to Lower Campanian.

3. Saghalinites zeugitanus (Pervinquiére) (= Tetragonites cala var. zeugitana Per-
vinquiére, 1907: 79, pl. 3 (fig. 3a—b)). ? Santonian.

4. Saghalinites zelandicus Shimizu, 1935 (= Gaudryceras politissimum Marshall
(non Kossmat), 1926: 143, pl. 20 (fig. 3), pl. 28 (figs 1-2), but is, in fact, a crushed
Anagaudryceras particostatum (Marshall); fide Henderson 1970).

5. Saghalinites kingianus (Kossmat, 1895) (= Ammonites cala Stoliczka gs
Forbes), 1865: 153, (pars) pl. 75 (fig. 4)). ? Santonian to Campanian.

6. Saghalinites kingianus (Kossmat) var. involutor Paulcke (1906: 174, pl. 17
(figs 3-4)). Campanian.

7. Saghalinites wrighti Birkelund (1965: 30, pl. 1 (fig. 5), pl. 2 (figs la—c, 5a-c),
pl. 3 (fig. 1), text-figs 14-25). Maastrichtian.

Occurrence

Saghalinites first appears in the Santonian and ranges to the Lower Maas-
trichtian. Species have a wide geographic distribution; there are records from
Antarctica, south Patagonia, Zululand, Madagascar, South Africa, Japan,
Sakhalin, southern India and Greenland.

Saghalinites cala (Forbes, 1846)
Figs 1OA-B, 11A-B, 12D-—G, 13A-B, E-K, ? C_-D, 14A—-F, 15A—-F

Ammonites cala Forbes, 1846: 104, pl. 8 (fig. 4a—c). Non Kossmat, 1895: 153, pl. 75 (fig. 4)
= Saghalinites kingianus (Kossmat).

Lytoceras (Tetragonites) cala Kossmat, 1895: 136, pl. 17 (fig. 12a—d).

non Tetragonites cf. cala Anderson, 1902: 84. Yabe, 1915: 16, pl. 1 (fig. 7), pl. 3 (fig. 2)
= Saghalinites nuperus Van Hoepen.

?Tetragonites aff. cala Woods, 1906: 335, pl. 41 (fig. 7a—c).

Tetragonites cala (Forbes) var. zeugitana Pervinquiére, 1907: 79, pl. 3 (fig. 30), text-fig. 18.

non Tetragonites cala Kilian, 1922: 176.

Tetragonites cala Shimizu, 1935: 181.

Saghalinites cala Spath, 1953: 9. Collignon, 1956: 99.

Tetragonites (Saghalinites) cala Howarth, 1958: 10, pl. 1 (fig. 11a—b).

CRETACEOUS FAUNAS FROM SOUTH AFRICA 169

Type
The lectotype is Forbes’s original figured specimen, BMNH C51057, from
the Valudayur Beds (Campanian—Maastrichtian) of Pondicherry, southern India.

Material

The authors have numerous specimens, as follows: SAS H163/E1, E2 and
E6, H163/3, H163B/2, 4 and 5, H163C/2 and 12, H163G/1, all from locality 20
south of Lake Mfuthululu, east-south-east of Mtubatuba (Maastrichtian [-II);
BMNH C78840-43 from locality 113 at the south-eastern corner of the Nibela
Peninsula, Lake St Lucia (Campanian IV); SAS Z2267—2267e from this locality
and to the area to the immediate west; SAS H104/1 from locality 117 at the
north-eastern tip of the southern Peninsula (Campanian IV); SAS Z2242 and
2248a—b, H119/11, H115/10 and 11 from locality 119; BMNH C78862 and
C78863 from locality 120; BMNH C78861 from locality 121, The Coves, on the
eastern shores of the southern Peninsula (Campanian IIJ-IV); BMNH C18857—60
from locality 124 north of Fanies Island Rest Camp on the eastern shores of the
southern Peninsula (Campanian IIJ-IV); BMNH C78856 from locality 126,
south of the Camp (Maastrichtian IT); BMNH C78853—C78855 from locality

a al
ee Te oe Mh

A B

Fig. 10. Saghalinites cala (Forbes). BMNH CS51058, the original of Kossmat (1895, pl. 17
(fig. 12a—d)), from the Valudayur Beds (Campanian—Maastrichtian) of Pondicherry, southern
Indian 1.

170 ANNALS OF THE SOUTH AFRICAN MUSEUM

132; BMNH C78844 and SAS specimens A1252, 2087, H60F/6, 60H/6 and
16217 from locality 133; BMNH C78845—C78852, and SAS specimens H63/2,
10, 13-14, 22-25, H61/1, 2, 5, 7, 9, 12; 18, H64/1-4, 6; 9, and 2221022) sirens
locality 134 in the area of Charters Creek Rest Camp, Lake St Lucia (Maastrich-
tian 1); and SAS H66/1 from locality 135, Makakatana Bay, Lake St Lucia
(Maastrichtian I). There is a single juvenile specimen, SAM-—4808, from the
Umzamba Formation at locality 1, the mouth of the Umzamba River, Transkei
(Pondoland) (Late Santonian to early Campanian) which may belong here.

Dimensions

D Wb Wh Wb/Wh U
Lectotype
BMNH C51057 S253 13,7(42) SIG) 1,38 14,4(45)
BMNH C51058 78,8 30 (38) 23,9(30) e2> 37,4(47)
SAS H60/12 66,8 — 20,5(30) — 31,2(46)

A B

Fig. 11. Saghalinites cala (Forbes). SAS H163b/2, from the St Lucia Formation, Maastrichtian

I-II, at locality 20, south of lake Mfuthululu, east south-east of Mtubatuba, Zululand. The

specimen shows coarse lateral folds associated with constrictions, a distinctly sulcate venter,
and spiral ridges on both flank and venter. x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA

jai

Lectotype D Wb Wh Wb/Wh U
SAS H61/11 76,5 29,2(38) 21,9(28) 1538 36,1(47)
SAS H163/D 89,0 38,5(43) DS 2311) 1,36 40,5(45)
BMNH C78855 at 28,2 Iie tGC9) 9,4(33) 1218 12,4(44)

at 63,0 DSO) 18,5(29) 125 30,0(47)
BMNH C78843 91,8 — 29,132) - 42,0(46)
BMNH C78863 oS = 31,8(32) - 46,4(46)
SAS Z2270k 60,5 DASG5) 18,5(31) 1,16 28,5(47)
SAS A1214 88,5 31,5(36) 28,0(32) 112 43,0(49)
Description

The bulk of the material occurs either as composite or internal moulds.

Juveniles, up to 30 mm diameter (Figs 12E-F, 13A—D, G-K)

The coiling is evolute, slowly expanding, the whorl section depressed and
rounded, the whorl breadth : height ratio being around 1,2, the greatest breadth
some distance below mid-flank. The umbilicus is broad, 44 per cent of the
diameter, shallow, with a low, rounded, undercut wall. The flanks are rounded,
and there is a broad, rounded venter. The surface of the mould is smooth, with
six prominent, narrow constrictions. The constrictions arise at the umbilical
seam, pass straight up the umbilical wall, and straight across the inner flanks in
a markedly prorsiradiate direction, flexing gently backwards across the upper
flank and shoulder to form a shallow concave sinus over the siphonal area.

Middle growth stages, 30 to 60 mm diameter (Figs 12D, 14A—F)

The coiling remains evolute, but the whorl section becomes more depressed,
the whorl breadth : height ratio being up to 1,25 and the whorl section changing
from depressed oval to polygonal.

The umbilical wall increases in height, is flat, and inclined outwards, the
umbilical shoulder is abruptly rounded, the whorl sides flattened and convergent,
the ventrolateral shoulder abruptly rounded, the venter, broadly rounded at
first, becomes flattened, and in some cases concave. Constrictions become
increasingly flexed, sweeping forwards over the inner flanks, flexing backwards
at mid-flank and developing a broad, shallow but distinct ventral sinus. The
number of constrictions increases, with up to eight per whorl in some specimens.

Many specimens show a range of structures associated with the siphonal
band (Fig. 14A, D), as discussed below.

60 mm Onwards (Figs 11A—B, 12G)

The largest specimens present are still incomplete at 100mm and are the
largest known representatives of the genus. Up to two-thirds of the outer whorl
is body chamber in these specimens. The whorl section is polygonal, as in middle
growth stages, but there is an increase in the degree of ventral concavity, and

eye ANNALS OF THE SOUTH AFRICAN MUSEUM

many specimens develop a low, but distinct, rounded siphonal ridge and related
structures.

Constrictions become stronger, deeper, closer spaced and more markedly
flexed on body chambers, and there is a tendency for the lower part of the flank
between constrictions to become irregularly swollen (Fig. 11A—B). In other
specimens, broad, flexuous folds are present on the flanks, and seem to represent
an exaggerated development of this feature. Yet other specimens show low,
rounded, spiral ridges on their flanks (Fig. 13E-F).

The few external moulds of the outer shell surface available suggest that it

D G

Fig. 12. A-C. Tetragonites superstes Van Hoepen. TM 7029, from the late Santonian—early
Campanian Umzamba Formation at locality 1, the mouth of the Umzamba River, Pondoland.
D-G. Saghalinites cala (Forbes). D. A silicone mould taken from BMNH C78841, from the
St Lucia Formation, Campanian IV, at the south-eastern corner of the Nibela Peninsula, Lake
St Lucia, Zululand. E-F. BMNH C78856, from the St Lucia Formation, Maastrichtian II,
at locality 126, south of Fanies Island Rest Camp, on the eastern shores of the southern
Peninsula, Lake St Lucia, Zululand. G. BMNH C78863, from the St Lucia Formation,
Campanian III-IV, locality 120, The Coves, on the eastern side of the southern Peninsula,
Lake St Lucia, Zululand. A-—D, G, x 1; E-F, x 2.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 173

bore faint growth striae, parallel to the constrictions, whilst the constrictions are
themselves rather less conspicuous on the shell exterior, being associated with a
low, collar-like rib (Fig. 12D).

The external suture line (Figs 14A—C, 15) is rather simple, with a large first
lateral saddle (E/L), and a smaller second lateral saddle (L/U,), both of which
are irregularly trifid. The suspensive lobe is strongly retracted, the auxiliary lobes
and saddles decreasing rapidly in size. The lateral lobe (L) is markedly bifid.

The internal suture has a deep dorsal lobe and a narrower lateral lobe

G H J K

Fig. 13. Saghalinites cala (Forbes). A-B, G-K. BMNH C78855, from the St Lucia Formation,
Maastrichtian I, at locality 132, near Charter’s Creek Rest Camp, Lake St Lucia, Zululand.
E-F. BMNH C78851, from the St Lucia Formation, Maastrichtian I, near Charter’s Creek
Rest Camp, Lake St Lucia, Zululand. The specimen is an internal mould of a body chamber
lacking constrictions, and showing lateral and ventral spiral ridges. C-D. SAM-—4808, best
referred to as Saghalinites cf. cala (Forbes) ?, from the late Santonian to early Campanian
Umzamba Formation at locality 1, the mouth of the Umzamba River, Pondoland. A-B, x 2;
C-K, x 1.

174 ANNALS OF THE SOUTH AFRICAN MUSEUM

extending to about half the depth. The first saddle is tall and narrow, the second
much smaller (Fig. 14B, F).

Discussion

The large number of specimens available show very variable adult features,
especially the nature and development of constrictions and lateral folds on the
body chamber, the degree of concavity of the venter and the extent of the
siphonal band. In spite of this, the authors feel confident in referring their
specimens to Forbes’ species on the basis of the comparable and distinctive
ontogenetic changes, the markedly similar polygonal whorl section of adults,
distinctly sigmoidal constrictions and overall comparable relative proportions.

Comparable variability in whorl section and venter form has been
described in Saghalinites wrighti from the Maastrichtian of west Greenland
(Birkelund 1965: 30 et seq., especially text-figs 14-20). S. cala and S. wrighti
can, however, be differentiated on the basis of the absence of constrictions
on the early whorls of S. wrighti, the fewer constrictions on the later whorls,
and their straight rather than flexuous course.

S. cala can readily be separated from both S. nuperus and the doubtful
S. kingianus on the basis of its sinuous rather than straight constrictions, and
the development of a polygonal whorl section during later growth stages rather
than the rounded section retained by these forms.

Pervinquiére’s Lytoceras (Tetragonites) cala var. zeugitana (1907: 79, pl. 3.
(fig. 30a—b)) is based on a specimen only 13 mm in diameter and was separated
from the typical form on the basis of the more angular, trapezoidal whorl section
and more flexuous constrictions. It seems doubtful if it indeed merits separation
in view of the great variability as described above, but since it is said to be of
Santonian age, it may conceivably be the juvenile of some other species.

As already noted, many of the present specimens show beautifully ventral
structures resembling those described by Grandjean (1910: 502-503), Neaverson
(1927), Hélder (1955), Vogel (1959: 510-511), Jordan (1968: 28) and Birkelund
(1965: 36).

On internal moulds of body chambers (Figs 11A, 13E—F), the chief structure
is a low, flat-topped ridge, marked off on either side by a distinct narrow groove
and extending from the aperture to the last septum. When intersected by con-
strictions, the ridge is weakened, but nevertheless continuous across the constric-
tion. It thus appears that this structure corresponds to the presence of a pair of
parallel ridges on the shell interior. On the phragmocone during later growth
stages (Fig. 14A, D), this ridge is usually subdued, or its site marked by a band
corresponding in width to the ventral lobe. The band is bisected by a continuous
median groove, and the surface is covered by fine, longitudinal striae which
converge slightly when traced in an apical direction from one suture to the next.
In some cases, concave transverse striae are present in the area enclosed by the
siphonal lobe, giving rise to a distinctive reticulate pattern (Schleppstreifen of
Holder 1955: 374), whilst the ventral band in some juveniles is ornamented by

CRETACEOUS FAUNAS FROM SOUTH AFRICA M75

a curious chevron-like striation. The interpretation of these structures is far from
clear; Jordan (1968) suggested that the ventral ridge represents the trace of a
muscle system controlling a pre-septal gas and fluid-filled space, but it is difficult
to reconcile this with the extension of the band throughout the length of the
body chamber. The striations of the siphonal band on the phragmocone are
typically interpreted as the site of muscle or ligament insertion associated with

E

Fig. 14. Saghalinites cala (Forbes). BMNH C78855, from the St Lucia Formation, Maastrich-
tian I, at locality 132 near Charter’s Creek Rest Camp, Lake St Lucia, Zululand. A and D
show details of ventral structures; B and F the internal suture. A-D, x 2; E-F, x 1].

176 ANNALS OF THE SOUTH AFRICAN MUSEUM

L E L

Ay

a A
7 : :
KA
5 ;

Fig. 15. External and internal sutures of Saghalinites cala (Forbes). A-E. BMNH C7854S5.
F. BMNH 78844. All x 6.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 77,

the attachment of the siphuncle to the shell, but again there is no convincing
evidence for this.

Occurrence

Saghalinites cala is common in Zululand, ranging from the Upper Cam-
panian (Campanian IV) to the Lower Maastrichtian (Maastrichtian II). The
doubtful Pondoland occurrence cannot be dated more precisely than Late
Santonian to Early Campanian. The type, from southern India, is of Santonian
or Campanian age, whilst there are also records from the Campanian of Antarc-
tica and the ?Santonian of Tunisia.

Saghalinites nuperus (Van Hoepen, 1921)
Figs 16A—E, 17A-B, 18

Tetragonites cf. cala Yabe, 1915: 16, pl. 1 (fig. 7), pl. 3 (fig. 2).

Tetragonites nuperus Van Hoepen, 1921: 13, pl. 3 (figs 3-4), text-fig. 8. Besairie, 1930: 224,
pl. 21 (fig. 3-3a).

Tetragonites cala Basse, 1931: 17, pl. 1 (figs 27-28), pl. 10 (fig. 7).

Saghalinites nuperus Collignon, 1956: 95, pl. 11 (fig. la—b); 1966: 3, pl. 456 (fig. 1856); 21,
pl. 463 (fig. 1893).

Type
The holotype is TM 532, the original of Van Hoepen (1921, pl. 3 (figs 3-4))
by original designation.

Material

In addition to the holotype, the authors have seen three paratypes, TM 536
and 539, both from the Umzamba Formation of Late Santonian to Early
Campanian age, locality 1, the mouth of the Umzamba River, Transkei (Pondo-
land), and a specimen, BMNH C78839, from locality 84, False Bay, Zululand
(Santonian I).

a = B C D E

Fig. 16. Saghalinites nuperus (Van Hoepen). A-B. The holotype, TM 532. C-—E. Paratype,
TM 535. Both specimens are from the Umzamba Formation, of late Santonian to early
Campanian age at locality 1, the mouth of the Umzamba River, Pondoland. x 1.

178

ANNALS OF THE SOUTH AFRICAN MUSEUM

Dimensions

D Wb Wh Wb/Wh U
Holotype 37,6 15,0(40) 14,0(37) 11 15,5(41)
Paratype TM 539 USD} 10,2(40) D0G>) als 10,6(42)
Paratype TM 536 15,0 6,0(40) 5,6(37) lie 6,2(41)
(crushed)
BMNH C78839 92,0 42,0(46) 35,8(39) ay) 32,5(35)
From Collignon 1955: 95
MHNP 2173 53,0 21,0(40) 19,0(36) Pl 21,0(40)
MHNP 2177 530) 25,0(42) 21,0(36) ee 27,0(46)
MHNP 2179 1220 31,0(43) D065) 1,24 30,0(42)
Description

Early whorls, 10-30 mm

The shell is of medium size, evolute, only 25 per cent of the previous whorl
being covered, slowly expanding, with a wide umbilicus (ca. 40% of diameter).
The whorl section is rounded at first in the smallest paratypes, but is depressed
throughout later growth, becoming somewhat less depressed as diameter
increases; the greatest breadth is a little below mid-flank. The umbilicus is
shallow, with a low subvyertical wall which merges into a rounded shoulder,
which in turn grades imperceptibly into the rounded convergent flanks. The
venter is broadly rounded.

The test is ornamented by very fine, dense striae which arise at the umbilical
seam, run at first normal to the seam but then sweep strongly forwards over the
shoulder and are gently convex and strongly prorsiradiate on the flanks. They
sweep gently back across the ventro-lateral shoulder to form a gentle concave
ventral sinus. The internal mould is smooth. Four to five constrictions per whorl
are present, and occur from a diameter of 5 mm onwards, although initially
rather faint. They are rather narrow, and follow a course parallel to the growth
striae. On the test, their site is marked by a faint collar.

Adults

The larger specimen, 98,5 mm in diameter, is the largest described individual
referred to this species and appears to be adult. The coiling is moderately
involute, with depressed whorls and a moderately high expansion rate, a rela-
tively deep umbilicus equal to 35 per cent of the diameter, with a high, subvertical
wall. The umbilical shoulder is fairly abruptly rounded, the flanks gently inflated,
convergent, with the greatest breadth some way below mid-flank. The ventro-
lateral shoulders are rounded, merging imperceptibly with a fairly broad, rounded
venter. The test is ornamented by fine, dense striae, of rather variable strength.
These arise at the umbilical seam, run normally across the inner part of the
umbilical wall, but sweep strongly forwards on the shoulder. They are straight
and strongly prorsiradiate on the inner flank. weakly convex at mid-flank, flexed

CRETACEOUS FAUNAS FROM SOUTH AFRICA 179

gently backwards across the shoulder and pass across the venter with a slight
convex peak. There are five well-developed constrictions on the last half-whorl,
which run parallel to the growth striae. On the mould they are relatively deeply
incised at the umbilical shoulder, and are narrow and sharply demarcated on
flank and venter. They are rather less prominent on the test.

The suture line (Fig. 18) is rather simple, with a large, trifid first lateral
saddle (E/L) and a smaller, virtually identical second lateral saddle (L/U,). The
lateral lobe (L) is bifid, the first auxiliary lobe trifid. The suspensive lobe is
retracted with several auxiliaries.

Discussion

The holotype and paratypes of this species are rather small, but the larger
individuals figured by Collignon (1956, 1966) and the present adult specimen

Fig. 17. Saghalinites nuperus (Van Hoepen). BMNH C78839, from the St Lucia Formation,
Santonian I, locality 84, False Bay, St Lucia, Zululand. x 1.

180 ANNALS OF THE SOUTH AFRICAN MUSEUM

show that this species retains a rounded whorl section throughout ontogeny,
whilst constrictions become more closely spaced as diameter increases.

Saghalinites nuperus is thus readily separated from S. cala and S. wrighti,
both these species developing a polygonal whorl section at large diameters. The
constrictions of S. cala are, in addition, markedly flexed with a concave ventral
sinus, rather than the peaked constrictions of the adult S. nuperus.

Saghalinites kingianus is a difficult species to interpret, being based upon a
composite drawing taken from more than one specimen. It appears, however, to
lack constrictions to a diameter of 40-50 mm, and when they do appear they
are strongly prorsiradiate and straight rather than gently curved as in S. nuperus.
Until further material is described and adequately figured, S. kingianus is perhaps
best regarded as a nomen dubium.

Paulcke’s Saghalinites kingianum var. involutor (1906: 175, pl. 17 (figs 3—-4)) is
based on two juveniles having the following dimensions:

D Wb Wh Wb|Wh U
Example | 11,0 9,0(81) 6,0(54) 1,5 3,5(31)
Example 2 22,0 17,0(77) 12,0(55) 1,42 O62)

It thus differs markedly from S. nuperus in relative proportions and, like
the typical form, lacks constrictions at this size.

Occurrence

The type material from Pondoland is from an unknown horizon within the
Umzamba Formation of Late Santonian to Early Campanian age. Detailed
collecting by one of the authors (H.C.K.) at the type section yielded one fragment
of S. nuperus in the uppermost Santonian just below the Santonian/Campanian
boundary. The species is also recorded from the Lower and Middle Santonian
and possibly the Upper Santonian/Lower Campanian of Madagascar, and is
known from the ‘Senonian’ of Japan.

Genus Pseudophyllites Kossmat, 1895
Type species

Ammonites indra Forbes, 1846 by original designation.

Diagnosis

Tetragonitids with moderately involute whorls when young, expanding to
become very involute when adult. Early whorls depressed, with greatest breadth
close to mid-flank, later whorls becoming rounded and varying from slightly
compressed to slightly depressed. No constrictions; surface of test ornamented
by fine transverse growth lines and spiral striae which combine to produce a
typical reticulate pattern. Suture very finely divided with asymmetrically trifid or

CRETACEOUS FAUNAS FROM SOUTH AFRICA 181

asymmetrically bifid major saddles having subphylloid terminations; suspensive
lobe retracted.

Discussion

Pseudophyllites is readily separated from Saghalinites Wright & Matsumoto,
1954 in that that genus is very evolute, has a low expansion rate, a rounded to
polygonal whorl section, a simpler suture line and prominent constrictions.
Tetragonites Kossmat, 1895 typically has a rounded to squarish whorl section,
a simpler suture, and generally bears striking constrictions throughout ontogeny.

The origin of Pseudophyllites clearly lies in Tetragonites, from which it
evolved in the Late Santonian. Collignon (1956) lists six species which have been
referred to that genus, and a further species, Pseudophyllites skoui Birkelund
(1965: 37, pl. 3 (figs 2-6), text-figs 26-33), has since been added. Species are
separated chiefly upon details of whorl section and suture line.

Occurrence

Pseudophyllites species are best known from the Campanian and Maastrich-
tian, the geographic distribution including Antarctica, South Africa, Madagas-
car, southern India, New Zealand, northern Australia, Japan, Sakhalin, southern
and central Europe, west Greenland, Alaska, British Columbia, California and
Brazil. The genus is also said to occur in the upper Santonian of Madagascar
(Collignon 1956).

Fig. 18. Saghalinites nuperus (Van Hoepen). External suture of TM 532.
x T0:

182 ANNALS OF THE SOUTH AFRICAN MUSEUM

Pseudophyllites indra (Forbes, 1846)
Figs 19A—F, 20-22

Ammonites indra Forbes, 1846: 105, pl. 11 (fig. 7). Stoliczka, 1865: 112, pl. 58 (fig. 2a—b).
Whiteaves, 1879: 105, pl. 13 (fig. 2—2a).

?Gaudryceras colloti De Grossouvre, 1894: 229, pl. 27 (fig. 8a—b).

Pseudophyllites indra Kossmat, 1895: 137, pl. 16 (figs 6-9), pl. 17 (figs 6-7), pl. 18 (fig. 3).
Whiteaves, 1903: 331. Woods, 1906: 334, pl. 41 (fig. 6a—b). ?Non Kilian & Reboul, 1909:
14, text-fig. 3 = ?P. latus. Spath, 1921b: table opposite page 50; 1922: 119 (pars).
Non Marshall, 1926: 152, pl. 20 (fig. 1), pl. 29 (figs 3-5) = P. latus. Nagao & Saito,
1934: 359, text-fig. 10. Collignon, 1938: 24, text-fig. E. Usher, 1952: 57, pl. 3 (figs 2-13),
pl. 21 (fig. 17). Collignon, 1956: 90. Matsumoto, 1959: 134. Jones, 1963: 25, pl. 7
(figs 6-7), pl. 8, pl. 29 (figs 7-12), text-fig. 10. Collignon, 1969: 12, pl. 516 (fig. 2032).

Lytoceras indra Boule, Lemoine & Thévenin, 1906: 2, pl. 1 (fig. 1-1b).

Tetragonites virgulatus Van Hoepen, 1921: 11, pl. 3 (figs 1-2), text-fig. 7.

?Pseudophyllites amphitrite Maury, 1930: 167, pl. 27 (fig. 1), pl. 28 (fig. 1).

Parapachydiscus catarinae Anderson & Hanna, 1935: 19 (pars), pl. 3 (figs 2-3).

Pseudophyllites aff. indra Spath, 1940: 43.

Type

Lectotype herein designated, BMNH C51068, figured by Forbes (1846,
pl. 11 (fig. 7)) from the Valudayur Group near Pondicherry, southern India.

Material

Six specimens, BMNH C19417 and C19418, TM 531 and 526 (the latter two
being the types of Tetragonites virgulatus Van Hoepen) from the Late Santonian
to Early Campanian, Umzamba Formation at locality 1, the mouth of the
Umzamba River, Transkei (Pondoland), SAS H126/2 from locality 106 at the
mouth of the Nyalazi River (Campanian I), and SAS H150, from locality 16,
south of Mtubatuba (age uncertain).

Dimensions
D Wb Wh Wb/Wh U
Lectotype
BMNH C51068 at 109,3 — 61,5(56) = 19,4(17)
at 94,5 44,5(47) 50,0(53) 0,89 15,5(16)
C19417 296,0 148,5(50) 171(57) 0,87 42,0(14)
C19418 Oss — 35,10) — 13,018)
H126/2 at Bh3)55) 53,0(54) SES) 1,02 18,6(18)
at TAGS) 40,3(52) 43,8(56) 0,92 14,3(18)
H150 109,0 60,0(55) 60,0(55) 1,0 =
Description

The coiling is very involute, rapidly expanding, typically with a compressed
whorl section. The umbilicus is small and deep, conical, with a flat, outwards
sloping wall. The umbilical shoulder is abruptly rounded, with initially rather
flattened, convergent flanks, and an arched, rounded venter. The test is orna-
mented by fine striae and ridges which arise at the umbilical seam, sweep slightly

CRETACEOUS FAUNAS FROM SOUTH AFRICA 183

backwards across the umbilical wall, flex backwards over the shoulder, pass
across the flanks in a prorsiradiate direction and may be faintly convex. They
sweep backwards across the ventro-lateral shoulder, and cross the venter with a
broad, faint, concave ventral sinus. In addition, there are faint, closely spaced
spiral striae which combine with the transverse ornament to produce a reticulate
pattern on the shell surface.

Internal moulds are smooth, or may bear faint traces of the reticulate orna-
ment. The suture line (Fig. 22) is highly subdivided, with a rather variably sub-

C D E F

Fig. 19. Pseudophyllites indra (Forbes). A-B. BMNH C19418. C-E. TM 431, the holotype of

Tetragonites virgulatus Van Hoepen. F. TM 526, a paratype of T. virgulatus. All specimens

are from the late Santonian to early Campanian Umzamba Formation at locality 1, the mouth
of the Umzamba River, Pondoland. x 1.

184 ANNALS OF THE SOUTH AFRICAN MUSEUM

& -~ os
: #

Fig. 20. Pseudophyllites indra (Forbes). BMNH C19417, from the Umzamba Formation, of
late Santonian to early Campanian age at locality 1, the mouth of the Umzamba River,
Pondoland. Reduced x 0,5. (British Museum photograph.)

CRETACEOUS FAUNAS FROM SOUTH AFRICA 185

Fig. 21. Pseudophyllites indra (Forbes). BMNH C19417, from the Umzamba Formation, of
late Santonian to early Campanian age at locality 1, the mouth of the Umzamba River,
Pondoland. Reduced x 0,5. (British Museum photograph.)

186 ANNALS OF THE SOUTH AFRICAN MUSEUM

divided ventral saddle (E) which is often lanceolate in broad outline, a large,
irregularly trifid first lateral saddle (E/L), a smaller bifid, second lateral saddle
(L/U,), a deeply incised bifid lateral lobe (L) deeper than the ventral lobe (E),
and a retracted suspensive lobe with a large bifid first auxiliary saddle. Saddle
terminations are typically subphylloid. The septal face shows two lateral saddles
on either side of the internal lobe, and there is a massive septal lobe, well dis-
played on several of the present specimens.

Discussion

This classic species is characterized by a high expansion rate and rapid
increase in whorl height, flattened flanks, fairly narrow venter, and a small,
conical umbilicus in which the umbilical wall slopes outward to an abruptly
rounded umbilical shoulder.

Pseudophyllites latus (Marshall) (1926: 152, pl. 20 (fig. 1), pl. 29 (figs 3-5)),
of which Pseudophyllites whangaroaensis (Marshall) (1926: 153, pl. 20 (fig. 2),
pl. 21 (fig. 11), pl. 32 (figs 5—6)), Pseudophyllites peregrinus Spath (1953: 7, pl. 1
(figs 6-9)) and Pseudophyllites skoui Birkelund (1965: 37, pl. 3 (figs 2-6), text-
figs 26-33) are synonyms, is a species known from the Campanian to Maastrich-
tian of New Zealand, Antarctica, Madagascar and Greenland. From Henderson’s
(1970: 12 et seq.) recent discussion this form has a generally broader venter than
P. indra, but shows identical ontogenetic changes and style of shell ornament.
The sutures are said to differ, however, the ventral saddle of P. indra being ©
lanceolate, that of P. Jatus being spatulate. The umbilical walls also differ, that
of P. indra sloping distinctly outwards, that of P. /atus being subvertical.

P. teres (Van Hoepen), a species based chiefly on small specimens, has been
separated from P. indra on the basis of the compressed, flattened and subparallel
flanks and consequently subrectangular whorl section, whilst the umbilical wall
is subvertical.

Gaudryceras colloti De Grossouvre (1894: 229, pl. 37 (fig. 8a—b)) from the
Upper Campanian of the Basses-Pyrenées, in southern France, is based upon a
specimen just under 60 mm in diameter, with the following proportions:

D Wb Wh Wb|Wh U
58,0 —-27,0(46) ~—-29,,5(51) 0,92 10,5(18)

It is preserved as a composite mould, and is deformed. The ornament is of
Pseudophyllites type, and it is best regarded as a synonym of Pseudophyllites indra.

Ammonites postremus Redtenbacher (1873: 115, pl. 26 (fig. 3a—d)) is based
upon a series of specimens, the figure showing what may be a Pseudophyllites.
It has a subrectangular whorl section and vertical umbilical wall; both features
clearly separate it from P. indra.

Tetragonites virgulatus Van Hoepen (1921: 11, pl. 3 (figs 1-2), text-fig. 7) is
based on juvenile P. indra; the types are figured here as Figure 19C—E.

Pseudophyllites amphitrite Maury (1930: 167, pl. 27 (fig. 1), pl. 28 (fig. 1))
is based upon a specimen having the following dimensions:

CRETACEOUS FAUNAS FROM SOUTH AFRICA 187

D Wb Wh Wb|Wh U
BIO 7565) 120(51) 0,63 45(21)

The ornament matches that of P. indra, whilst there is a comparable ventral
profile. The whorls are highly compressed, however, but this appears to be due
to post-mortem crushing; the species is also best considered a synonym of P. indra.
Pseudophyllites nereidideditus Maury (1930: 169, pl. 29 (fig. 1)) appears to be a
crushed Pseudophyllites teres, as discussed below. The umbilicus and relative
proportions readily separates it from P. indra.

Occurrence

Pseudophyllites indra ranges from the late Santonian to early Maastrichtian.
Its geographic distribution includes South Africa (Zululand and Pondoland),
Madagascar, southern India, northern Australia, Japan, Sakhalin, Alaska,
British Columbia, California and possibly south-eastern France and Brazil.

U, L E

Fig. 22. External suture of Pseudophyllites indra (Forbes). TM 531. x 5.

Pseudophyllites teres (Van Hoepen, 1920)
Figs 23A—B, 24A-B

Tetragonites teres Van Hoepen, 1920: 144, pl. 25 (figs 1-2).

Pseudophyllites indra Spath, 1922: 119 (pars).

?Pseudophyllites nereidideditus Maury, 1930: 169, pl. 29 (fig. 1).

Pseudophyllites teres Collignon, 1956: 94, pl. 9 (fig. 2-2b); 1969: 14, pl. 517 (fig. 2034).

Type
The holotype is Van Hoepen’s original specimen, TM 562, by original
designation.

188

Material

ANNALS OF THE SOUTH AFRICAN MUSEUM

Two specimens, the holotype and BMNH C19415, both from the late
Santonian to early Campanian Umzamba Formation at locality 1, the mouth of
the Umzamba River, Pondoland.

Dimensions
D Wb Wh Wb/Wh U

Holotype
TM 562 (a) 69 — 390) — 14(20)

(5) 57 26(46) 31(54) 0,84 —
From Collignon (1956: 94)
MHNP 2160 51,0 25,0(49) 26,0(51) 0,96 eOQ2)
MHNP 2163 78,0 39,0(50) 42,0(54) 0,93 16,0(21)
MHNP 2164 98,0 43,0(44) 52,0(53) 0,83 19,0(19)
MHNP 2166 130 54,0(47) 63,0(55) 0,86 21,0(18)
Description

The coiling is involute, rapidly expanding, and compressed (whorl breadth
to whorl height ratio is less than 0,96, decreasing with age). The umbilicus is
small (20 % of diameter), deep, with a high, subvertical wall and abruptly rounded

A B

Fig. 23. Pseudophyllites teres (Van Hoepen). TM 535, the holotype from the Umzamba Forma-
tion, of late Santonian to early Campanian age, locality 1, the mouth of the Umzamba River,
Pondoland. x 1.

189

CRETACEOUS FAUNAS FROM SOUTH AFRICA

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Ajre9 O} URIUOJURG 9k] JO ‘UONBWIOY eqUIeZWIY, 94} WO ‘SI6b1D HNWa ‘(usds0x ueA) sasaz sajyjdydopnasg ‘pZ ‘314

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190 ANNALS OF THE SOUTH AFRICAN MUSEUM

shoulder. The greatest breadth is at the umbilical margin, the inner flanks being
flattened and subparallel, with broadly rounded shoulders and venter. Ornament
is not well preserved, but consists of fine, dense growth striae which pass back
across the umbilical wall, sweep forwards over the shoulder and are strongly
prorsiradiate and faintly convex on the flank, passing backwards across the
shoulder and running almost normally across the venter.

There is a marked apertural constriction. The suture line is poorly visible,
but of the Pseudophyllites type.

Discussion

The largest specimens the authors have seen of this species are adult at
approximately 100 mm diameter, and at this size the compressed whorls with
flattened, subparallel sides, plus the umbilicus with subvertical wall, clearly
differentiates this species from Pseudophyllites indra, P. latus and P. postrematus.
Pseudophyllites nereidideditus Maury (1930: 196, pl. 29 (fig. 1)) appears to be a
large crushed example possibly referable to this species. The dimensions given
by Maury are as follows:

D Wb Wh Wb/Wh U
210,0 70,0(33) 1IS0G5) 0,61 45,0(21)
Occurrence

Santonian—Campanian of Pondoland (Umzamba Formation) and
Madagascar. ? Maastrichtian of Brazil.

Pseudophyllites latus (Marshall, 1926)
Figs 25-26

Pseudophyllites indra Kilian & Reboul, 1909: 14. Marshall, 1926: 152, pl. 20 (fig. 1), pl. 29
(figs 3-S).

Tetragonites latus Marshall, 1926: 149, pl. 20 (fig. 6), pl. 32 (figs 1-2).

Pseudophyllites whangaroaensis Marshall, 1926: 153, pl. 20 (fig. 2), pl. 21 (fig. 11), pl. 32
(figs 5-6).

Pseudophyllites peregrinus Spath, 1953: 7, pl. 1 (figs 6-9). Collignon, 1956: 92, text-fig. 12.

Pseudophyllites latus Henderson, 1970: 12, pl. 1 (fig. 10), pl. 2 (fig. 3), text-fig. 4a—c.

Pseudophyllites skoui Birkelund, 1965: 37, pl. 3 (figs 2-6), text-figs 26-33.

Type

The lectotype, designated by Henderson (1970: 14) is the original of Marshall
(1926, pl. 32 (fig. 1)), from the Mata Series (Campanian) of New Zealand.

Material

One specimen only, SAS Z1114, from locality 106 at the mouth of the
Nyalazi River (Campanian I).

CRETACEOUS FAUNAS FROM SOUTH AFRICA

Dimensions
D

SAS Z1114 172,0
From Collignon (1956: 93)
MHNP 2152 80,0
MHNP 2154 90,0
MAAINP 2155: 127,0
MIENP 2157 + =6181,0

Wb
96,4(56)

44,0(55)
51,0(57)
73,0(57)

113,0(62)

Wh
94,8(55)

40,0(50)
48.0(53)
69,0(54)

101,0(56)

Wb|Wh
1,02

test
ite
1,05
1,1

19]

U
28,5(17)

16,0(20)
16,0(18)
24,0(19)
33,0(18)

Fig. 25. Pseudophyllites latus (Marshall). SAS Z1114, from the St Lucia Formation, Campanian

I, locality 106, at the mouth of the Nyalazi River, Zululand. x 0,8.

192 ANNALS OF THE SOUTH AFRICAN MUSEUM

Description

The specimen is wholly septate, and retains much of the original aragonitic
shell. The coiling is very involute, more than four-fifths of the previous whorl
being covered, with a slightly depressed whorl section throughout ontogeny.
The whorls expand rapidly, the greatest breadth being at the umbilical shoulder.
The umbilicus is deep, conical and narrow (17% of diameter), with a high, flat,

Fig. 26. Pseudophyllites latus (Marshall). SAS Z1114, from the St Lucia Formation, Campanian
I, locality 106, at the mouth of the Nyalazi River, Zululand. The specimen shows the reticulate
ornament clearly, and has the septal lobe well preserved. x 0.8.

i gee
<_+ ie

:
é

CRETACEOUS FAUNAS FROM SOUTH AFRICA 193

subvertical wall. The umbilical shoulder is abruptly rounded, the flanks broadly
rounded and convergent, merging with a broadly rounded venter. Ornament
consists of fine transverse growth striae and longitudinal ridges which combine
to produce a reticulate pattern which is particularly conspicuous on the venter.
The transverse striae arise at the umbilical seam, pass straight up the umbilical
wall with a shallow concavity, are slightly prorsiradiate and weakly convex across
the flanks, and sweep backwards across the ventrolateral shoulders to form a
shallow, broad ventral sinus.

The suture consists of a broad, short, moderately subdivided spatulate
ventral saddle, a massive, highly subdivided bifid first lateral saddle (E/L) and
a smaller bifid second lateral saddle (L/U,) separated by a symmetrical bifid
lateral lobe (L) which is deeper than the external lobe (E). The suspensive lobe
includes five or six auxiliary lobes; there are two internal saddles and a massive
septal lobe (Fig. 26).

Discussion

P. latus is readily separated from P. indra on the basis of its broader whorls,
less rapidly increasing height, and much steeper umbilical wall. It differs from
P. teres in being broader and having convergent rather than subparallel flanks.
The spatulate rather than lanceolate ventral saddle is also distinctive. P. post-
rematus has a distinctly rectangular whorl section, as noted above.

The authors agree with Henderson (1970) in regarding Pseudophyllites
peregrinus Spath, P. skoui Birkelund and P. whangaroaensis Marshall as syno-
nyms of P. latus.

Occurrence

Campanian of New Zealand and Antarctica, Upper Santonian and Cam-
panian of Madagascar, Maastrichtian of Greenland, and Campanian of Zululand.

STRATIGRAPHY

The stratigraphic distribution in south-eastern Africa of the species
described herein is illustrated in Figure 27.

ACKNOWLEDGEMENTS

The authors are grateful to Dr H. W. Ball, Dr M. K. Howarth and Mr D.
Phillips of the British Museum (Natural History), Dr M. Cooper of Wolfson
College (Oxford), General M. Collignon (Moirans), Mr C. W. Wright (London),
Professor Dr J. Wiedmann (Tiibingen), Mr P. J. Rossouw, Dr C. K. Brain and
Mrs E. Voigt (Pretoria), and the technical staff of the Department of Geology,
Oxford University, Geologisch-Palaontologisches Institut (Tiibingen), South
African Geological Survey (Pretoria), Transvaal Museum (Pretoria), and South
African Museum (Cape Town) for their help and assistance in many ways.
Thanks are due to the Director, Geological Survey of South Africa for loan of
part of the material described.

ANNALS OF THE SOUTH AFRICAN MUSEUM

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CRETACEOUS FAUNAS FROM SOUTH AFRICA 195

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Haas, O. 1952. Some Albian desmoceratid and lytoceratid ammonites from Angola. Am.
Mus. Novit. 1561: 1-17.

HENDERSON, R. A. 1970. Ammonoidea from the Mata Series (Santonian—Maastrichtian) of
New Zealand. Palaeontology Spec. Pap. 6: 1-81.

Ho per, H. 1955. Uber die Sipho-anheftung bei Ammoniten. Neues Jb. Geol. Paldont. Mh.
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Howarth, M. K. 1958. Upper Jurassic and Cretaceous ammonite faunas of Alexander Land
and Graham Land. Scient. Rep. Falkld. Isl. Dep. Surv. 21: 1-16.

IMLAY, R. W. 1960. Early Cretaceous (Albian) Ammonites from Chitina Valley and Talkeena
Mountains, Alaska. Prof. Pap. U.S. geol. Surv. 354D: 87-114.

JONES, D. L. 1963. Upper Cretaceous (Campanian and Maastrichtian) ammonites from southern
Alaska. Prof. Pap. U.S. geol. Surv. 432: 1-53.

JORDAN, R. 1968. Zur Anatomie Mesozoischer Ammoniten nach den Strukturelementen der
Gehause-innenwand. Beih. geol. Jb. 77: 1-64.

KENNEDY, W. J. & KLINGER, H. C. 1975. Cretaceous faunas from Zululand and Natal, South
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KILIAN, W. 1922. Note sur une faune d’Ammonites de Nouvelle-Zelandie decouverte par
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KILIAN, W. & REBOUL, P. 1909. Les céphalopodes néocrétacés des iles Seymour et Snow Hill.
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KRENKEL, E. 1910. Die untere Kreide von Deutsch Ostafrika. Beitr. Paldont. Geol. Ost-Ung.
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Paleont. Contrib. Univ. Kans. 44: 1-32.

MARSHALL, P. 1926. The Upper Cretaceous ammonites of New Zealand. Trans. N.Z. Inst. 56:
129-210.

Matsumoto, T. 1942. A note on the Japanese ammonoid species belonging to the Tetragoni-
tidae. Proc. Imp. Acad. Tokyo 18: 671-673.

Matsumoto, T. 1959. Upper Cretaceous ammonites of California. Part 2. Mem. Fac. Sci.
Kyushu Univ. (D) spec. vol. 1: 1-172.

Maury, C. J. 1930. O cretaceo da Parahyba do norte. Monogr. Serv. geol. min. Brasil 8: 1-305.

Murpny, M. A. 1967a. The Aptian—Cenomanian members of the ammonite genus Tetragonites.
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Murpny, M. A. 19676. Aptian and Albian Tetragonitidae (Ammonoidea) from Northern
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NAGAO, T. & SaiTo, R. 1934. Peculiar septal features observed in ammonites of certain Lyto-
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NEAVERSON, E. 1927. The attachment of the ammonite-siphuncle. Proc. Lpool geol. Soc. 14:
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PAULCKE, W. 1906. Die Cephalopoden der oberen Kreide Siidpatagoniens. Ber. naturf. Ges.
Freiburg i.B. 15: 167-248.

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secondaires. Mem. Carte géol. Tunis. 1907: 1-438.

PicTET, F. J. 1848. Description des Mollusques fossiles qui se trouvent dans les Gres Verts
des environs de Géneve. 1. Céphalopodes. Mem. Soc. Phys. Hist. nat. Geneve 11: 257-412.

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Alpen. Abh. geol. Bundesanst. Wien 5: 91-140.

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and Palaeontology. Tokyo. (In Japanese.)

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(2) 2: 159-226.

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SPATH, L. F. 19216. On Upper Cretaceous Ammonoidea from Pondoland. Ann. Durban Mus.
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SPATH, L. F. 1922. On the Senonian Ammonite fauna of Pondoland. Trans. R. Soc. S. Afr. 10:
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SpaTuH, L. F. 1925. On Senonian Ammonoidea from Jamaica. Geol. Mag. 62: 28-32.

SpATH, L. F. 1940. On Upper Cretaceous (Maastrichtian) Ammonoidea from Western
Australia. J. Proc. R. Soc. West. Aust. 26: 41-57.

SpaTH, L. F. 1953. The Upper Cretaceous Cephalopod fauna of Grahamland. Scient. Rep.
Falkld. Isl. Dep. Surv. 3: 1-60.

STOLICZKA, F. 1863-1866. The fossil Cephalopoda of the Cretaceous rocks of southern India.
Palaeont. Indica (3) 1: 41-56 (1863); 2-5: 57-106 (1864); 6-9: 107-154 (1865); 10-13:
155-216 (1866).

Usuer, J. L. 1952. Ammonite faunas of the Upper Cretaceous of Vancouver Island, British
Columbia. Bull. Geol. Surv. Can. 21: 1-182.

VAN HOoePEN, E. C. N. 1920. Description of some Cretaceous ammonites from Pondoland.
Ann. Transy. Mus. 7: 142-147.

VAN HOoePEN, E. C. N. 1921. Cretaceous Cephalopoda from Pondoland. Amn. Transy. Mus. 8:
1-48.

VoGEL, K. P. 1959. Zwergwuchs bei Polyptychiten (Ammonoidea). Geol. Jahrb. 76: 469-540.

WEDEKIND, R. 1916. Uber Lobus, Suturallobus und Inzision. Zentb/. Geol. Paldont. 1916:
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WHITEAVES, J. F. 1903. On some additional fossils from the Vancouver Cretaceous with a
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WIEDMANN, J. 19625. Unterkreide-ammoniten von Mallorca 1. Liefr. Lytoceratina, Aptychi.
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WIEDMANN, J. 1973. The Albian and Cenomanian Tetragonitidae (Cretaceous Ammonoidea)
with special reference to the circum-indic species. Eclog. geol. Helv. 66: 585-616.

WIEDMANN, J. & DIENI, I. 1968. Die Kreide Sardiniens und ihre Cephalopoden. Palaeontogr.
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Woops, H. 1906. The Cretaceous fauna of Pondoland. Amn. S. Afr. Mus. 4: 275-350.

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YABE, H. 1903. Cretaceous Cephalopoda from Hokkaido. Part 1. J. Coll. Sci. Tokyo 18: 1-55.

YABE, H. 1915. Notes on some Cretaceous fossils from Anaga on the Island of Awaji and
Toyajo in the Province of Kii. Sci. Rep. Tokohu Univ. Geol. (2) 4: 13-24.

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Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (figs 8a—b).

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WILLIAM JAMES KENNEDY &
HERBERT CHRISTIAN KLINGER

CRETACEOUS FAUNAS FROM ZULULAND
AND NATAL, SOUTH AFRICA

THE AMMONITE FAMILY TETRAGONITIDAE
HYATT, 1900

Be. or

VOLUME 73 PART 8 AUGUST 1977 ISSN 0303-2515

ey THE SOUTH AFRICAN |
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BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.

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

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

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

Konn, A. J. 1960b. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4): 1-S1.

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 73 ~~ Band
August 1977 Augustus
Part 8 Deel

NOTES ON THE SCORPION FAUNA OF THE CAPE
PART 2
THE PARABUTHUS CAPENSIS (EHRENBERG)
SPECIES-GROUP; REMARKS ON TAXONOMY AND
BIONOMICS (ARACHNIDA, SCORPIONIDA, BUTHIDAE)

By

E. B. EASTWOOD

Cape Town Kaapstad

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NOTES ON THE SCORPION FAUNA OF THE CAPE
PART 2

THE PARABUTHUS CAPENSIS (EHRENBERG) SPECIES-GROUP;
REMARKS ON TAXONOMY AND BIONOMICS
(ARACHNIDA, SCORPIONIDA, BUTHIDAE)

By

E. B. EASTWOOD
South African Museum, Cape Town

(With 7 figures and 1 table)

[MS. accepted 31 March 1977]

ABSTRACT

Parabuthus neglectus Purcell was synonymized with P. capensis (Ehrenberg) by Hewitt in
1918. A re-examination of the types of these two species and material in the South African
Museum collection revealed significant morphological and geographical differences. P. capensis
is redescribed and P. neglectus is reinstated and redescribed. Distribution, morphology and
habitat of these two species are discussed, with notes on a black form of P. capensis.

CONTENTS

PAGE

Introduction . : : . ; : , : i l99
Redescription

Parabuthus capensis (Ehrenberg, 1828) . . 200

Parabuthus neglectus Purcell, 1899 . : ~ 205

Discussion . : ; ‘ : ; : ; 2 e207

Material examined : : : ' : ’ >, wi2

Acknowledgements. : 4 : ‘ . 214

References . : : : : : : : . 204

INTRODUCTION

Androctonus capensis was originally described by Ehrenberg (1828), not
Hemprich and Ehrenberg as is recorded in most literature. The original descrip-
tion is to be found in an account of the travels of Hemprich and Ehrenberg,
hence the mistake made by subsequent workers in referring the description to
both these people.

Purcell (1902) redescribed P. capensis from specimens in the South African
Museum collection. Pocock (1902) compared the type of P. planicauda (Pocock)
with Purcell’s capensis and found them to be identical. He suggested that Purcell
(1899) had redescribed P. capensis as P. neglectus and P. planicauda (Pocock)
as P. capensis (Purcell 1902). Hewitt (1918) synonymized P. neglectus with
P. capensis.

199

Ann. S. Afr. Mus. 73 (8), 1977: 199-214, 7 figs, 1 table.

a a e”6~—SCO

200 ANNALS OF THE SOUTH AFRICAN MUSEUM

In this paper P. capensis is redescribed with notes on its distribution and
habitat, and a black form which is morphologically identical is discussed.
P. neglectus is reinstated and redescribed, since the original description was
inadequate, with notes on its distribution, morphology and habitat.

REDESCRIPTION

Parabuthus capensis (Ehrenberg, 1828)

HOLOTYPE FEMALE (Fig. 1)
Measurements are given in Table 1.

Colour

In the alcohol-preserved specimen the tergites, carapace and telson are dark
reddish-brown, the cauda, sternites and legs yellowish-brown.

TABLE |

Measurements in millimetres of Parabuthus capensis (Ehrenberg) and P. neglectus Purcell.

P. capensis P. neglectus ‘P. neglectus
Holotype ° Lectotype 3 Paralectotype 2 9
Total length A ; : 73,6 79,0 88,0—100,0
Carapace length . i : 5 8,6 6,0 6,5-8,2
width (max.) . : 9,0 6,8 7,0-9,5
width (min.) : Sp) 4,0 3,5-6,0
Pedipalps
Femur length . d : 6,2 4,5 5,0-6,0
Patella length . ‘ 5 6,8 4,8 5,2-7,0
Chela length ; : : 11,0 9,8 9,2-13,0
width : ; : 3,0 2,0 2,0-3,0
Movable finger length . 7,9 5,0 6,1-8,2
Mesosoma length . : : 18,2 1s) 19,2-21,0
Metasoma length : : é 46,8 32.0 38,0—-52,0
Caudal segment I
length . ' : : 6,0 5,0 -4,0-7,0
width . : : 5,8 4,0 4,0-6,0
Caudal segment II
length . : : Us2 555 4,8-7,5
width . : ‘ . 6,0 4,2 4,2-6,2
Caudal segment III
length . : A Tz a5 4,8-8,0
width . ‘ ; 6,0 4,2 4,2-6,0
Caudal segment IV
length . : 5 8,0 6,0 5,0-7,8
width . : : ; 6,0 4,0 4,5-6,0
Caudal segment V
length . : ; : 9,0 6,5 6,0-8,8

WIGS Ton siamese 5 6,0 3,6 4,0-5,0

201

NOTES ON THE SCORPION FAUNA OF THE CAPE

“(dINQUOIYA) SisuadDd snyinqosévg JO MOIA [eIUIA ‘G

‘oyeuoy odAVO[OY
‘gyeuiay odAjojoy “(s19quoi1yq) sisuadvd snyinqvivg jO MdIA |[eSIOG ‘ek ‘[ ‘3Iy

202 ANNALS OF THE SOUTH AFRICAN MUSEUM

Carapace

Slightly wider than long; anterior—posterior divergence 0,47; densely and
finely granular throughout.

Tergites

Densely granular throughout, the granules becoming coarser posteriorly in
each segment; median keel smooth, extending two-thirds the length of segments
I to VI posteriorly; two pairs of strongly granular lateral keels on segment VII;
a few short microtrichia on the lateral margins of segments I to VI.

Sternites

The last four segments smooth and polished; the last segment with sparse,
rounded granules laterally; rows of microtrichia laterally and posteriorly on each
segment.

Cauda

Segment I sparsely granular; ventral and ventrolateral keels smooth, lateral,
dorsolateral and dorsal keels granular; dorsal stridulatory area extending the
whole length and almost three-quarters of the breadth of the segment, with finely
granular ridges laterally.

Segment II granular, ventral keels consist of long contiguous granules,
terminal granule slightly enlarged, rounded; dorsal stridulatory area medially
depressed; the posterodorsal edge of the segment is elevated and slightly curved
forward medially, forming a subtriangular ‘lip’.

Segment IV granular, dorsal keel granular, posteriorly continuous with the
dorsolateral keel; accessory keel consists of 4-6 blunt granules; ventral keels
posteriorly obsolete; dorsal surface medially depressed along its whole length,
lightly shagreened medially with finely granular ridges laterally.

Segment V granular; the single ventral keel is posteriorly obsolete; well-
developed ventrolaterals; dorsal keels well developed anteriorly; accessory keels
represented by two or three low, rounded larger granules; dorsally the segment
is smooth. Vesicle coarsely and sparsely granular.

The entire cauda is very sparsely setiferous.

Pedipalps

Femur finely granular, with a few larger granules internally; anterodorsal,
anteroventral and posterodorsal keels distinctly granular. Patella finely granular;
keels distinctly granular. Chela slender, movable finger with 12 inner flanking
teeth; the median series consisting of 8-10 teeth, including the external lateral
teeth of the outer series of each row; the distal end with 3 enlarged teeth.

Trichobothriotaxy
See Figure 4.

NOTES ON THE SCORPION FAUNA OF THE CAPE 203

Pectines

30/30 teeth; the basal lamellae greatly enlarged and subrectangular in shape.

Operculum
Subcordiform; fused medially.

TYPE MATERIAL

Holotype female

ZMB 133. Cape of Good Hope, Lichtenstein legit., deposited at the Zoo-
logisches Museum, Berlin, DDR.

Parabuthus neglectus Purcell, 1899

LECTOTYPE MALE (Fig. 2)
Measurements are given in Table 1.

Colour

In alcohol-preserved specimens the colour is yellow, the mesosoma and
metasoma yellowish-brown.

Carapace

Wider than long; anterior—posterior divergence 0,5; densely and finely
granular throughout.

Tergites

Densely and finely granular throughout, with coarser granulation on each
segment posteriorly; median keel represented by a granular elevation in the
middle of the first six segments; two pairs of strongly granular lateral keels on
the seventh segment; fairly long microtrichia on the lateral and posterior margins
of all segments.

Sternites

The last four segments smooth and polished; the last segment with sparse,
rounded granules laterally; sparse rows of microtrichia laterally and posteriorly
on all segments.

Cauda

Segment I sparsely granular; ventral and ventrolateral keels smooth; lateral,
dorsolateral and dorsal keels well developed and granular; posterior granule of
dorsal keel not enlarged; dorsal surface with a slightly concave shagreened area,
extending the whole length and almost three-quarters of the breadth of the seg-
ment, laterally with finely granular ridges.

Segment II sparsely granular; ventral and ventrolateral keels consist of

ANNALS OF THE SOUTH AFRICAN MUSEUM

204

“aye 9d AjO1DOf “[OdIN SMJIa/saul SNYINGDADT JO MIA [eIUSA *Q “sfeUT 9dA4O}99] “[[OdINg SHJIa/BaU SNYINGV.1D_ JO MIA [eSIOG *B “7 “BIA

205

NOTES ON THE SCORPION FAUNA OF THE CAPE

“SJUSLUSOS [BPN OA} JSE PUB UOS|d} SY} JO UONVIOJOS JOyxIVp dy} SUIMOYsS puLjenbeweN UsoYyJIOU WOI s[eUIOy
‘[9OAN, SNJIAPGIU SNYINGDADG JO MIA |CSIOG ‘q “a]BUIAay “(S1aquory_) sisuvadvs snyinqoavg JO WUIOJ YORIQ 9Y} JO MOIA [SIO “ev *¢ ‘BIA

206 ANNALS OF THE SOUTH AFRICAN MUSEUM

separate well-defined and prominent granules, increasing in size posteriorly;
other keels not as well defined; dorsal surface with the shagreened area forming
a deep depression; the posterodorsal edge of the segment is strongly elevated and
curved forward medially, forming a very distinctly subtriangular ‘lip’.

Segment II granular; keels well developed and granular; ventral and ventro-
lateral keels very prominent; the posterior granule of the dorsal keel slightly
enlarged; the dorsal shagreened area forming a deep oval depression anteriorly ;
finely granular ridges laterally; the postercdorsal edge of the segment is curved
forward medially to form a well-developed subtriangular ‘lip’.

Segment IV granular; keels granular, not prominent; posterior granule of
dorsal keel slightly enlarged; dorsal surface medially depressed along its whole
length, lightly shagreened medially with finely granular ridges laterally.

Segment V granular; the single ventral keel distinct; well-developed ventro-
laterals; dorsal keels well developed anteriorly, accessory keels represented by
three or four sharply pointed spines; dorsally the segment is smooth.

Vesicle granular; ventrally some of the granules are very prominent. The
metasoma is covered with long setae ventrally and laterally.

Pedipalps

Femur finely granular, with a few larger granules internally; anterodorsal,
anteroventral and posterodorsal keels distinctly granular. Patella finely granular;
keels distinctly granular. Chela slender, movable finger with 10-inner flanking
teeth; the median series consisting of 6-10 teeth, including the external lateral
teeth of the outer series of each row; the distal end with 3 enlarged teeth.

Trichobothriotaxy
See Figure 5.

Pectines
38/38 teeth; the basal lamellae not enlarged.

Operculum
Subcordiform; cleft longitudinally, with a pair of genital papillae below the
operculum.

PARALECTOTYPE FEMALES

Measurements given in Table 1. Morphologically the same as the holotype
except for the following:

Pectines
34-36 teeth, with basal lamellae greatly enlarged and subrectangular in
shape.

Operculum
Subcordiform; fused medially.

NOTES ON THE SCORPION FAUNA OF THE CAPE

TYPE MATERIAL
A lectotype was chosen from the syntypes designated by Purcell (1899).

Lectotype male

SAM-—1197/1. Between Pakhuisberg and Oorlogskloof, Clanwilliam district,
Cape. (32°0'S 19°15’E) M. Schlechter legit. 1897.

Paralectotypes

SAM-1197/2. 3 3 5, 11 2 2. Data as for lectotype. All type material
deposited at the South African Museum.

DISCUSSION

The P. capensis group is distinguished from other species by having the
postero-dorsal edge of caudal segments II and III curved forward medially,
forming a subtriangular ‘lip’.

P. capensis and P. neglectus are separable by the following combination of
characters: the structure of the accessory keels of caudal segment V, the structure
of the ventral and ventrolateral keels, and posterior dorsal lip of caudal segments
II and III, the density and length of meso- and metasomal setae, and number of
pectinal teeth. The distribution of trichobothria on the pedipalpal segments is
orthobothriotaxic for both species (Figs 4-5), with little variation in position.
These two species may be separated by the differences in characters set out in
the key provided below.

KEY TO THE PARABUTHUS CAPENSIS SPECIES-GROUP

1. Granules of accessory keels of caudal segment V low and rounded; ventral and ventrolateral
keels of caudal segments II and III consist of low, almost contiguous granules; posterior
dorsal lip of caudal segments II and III not well developed; meso- and metasomal setae
sparse and short; pectinal teeth 33-37 for males, 31-35 for females

capensis (Ehrenberg)

— Granules of accessory keels of caudal segment V elongated and sharply pointed; ventral
and ventrolateral keels of caudal segments II and JII separate and prominent; posterior
dorsal lip of caudal segments II and III well developed, extending well forward; meso- and
metasomal setae long and fairly dense; pectinal teeth 36-38 for males, 35-36 for females

neglectus Purcell

P. capensis is found in the south-western corner of the Cape in the winter-
rainfall region, extending from Clanwilliam in the north to Cape Town in the
south and eastwards to Tulbagh. In the north (Clanwilliam and Laaiplek), a
form is found which is entirely black (see Fig. 3a). This form cannot be morpho-
logically distinguished from the yellow form of P. capensis and ecological data
give no indication that it is a distinct species. Specimens of the yellow form have
been found under rocks or other debris in areas of hard-packed soil. The black
forms from Laaiplek were found under rubbish on coastal sand dunes, while the
specimen from Clanwilliam was found under a stone in a shallow burrow in an

208 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 4. Trichobothriotaxy of Parabuthus capensis (Ehrenberg). a. External aspect of male chela.
b. External aspect of female chela. c. Ventral aspect of female chela. d. Dorsal aspect of patella.
e. External aspect of patella. f. Dorsal aspect of femur.

209

NOTES ON THE SCORPION FAUNA OF THE CAPE

2mm

e® POO SO gRengr?’

Fig. 5. Trichobothriotaxy of Parabuthus neglectus Purcell. a. External aspect of male chela.
b. External aspect of female chela. c. Ventral aspect of female chela. d. Dorsal aspect of
patella. e. External aspect of patella. f. Dorsal aspect of femur.

210 ANNALS OF THE SOUTH AFRICAN MUSEUM

area of hard-packed soil. The author has found P. capensis in association with
Opisthophthalmus pallidipes Koch, O. capensis (Herbst) and Uroplectes
carinatus (Pocock).

A laboratory specimen of P. capensis has been observed to excavate a run
beneath ground cover as follows: the tail was curved either sideways or directly
over the mesosoma so that the telson lay above the carapace, and the first two
pairs of walking legs were used in alternative scraping motions to loosen the sand.
The body was then raised by stilting on the hind legs while the first two pairs of
legs and sometimes the third pair scraped the sand very vigorously so that it
sprayed out behind the scorpion; the pedipalps and fourth pair of legs were used

5 mm
ee es \
oa |
sae Oe
—-—- eos ‘ x fas
£ é 2 ig Ys ° oF /

Fig. 6. Fifth caudal segment and telson of a. Parabuthus capensis (Ehrenberg) and b. Para-
buthus neglectus Purcell.

NOTES ON THE SCORPION FAUNA OF THE CAPE oti

as supports during this operation. To remove small stones or compacted soil the
first two pairs of legs were tucked up under the body and the tail extended
straight out backwards while the debris was transported rapidly backwards in a
single movement, with the last two pairs of legs being used for traction. Only
this latter part of the process is similar to the burrowing behaviour of Opistho-
phthalmus. The chelicerae were not observed to play any role in loosening the
sand.

The distribution of P. neglectus extends from Clanwilliam northward to the
Orange River, and eastward from De Doorns to Hanover and Graaff-Reinet in
the west. This species appears to be randomly distributed in the desert and semi-
desert regions of the Cape. In Namaqualand, the author has found this species
in close association with Opisthophthalmus granifrons Pocock, and Uroplectes
carinatus (Pocock).

P. neglectus also shows some colour variation. In northern Namaqualand
specimens may have the last two caudal segments and telson darkly pigmented
(Fig. 3b). This is also a feature of several other species in this area, P. schlechteri
Purcell, O. wahlbergi nigrovesicalis Purcell and O. w. gariepensis Purcell.

capensis

O| negiectus

Fig. 7. Map showing distribution of the Parabuthus capensis species-group.

We ANNALS OF THE SOUTH AFRICAN MUSEUM

In the author’s experience P. capensis has always been collected in areas of
hard-packed soil, with the exception of the black specimens from Laaiplek,
whereas P. neglectus has always been found in a sandy habitat. The longer and
denser setae of P. neglectus indicates that this species is psammophilous, whereas
the short and sparse setae of P. capensis indicates pelophily.

MATERIAL EXAMINED
All reference numbers refer to the South African Museum collection.
Parabuthus capensis (Ehrenberg) (yellow form)
Gouda (33°15’S 19°0’E)
I 2G758)
Tulbagh (33°20’S 10°10’E)
1 2 (504)
St Helena Bay (32°45’S 18°0’E)
356) Ga & Sef COOOT 15005 12807)
Eendekuil (32°40’S 18°45’E)
2 2 8 (B8934)
Jacobsbaai (32°55’S 17°55’E)
(C27)
Sak River
12 (Bits3)
Hermanus (34°25’S 19°15’E)
234,42 2 (11506)
Cape Peninsula (34°0’S 18°25’E)
1 g, 1 2 2813, B545)

Parabuthus capensis (Ehrenberg) (black form)
Laaiplek (32°15’S 18°5’E)

AS By 2 2 EC)

Clanwilliam (32°5’S 18°50’E)

1 2 (C36)

Parabuthus neglectus Purcell
Clanwilliam (32°5’S 18°50’E)
33 5,4 2 & (3760, 1199, 3754)

NOTES ON THE SCORPION FAUNA OF THE CAPE

Steinkopf (29°10’S 17°40’E)

ee. 4 2 2 (567, 1704, 1711, 1713)
Concordia (29°30’S 18°0’E)

ae.) 9? (1702, 5198)

Garies (30°30’S 18°0’E)

wea, > © 2 (1703, B7295, C43)
Okiep (29°35’S 18°0’E)

56,2 2 2 (1701)

Aggenys (29°10’S 18°50’E)

1 3 (2948)

Port Nolloth (29°10’S 17°55’E)
1 2 (BS68)

Kamieskroon (30°10’S 17°55’E)
fae.) 2 2 (BI314, B7338, C37)
Kliprand (30°30'S 18°40’E)

1 3 (C41)

Kuboos, Richtersveld (28°25’S 17°0’E)
8 ¢ 34,3 2 2 (B571, B8944)

Jakkalsputs, Richtersveld (28°35’S 17°0’E)
1 2 (C49)

Upington (28°30’S 21°15’E)

1 3, 1 2 (B8943)

Touws River (33°15’S 20°0’E)

3 2 2 (14263, 14365, 14378)

Van Rhynsdorp (31°35’S 18°40’E)
1 2 (1209)

Hanover (31°35’S 18°40’E)

1 3, 1 2 (9989, 9992)
Matjesfontein (33°15’S 20°35’E)
2 2 2 (12809, 14363)

Montagu (33°40’S 19°10’E)

1 2 (B3979)

213

214 ANNALS OF THE SOUTH AFRICAN MUSEUM

De Doorns (33°30’S 19°40’E)
1 3 (B609)

Calvinia (31°30’S 19°50’E)
1 g (12710)

Graaff-Reinet (31°30’S 24°30’E)
4 2 2 (12010)

ACKNOWLEDGEMENTS

I thank Dr V. Whitehead (Head, Department of Entomology, South African
Museum) for help and advice in the preparation of this paper, and the Director
for providing research facilities. Thanks are also due to Mr S. Kannemeyer for
photographic work, and Dr M. Moritz of the Zoologisches Museum, Berlin, for
the loan of type material.

REFERENCES

EHRENBERG, C. G. 1828. (Symbolae Physicae, seu Icones et descriptiones Corporum Naturalium
novorum aut minus cognitorum, quae ex itineribus per Libyam, Aegyptum, Nubiam
Dongalam, Syriam, Arabiam et Habessiniam. P. C. Hemprich et C. G. Ehrenberg...
studio annis 1820-25 dedierunt . . . Pars Zoologica 4 vol. fol. Berolini 1828-1845). Inverte-
brata other than Insecta. C. G. Ehrenberg: (122nd page). ;

Hewitt, J. 1918. A survey of the scorpion fauna of South Africa. Trans. R. Soc. S. Afr. 6:
89-192.

Pocock, R. I. 1902. A contribution to the systematics of scorpions. Ann. Mag. nat. Hist. (7)
10: 364-380.

PURCELL, W. F. 1899. New South African scorpions in the collection of the South African
Museum. Ann. S. Afr. Mus. 1: 433-438.

PURCELL, W. F. 1902. On some South African Arachnida belonging to the Orders Scorpiones,
Pedipalpi and Solifugae. Ann. S. Afr. Mus. 2: 137-225.

ABBREVIATIONS

d_ dorsals
db dorsal basals
dt dorsal terminals
e externals
eb, Eb external basals
esb, Esb_ external suprabasals
est, Est external subterminals
i internals
it internal terminals
V_ventrals
SAM_ South African Museum, Cape Town
ZMB_ Zoologisches Museum, Berlin

=
Ran oo

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

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

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

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

Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15A
Nucula (Leda) 6 aimee Gould, 1845: 37.
Leda plicifera A. Adams, 6: 50.
Laeda bicuspidata a 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (figs 8a—b).

Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:
comma separates author’s name and year
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figures of plates are enclosed in parentheses to distinguish them from text-figures
dash, not comma, separates consecutive numbers

Synonymy arrangement according to chronology of bibliographic references, whereby
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not acceptable.

In describing new species, one specimen must be designated as the holotype; other speci-
mens mentioned in the original description are to be designated paratypes; additional material
not regarded as paratypes should be listed separately. The complete data (registration number,
depository, description of specimen, locality, collector, date) of the holotype and paratypes
must be recorded, e.g.:

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

(c) Scientific names, but not their vernacular derivatives
e.g. Therocephalia, but therocephalian

Punctuation should be loose, omitting all not strictly necessary

Reference to the author should be expressed in the third person

Roman numerals should be converted to arabic, except when forming part of the title of a
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‘Revision of the Crustacea. Part VIII. The Amphipoda.’

Specific name must not stand alone, but be preceded by the generic name or its abbreviation
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Name of new genus or species is not to be included in the title: it should be included in the
abstract, counter to Recommendation 23 of the Code, to meet the requirements of
Biological Abstracts.

E. B. EASTWOOD

NOTES ON THE SCORPION FAUNA OF THE CAPE
PART 2

THE PARABUTHUS CAPENSIS (EHRENBERG)
SPECIES-GROUP; REMARKS ON TAXONOMY AND
BIONOMICS (ARACHNIDA, SCORPIONIDA, BUTHIDAE)

=. aa 7
a 7 a
- —— (ous y ~ e

/OLUME 73 PART 9 DECEMBER 1977 3 ISSN 0303-2515

ag ;

OF THE SOUTH

CAPE TOWN —

INSTRUCTIONS TO AUTHORS

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

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Examples (note capitalization and punctuation)

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,

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

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

THIELE, J. 1910. Mollusca: B. Polgulecenee, 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 73 ~~ + Band
December 1977 Desember
Part 9 Deel

CRETACEOUS FAUNAS FROM ZULULAND,
SOUTH AFRICA AND SOUTHERN MOZAMBIQUE
THE APTIAN ANCYLOCERATIDAE (AMMONOIDEA)

By

HERBERT CHRISTIAN KLINGER
&
WILLIAM JAMES KENNEDY

Cape Town Kaapstad

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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

CRETACEOUS FAUNAS FROM ZULULAND, SOUTH AFRICA, AND
SOUTHERN MOZAMBIQUE
THE APTIAN ANCYLOCERATIDAE (AMMONOIDEA)

By

HERBERT CHRISTIAN KLINGER

South African Museum, Cape Town
&

WILLIAM JAMES KENNEDY

Geological Collections, University Museum, Oxford

(With 89 figures)

[MS. accepted 14 April 1977]

ABSTRACT

Heteromorph ammonites referred to the subfamilies Ancyloceratinae, Heteroceratinae,
and Helicancylinae occur widely in rocks of Aptian age in northern Zululand and southern
Mozambique. The subfamily Ancyloceratinae is represented by Ancyloceras (Ancyloceras),
Ancyloceras (Adouliceras), Tropaeum, Australiceras and a specimen tentatively referred to
Lithancylus. The subfamily Helicancylinae, separated from the Ancyloceratinae with some
hesitation, is represented by Toxoceratoides and Tonohamites although doubt exists as to the
generic affinity of some of the species. The subfamily Heteroceratinae is represented in the
Upper Aptian by a helical hamitid-like form with or without tubercles, referred to a new
genus, Helicancyloceras. Dimorphism is tentatively recognized in Ancyloceras (Adouliceras)
and Helicancyloceras, whilst observations on the biogeography of the group are also included.

The following new species are described: Ancyloceras (Adouliceras) cooperi sp. nov.,
Tropaeum dayi sp. nov., Tropaeum obesum sp. nov., Australiceras wandalina (Boshoff MS)
sp. nov., Toxoceratoides ? haughtoni sp. nov., Tonohamites ? caseyi sp. nov., Helicancyloceras
(Helicancyloceras) densecostatum sp. nov., Helicancyloceras (Nonyaniceras) nonyani sp. nov.,
Helicancyloceras (Nonyaniceras) circulare sp. nov. and Helicancyloceras (Nonyaniceras)
crassetuberculatum sp. nov.

CONTENTS

PAGE

Introduction . : ' : : ; = ~2h6
Location of specimens . : : : mee Aly
Field localities : : : : : Coen 32) oF)
Measurements : : : , : 2 ets
Systematic palaeontology : ; : Ve e2ks
Subfamily Ancyloceratinae Meek, 1876. folks
Subfamily Helicancylinae Hyatt, 1894 . < 77305
Subfamily Heteroceratinae Spath, 1922. . *..) e825
Palaeobiogeography : : 4 . . 349
Acknowledgements . 3 : ; ; 395
References. : : : : : . + 356

DAS

Ann. S. Afr. Mus. 73 (9) 1977: 215-359, 89 figs.

216 ANNALS OF THE SOUTH AFRICAN MUSEUM

INTRODUCTION

Ammonites belonging to the family Ancyloceratidae are locally abundant
in the Aptian of northern Zululand and southern Mozambique. Most specimens
are fragmentary, and complete individuals rare, making positive identifications
difficult in many instances. In the descriptions given below, the authors have in
consequence made extensive use of open nomenclature in the sense of Richter
(1948; see also Matthews 1973).

The family Ancyloceratidae is represented by the following subfamilies in
the present collection: Ancyloceratinae Meek, 1876, Heteroceratinae Spath,
1922, and Helicancylinae Hyatt, 1900. Some of the Heteroceratinae have been
described elsewhere (Klinger 1976).

The subfamilies of the Ancyloceratidae are all intimately related, and some
may be superfluous; the authors retain them here as a working framework for
classification of their material, for they do seem to show a reasonable degree
of morphological homogeneity, and are of value when dealing with fragmentary
material, as Thomson (1974: 17) has noted. Extensive splitting of the hetero-
morphs at family level, as suggested by Dimitrova (1970), is to be avoided in
the authors’ view, given the current state of knowledge of several groups which
is still insufficient for a confident statement of their phyletic relationships,
especially as many classic species require reinvestigation.

The Ancyloceratidae have a world-wide distribution and, following the
systematic account, the authors give details of the palaeobiogeography of the
group.

Generic comparisons may be drawn on a global scale, but at specific level
most of the taxa encountered in the literature appear of more limited distribu-
tion, and some are markedly endemic. A close relationship exists, however,
between the Barremian and Aptian faunas of Zululand and southern Mozam-
bique and those of the southern U.S.S.R. as shown below, and as noted recently
by Forster (1975a: 256, 1975b: 270).

The following species are described below:

Ancyloceras? sp. ind.

Ancyloceras? sp. cf. humboldtiana (Lea)

Ancyloceras (Adouliceras?) sp. cf. ajax Anderson

Ancyloceras (Adouliceras) mozambiquense (Krenkel)

Ancyloceras (Adouliceras) cooperi sp. nov.

Ancyloceras (Adouliceras) sp. gr. ex. mozambiquense (Krenkel)—
cooperi Sp. NOV.

Lithancylus sp.

Tropaeum sp. aff. subsimbirskense subsimbirskense (Sinzow)

Tropaeum subsimbirskense compressum (Sinzow)

Tropaeum rossicum Casey

Tropaeum dayi sp. nov.

Tropaeum obesum sp. nov.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 217

Tropaeum sp. aff. undatum Whitehouse

Tropaeum sp. gr. ex. bowerbanki J. de C. Sowerby densistriatum
Casey —hillsi (J. de C. Sowerby)

Tropaeum sp. indet.

Australiceras ramososeptatum (Anthula)

Australiceras sp. aff. irregulare (Tenison Woods)

Australiceras wandalina (Boshoff MS) sp. nov.

Australiceras spp. indet.

Australiceras sp. cf. ‘Crioceras’ sarasini Favre

Toxoceratoides royerianus (d’Orbigny)

Toxoceratoides krenkeli Forster

Toxoceratoides ? haughtoni sp. nov.

Tonohamites koeneni Casey

Tonohamites sp. aff. aequicingulatus (von Koenen)

Tonohamites ? caseyi sp. nov.

Helicancyloceras (Helicancyloceras) vohimaranitraensis (Collignon)

Helicancyloceras (Helicancyloceras) densecostatum sp. nov.

Helicancyloceras (Helicancyloceras) spp. indet.

Helicancyloceras (Nonyaniceras) nonyani sp. nov.

Helicancyloceras (Nonyaniceras) circulare sp. nov.

Helicancyloceras (Nonyaniceras) crassetuberculatum sp. nov.

Helicancyloceras (Nonyaniceras) spp. fragments

Incertae sedis.

LOCATION OF SPECIMENS

The following abbreviations are used to indicate the source of the material:
SAS Geological Survey of South Africa, Pretoria
SAM South African Museum, Cape Town
UPG Department of Geology, University of Pretoria
BM(NH) British Museum (Natural History), London
NMB National Museum, Bloemfontein
OUM University Musem, Oxford
UP Palaeontological Institute, Uppsala University
MNHP Muséum d’Histoire Naturelle, Paris.

FIELD LOCALITIES

Details of field localities referred to in this paper are given by Kennedy
& Klinger (1975). Co-ordinates of additional localities in Zululand studied
since then, and localities in southern Mozambique collected by M. R. Cooper
are provided in the text.

218 ANNALS OF THE SOUTH AFRICAN MUSEUM

MEASUREMENTS

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

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

Figures in parentheses are dimensions expressed as a percentage of the
total diameter. As no standard set of abbreviations for heteromorph ammo-
noids has as yet come into common use, other dimensions are written out in full.

SYSTEMATIC PALAEONTOLOGY

Class CEPHALOPODA Zittel, 1884

Order AMMONOIDEA Zittel, 1884
Suborder ANCYLOCERATINA Wiedmann, 1966
Superfamily ANCYLOCERATACEAE Meek, 1876
Family Ancyloceratidae Meek, 1876
Subfamily | Ancyloceratinae Meek, 1876.

The features which allow separation of the Ancyloceratinae and Crio-
ceratitinae are very slight indeed. Casey (1960) accorded the Ancyloceratinae
and Crioceratitinae subfamilial rank. His extensive researches on the hetero-
morphs of the English Lower Greensand (Aptian to Lower Albian) have con-
tributed enormously to our knowledge of these ammonites, and without his
work the description of the Zululand and Mozambique species would have
been rendered much more difficult. In view of this, the subfamilies are separated
here also, although the differences in features which allow their distinction are
slight.

The Ancyloceratinae are not well represented in the Cretaceous of Zulu-
land and southern Mozambique, but few other ammonite groups present in
the area grow to such enormous sizes and show such distinctive and bizarre
coiling.

The subfamily is represented by the following genera: Ancyloceras s.1.
d’Orbigny, 1842, Tropaeum J. de C. Sowerby, 1837, Australiceras Whitehouse,
1926, and Lithancylus Casey, 1960.

Genus Ancyloceras d’Orbigny, 1842
Type species
Ancyloceras matheronianum d’Orbigny, 1842 from the Lower Aptian of
France by the subsequent designation of Haug (1889: 212).

Diagnosis

Small to large heteromorphs with ancyloceratid coiling; early whorls in
contact or widely separated. Ornament consists of trituberculate ribs separated

CRETACEOUS FAUNAS FROM SOUTH AFRICA 219

by a variable number of intermediate non-tuberculate ribs throughout, or only
in early and late growth stages.

Subgenus Ancyloceras (Ancyloceras) @Orbigny, 1842
Diagnosis
Ancyloceras in which trituberculate ribs are present throughout ontogeny.

Discussion

The genus Ancyloceras has become a receptacle for many heteromorphs
showing ancyloceratid coiling, and as Casey (1960: 21) indicated, most of the
Georgian (Gruzinian) ancyloceratids described by Rouchadzé (1933) are
generically distinct from Ancyloceras and should probably be referred to
Pedioceras or some genus within the Pedioceratinae. The same holds true for
some ancyloceratid forms from the U.S.S.R. and from Bulgaria described by
Drushchitz & Kudryavtsev (1960) and Dimitrova (1967) respectively. Inasmuch
as the family Pedioceratidae is not well defined, and Pedioceras is a poorly-
known genus (Yenne 1949 provides the most comprehensive discussion), it is
perhaps wisest to retain the forms noted above in Ancyloceras with a mark of
interrogation.

Ancyloceras (Ancyloceras ?) sp. indet.

Figs 1A-B, 18B, 79D
Material
SAS H54/39 and SAS B11 from the Lower Aptian of Locality 170,
Zululand.

Description

Only part of the initial coil and a non-septate fragment are known. Initial
coiling appears to have been relatively close, with the whorls just touching,
but not impressed. In SAS B11 (Fig. 1A—B) the two coils are not in one plane,
but are helicoid. This does not appear to be due to postmortem damage. The
whorl section is ovoid, wider than high (i.e. lateral diameter greater than
siphonal—antisiphonal diameter) with a broadly rounded venter and dorsum.
Ornament consists of major trituberculate ribs and a varying number (usually
one to three) of intermediaries. The tubercles are very prominent and were
originally spinose, and where the spines have broken off, low, round bosses
remain, indicating the presence of a basal septum separating spine from shell.

The suture line is too poorly preserved for comment.

Dimensions

Specimen D Wb Wh Wb/Wh U
SAS B11 c.36 8 16,5(c. 49) 12,6(e. 20) 13 17(c. 47)

220 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 1. A-B. Ancyloceras (Ancyloceras?) sp. indet. Dorsal and lateral views of specimen
SAS B11 from locality 170, Aptian I. x 1,5.

CRETACEOUS FAUNAS FROM SOUTH AFRICA BD)

Discussion

This is one of the few ammonites known in Bed 43 at locality 170 (see
Kennedy & Klinger 1975, fig. 11). The fact that the coils in specimen SAS B11
are not in one plane, but rather appear helical, is disturbing. According to
Clark (1958: 1076) the coiling in Ancyloceras is always planispiral, and is a
feature (amongst others), which serves to distinguish Ancyloceras from Aniso-
ceras. Unfortunately our material does not permit a more detailed investigation
to determine whether coiling is helical throughout the species or merely restricted
to this one specimen due to pathological reasons.

Ancyloceras helicoides Rouchadzé (1933: 217, pl. 8 (fig. 3)) from Kouthais,
Georgia (U.S.S.R.) is a helically coiled ancyloceratid, and is referred to a new
genus, Kutatissites, by Kakabadze (1970). If the Zululand species is indeed
coiled helically in all specimens, it should be referred to Kutatissites. This
would further accentuate the close similarity between the Lower Cretaceous
faunas of the southern part of the U.S.S.R. and south-eastern Africa.

Until further material becomes known the authors prefer to retain the
species in the genus Ancyloceras with a mark of interrogation.

Occurrence
Aptian I of Zululand.

Ancyloceras ? sp. cf. A. humboldtiana (Lea, 1841)

Fig. 2
Compare:

Ancyloceras humboldtiana (Lea, 1841) in Forbes 1845: 171-172.

Material
BMNH C79717 from locality 166, Mfongozi Creek, Zululand, Aptian III.

Description

The single specimen consists of a fragment of the inner whorls and part
of the outer whorl and straight shaft.

The inner whorl is ornamented by single, trituberculate ribs. On the outer
whorl and shaft ribbing becomes sparse, and tuberculation weakened and
possibly disappears. On the shaft the ribs are widely spaced with smooth
interspaces.

Discussion

Obviously specific identification based on this fragment alone is impossible.
The Zululand specimen bears some similarity to the specimen figured by
Forbes, in possessing widely-spaced ribs on the shaft, but differs in lacking the
fine intermediary ribs.

932 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 2. Ancyloceras ? sp. cf. A. humboldtiana (Lea, 1841). BMNH C79717 from locality 166,
Aptian III. x 1.

Occurence
Upper Aptian of Zululand.

Subgenus Ancyloceras (Adouliceras) Thomel, 1964
Type species

Ancyloceras adouli Astier, 1851 by the original designation of Thomel
(1964: 56).
Diagnosis

Coiling ancyloceratid, generally with small initial spire, the whorls of
which expand rapidly. Tubercles are linked by two or three ribs, but disappear

CRETACEOUS FAUNAS FROM SOUTH AFRICA 223

at varying diameters on the initial spire. Shaft generally ornamented by fine,
dense ribs only, without tubercles or differentiation into strong and weak ribs
for at least part of the length.

The crozier may be inflated and is generally ornamented by three strong
rows of tubercles (lateral, lower and upper ventro-lateral) at some stage.

Discussion

Adouliceras was erected as a subgenus of Ancyloceras by Thomel (1964: 55)
with Ancyloceras adouli Astier, 1851 as type species. The holotype, BMNH
C73806, reposited in the British Museum (Natural History), is here illustrated
as Figures 3-4. Thomel’s original diagnosis is as follows:

‘Le sous-genre Adouliceras, créé pour les formes du groupe d’Ancyloceras
adouli ASTIER, est caractérisé par une spire da croissance tres rapide, a
cette partie de la coquille, un port remarquable. L’ornamentation de la
spire est également particuliére; elle consiste en cétes fines, simples toutes
semblables, sur lesquelles on observe, de place en place, des tubercles, plus
ou moins volumineux 4 cheval sur deux ou trois consécutives. La hampe,
de section elliptique, est ornée de cdtes simples, obliques, vigoureuses,
généralement dépourvues de tubercules, du moins sur la partie cloisonnée.
La crosse est connue seulement chez A. adouli ou elle porte des tubercules
trés vigoureux, et chez A. renauxianum.’

In addition to the type species, A. kaliae (Sarkar), A. collignoni (Sarkar)
and A. renauxianum were referred to the subgenus. Comparison of A. adouli
and A. renauxianum shows the extreme variation encountered in A. (Adouli-
ceras). In A. (Ad.) adouli the whorl section becomes abruptly inflated towards
the hook. Furthermore, the initial spirally coiled section in the latter lacks
tuberculation altogether. (See d’Orbigny 1842, pl. 123.) Whether this is in fact
true, or merely another of d’Orbigny’s artist’s restorations is unknown.

Recently Murphy (1975) described a species from California under the
name of Ancyloceras thomeli. In all respects this species has the characteristics
of Adouliceras, apart from the fact that ‘the spire of A. thomeli tapers much less
rapidly than that of A. adouli which, according to Thomel (personal communi-
cation) is grounds for excluding the California form from Adouliceras’ (Murphy
1965: 25). Comparison of the holotype of A. (Ad.) adouli, Figures 3-4, with the
specimen figured by Thomel (1964, pl. 9 (fig. 2)) shows that the size of the initial
spire is variable. This is further borne out by the Zululand and Mozambique
material to be described below. The presence or absence of fine ribs on the hook
quoted by Murphy as being a reason for separating Ancyloceras thomeli from
A. (Ad.) adouli is just as variable.

It is here proposed to consider the absence of tubercles or stronger ribbing
on part of the shaft and the mode of ornament on the early whorls as consti-
tuting the main characteristic of Adouliceras. The size of the initial spire is
generally smaller than that of Ancyloceras s.s.

224 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 3. Ancyloceras (Adouliceras) adouli Astier, 1851. BMNH C73806, the original of Astier
(1851, pl. 6 (mo. 12), pl. 7 (no. 12 bis)), from the Neocomian of Cheiron, Basses Alpes (France).
x 0,45.

CRETACEOUS FAUNAS FROM SOUTH AFRICA

Fig. 4. Ancyloceras (Adouliceras) adouli Astier, 1851. As in Figure 3.

225

226 ANNALS OF THE SOUTH AFRICAN MUSEUM

In addition to the species mentioned above, the following may possibly
also be referred to Adouliceras: Ancyloceras attrox Anderson (1938: 209 pl. 69
(figs 1-3)) and Ancyloceras ajax Anderson (1938: 210, pl. 65 (figs 1-3)). Ancylo-
ceras ewaldi Dames (1880: 690, pl. 25, pl. 26 (fig. 1)) is probably also an example
of Adouliceras.

Adouliceras represents a line of development which resembles that of
Australiceras gr. ex. gigas, and some species were previously referred to that
genus (e.g. Casey 1961: 51). Thomel (1964: 60, table 2), however, has shown
that they are parallel but apparently unrelated lineages.

Adouliceras differs from Ancyloceras s.s. in lacking tubercles on part of
the shaft. It differs from Australiceras gr. ex. gigas mainly in having a smaller
initial spire, a different mode of tuberculation, looser coiling, and, most impor-
tant of all, in lacking major costae and tubercles on the shaft. It may be
distinguished from uncoiled species of Tropaeum e.g. T. gr. ex. hillsi by the
possession of three rows of tubercles on the early and late parts of the shell.
Non-tuberculate fragments of the ammonitic coils are generically indistinguish-
able from Tropaeum.

Unfortunately the South African and Mozambique material does not shed
any light on the phylogeny and relationship of Adouliceras to other genera of
the Ancyloceratinae.

Occurrence

Upper Barremian and Lower Aptian of western and central Europe,
California, Zululand and Mozambique.

Ancyloceras (Adouliceras ?) sp. cf. ajax Anderson, 1938

Figs 5-6A
Compare:
Ancyloceras ajax Anderson, 1938: 210, pl. 65 (figs 1-3).
Material

SAS Zo(1) from locality 162, Mfongozi Creek, Zululand. Lower ? Aptian.

Description

One large fragment consisting of the non-septate part of the shaft and
crozier is compared to Anderson’s species. The terminal septum is exposed at
the broken end of the shaft.

The section of the shaft is ovoid, with a greater dorso-ventral than lateral
diameter. Towards the crozier the whorl breadth increases rapidly, surpassing
the whorl height. On the recurved end the whorl section is depressed with a
flattened dorsum, moderately angular umbilical edge, and a broadly rounded
venter.

Ornament on the shaft consists of low, rounded, oblique, prorsiradiate ribs.
The ribs cross the dorsum with a slight forward curvature. The venter is abraded,

CRETACEOUS FAUNAS FROM SOUTH AFRICA 225),

Fig. 5. Ancyloceras (Adouliceras) sp. cf. ajax Anderson, 1938. SAS Zo(n) from locality 162,
Mfongozi Creek, Zululand, Lower? Aptian. x 0,4.

but it appears that the ribs crossed the venter without diminution. There are
about 25 ribs in a distance equal to the whorl height. At a point 70-80 mm
before the bend in the crozier, strong ribs appear, becoming progressively
stronger towards the bend. Umbilical, lateral and ventral tubercles develop on
the major ribs. The major ribs cross the flanks radially and curve forward over
the venter, but are effaced on the dorsum.

228 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 6. A. Whorl section of A. (Adouliceras) sp. cf. ajax Anderson, 1938. SAS Zo(n). x 1.
B. Whorl section of Australiceras wandalina sp. nov. UPG B8 from Manyola Drift, Aptian

Mr ><. 0%5;
Dimensions
Wb 80 101,5 89,5
Wh _~ 92,5 91,6 97,0
End of shaft. In crozier. Aperture.
Discussion

The specimen bears a strong resemblance to the Californian Ancyloceras
ajax, but lacks the inflated body chamber and the very strong tuberculation
on the crozier.

Ancyloceras thomeli Murphy (1975: 24, pl. 3 (figs 1, 5), pl. 11 (figs 1—-2)) also
differs on account of the presence of an inflated body chamber.

In A. (Ad.) renauxianum @Orbigny (1842: 499, pl. 123) non-tuberculate
intermediaries occur between the major costae in the crozier. A. (Ad.) ewaldi
Dames (1880: 690, pl. 25, pl. 26 (fig. 1)) has stouter ribbing on the shaft, and
more closely spaced ribs on the crozier. A. (Ad.) adouli Astier (1851: 23, pl. 6
(nr 12); pl. 7 (nr 12b); see also Thomel 1964, pl. 9 (fig. 2)) has a strongly inflated
body chamber and is readily distinguishable from the present specimen.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 229

Occurrence

The exact stratigraphic position of the Zululand specimen is unknown,
although definitely Aptian. Anderson’s species is said to come from close to the
Barremian/Aptian boundary.

Ancyloceras (Adouliceras) spp.
group of mozambiquense Krenkel—cooperi nov.

Abundant leached concretions yielding fragments of ancyloceratids litter
the surface and topsoil at locality 170. The stratigraphic interval represented by
these concretions is at least 10 m, whilst similar material from Da Silva’s (1962)
locality A (26°49'50”S 32°12’55”E) herein referred to as Lubemba, in southern
Mozambique is available. Apart from the ancyloceratids, the associating fauna
consisting of Cheloniceras spp., Neosilesites and Valdedorsella indicates a Lower
Aptian age in both cases, and subsequent work by Forster (1975a) confirms this.

All the ancyloceratid fragments consist either of parts of the early whorls,
parts of the uncoiled shafts and/or the crozier, and at the time of writing, no
complete specimen has been found. Two different kinds of early whorls can be
identified; one with a circular, the other with a subtrigonal whorl section, the
latter type being commonest. The shaft and croziers, on the other hand are so
varied that hardly two specimens are alike. In addition to differences in ornamen-
tation, remarkable differences in size occur, Macro- and micro-conchs being
present.

The early whorls with subtrigonal whorl section can be identified with
A. (Ad.) mozambiquense Krenkel; those with circular whorl section are referred
to a new species, A. (Ad.) cooperi sp. nov. Some types of shaft and crozier may
tentatively be referred to the two species. However, due to the extreme variability,
and the absence of any complete specimens for definite allocation, the shafts and
croziers are referred in open nomenclature to A. (Ad.) gr. ex. mozambiquense—
cooperi. Admittedly, this may be interpreted as vertical systematics par excellence,
but in the authors’ opinion this is preferable to the other alternatives—to ignore
the material until such time as future collecting may hopefully yield a complete
specimen, or to erect a number of species based on incomplete specimens as has
so often been done in the past. Moreover, identification based on the shaft and
crozier alone is impracticable as will be shown in the discussion below, for even
unrelated groups of heteromorphs may have the same type of crozier.

Ancyloceras (Adouliceras) mozambiquense Krenkel, 1910
Figs 7, 8A—B, 9B, 10B, 11B, 12A, D

Ancyloceras sp. Kilian, 1902: 465.

Ancyloceras fallauxi Uhl. n. var. mozambiquense Krenkel, 1910: 153, pl. 17 (figs 2-3).

? Ancyloceras fallauxi Uhlig var. mozambiquense: Haughton & Boshoff, 1956: 14.

Tropaeum mozambiquense: Wachendorf, 1967: 292, pl. 34 (fig. 1). Non Da Silva 1962: 21,
pl. 9 (figs 1-2), pl. 10 (fig. 1), pl. 11 (fig. 1), pl. 12 (fig. 1), pl. 13 (fig. 1), pl. 14 (fig. 1).

? Tropaeum cf. hillsi: Forster, 1975a: 151, pl. 2 (fig. 1), text-fig. 28.

Australiceras mozambiquense: Forster, 1975a: 155, pl. 3 (fig. 3), text-fig. 31.

230 ANNALS OF THE SOUTH AFRICAN MUSEUM

Holotype

The specimen figured by Krenkel (1910, pl. 17 (figs 2-3)) reported to have
been collected from near Delagoa Bay, Mozambique.

Neotype

SAM-PCM5349 (Fig. 7) from Lubemba, southern Mozambique. The
holotype was destroyed by bombing during 1944.

Material

SAM-PCM5314, SAM-PCM5351, SAM-PCM5438 from Lubemba,
southern Mozambique; SAS 54/45; SAS 54/41/4; SAS 54/45/2; BMNH C78883
and BMNH C78884 from locality 170, Mlambongwenya Creek, Zululand.
Aptian J-II.

Description

Coiling is crioceratitid but very variable. On the most complete specimen,
SAM-PCM5349 (Fig. 7), the inner whorl is not in contact with the outer. Some
specimens, however, have a distinct dorsal impression of the ventral tubercles
on the preceding whorl. The whorl section is typically subtrigonal to rectangular,
higher than wide with a flattened but not impressed dorsum and slightly
inflated flanks converging to a moderately rounded venter. In some specimens
the dorsum is rounded and convex (Fig. 8A).

As can be seen from the impression of the inner whorl of specimen
SAM-PCM5349 (Fig. 7), juvenile ornament consists of very fine, radial ribs
which cross the dorsum with traces of duplication. The umbilical tubercles
appear to be most prominent, and appear as low, rounded bosses, covering one
to three ribs. The bosses probably indicate the former presence of long spines,
with a basal septum. The lateral tubercles are smaller and more pointed.
Ventral tubercles are not preserved, but judging by their impression on the
dorsae of some of the larger whorls, were quite prominent and situated close to
each other on either side of the siphonal line.

At larger diameters, ribbing becomes weaker and curves forward over the
dorsum, although radial to slightly sinusoidal across the flanks. The ribs cross
the venter without visible sign of interruption. Bifurcation of ribbing occurs at
the umbilical edge, or on the dorsal third of the flanks. The diameter at which
tuberculation disappears is very variable, the ventral and lateral tubercles
disappearing first. At a whorl height of approximately 45 mm, the last umbilical
tubercles are visible. There are about 11 to 13 ribs in a distance equal to the
whorl height.

The suture is highly incised and dendritic. Unfortunately, however, none of
the specimens is suitable for reproduction of the whole suture line.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 231

Fig. 7. Ancyloceras (Adouliceras) mozambiquense Krenkel, 1910. SAM-PCM5349, neotype
from the Lower Aptian of Lubemba, Mozambique. Collected by M. R. Cooper. x 1,3.

232 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 8. Ancyloceras (Adouliceras) mozambiquense Krenkel, 1910. A. SAS 54/45. B. SAS 54/1.
Both from the Lower Aptian, Aptian I-II of locality 170. x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 233

Fig. 9. A. Ancyloceras (Adouliceras) gr. ex mozambiquense-cooperi. SAM-PCMS5328, crozier

from the Lower Aptian of Lubemba, Mozambique. Note the strong development of the

umbilical and lateral tubercles. x 0,85. B. Ancyloceras (Adouliceras) mozambiquense Krenkel,
1910. SAS 54/45/2 from locality 170, Zululand, Lower Aptian, Aptian I-II. x 1.

i ee ee ee ee ce | ae

234 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 10. A. Australiceras ? sp. cf. ‘Crioceras’ sarasini Favre, 1908. SAS H71D/18 from Nhlohlela
pan, Mkuze Game Reserve, Zululand, Aptian IIJ-IV. x 1. B. Ancyloceras (Adouliceras)
mozambiquense. SAS 54/45 from locality 170, Zululand, Aptian I-II. x 1,2.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 235

Fig. 11. A. Ancyloceras (Adouliceras) gr. ex mozambiquense-cooperi. SAS 54/45/3, crozier

from locality 170, Zululand, Aptian I-II. Form 3 with only laterally developed tubercles in

crozier. x 0,54. B. Ancyloceras (Adouliceras) mozambiquense Krenkel, 1910. SAS 54/41/3
from locality 170, Zululand, Aptian I-II. x 1,2.

236 ANNALS OF THE SOUTH AFRICAN MUSEUM

Dimensions
Specimen Wb Wh Wb/Wh
SAM-PCM 5349 44 47 0,93
SAS 54/41/3 51,6 56,4 0,91
Discussion

It is difficult to understand how Krenkel’s (1910: 153) original description
of the species was so consistently misinterpreted by subsequent workers;
Krenkel states of the unique holotype (destroyed during the Second World
War):

‘.. . das aus dem Ubergang des spiral eingerollten Teiles zum Schaft
stammt. Die Innenseite zeigt keine Spur einer Beriihrung mit dem friiheren
Umgange.’

Da Silva’s interpretation of Krenkel’s species is unacceptable, and his
Tropaeum mozambiquense was merely a catch-all for fragments of a variety of
Tropaeum. The specimen which Da Silva (1962: 21, pl. 9, figs 1-2) described as
corresponding to Krenkel’s original, has a distinctly impressed dorsal zone, and
this is a feature Krenkel explicitly stated to be absent. Forster (1975a) has
recently described material from the same area in Mozambique, and he realized
that most of Da Silva’s specimens of ‘Tropaeum mozambiquense’ were better
referred to Tropaeum subsimbirskense compressum (Sinzow). Forster also noted
the presence of three rows of tubercles on some of his material and concluded
that Krenkel’s species must be an australiceratid, with affinities with
A. rabenjanaharyi Collignon and A. ramososeptatum (Anthula), again a
departure from Krenkel’s original concept.

Other finely ribbed species of Tropaeum bear a strong similarity to the
non-tuberculate fragments of A. (Ad.) mozambiquense. On the basis of the
present material, however, the ancyloceratid character of the species is firmly
established.

A. (Ad.) mozambiquense is easily distinguished from other species of
A. (Adouliceras) by the sub-triangular whorl section and by the relatively large
planispirally coiled immature stage which is a departure from the normal
Adouliceras pattern.

Occurrence
Lower Aptian, Aptian I-II of Zululand and southern Mozambique.

Ancyloceras (Adouliceras) cooperi sp. nov.
Figs 12B—C, 13A-C, 14A—C
Holotype

SAM-PCMS317 in the South African Museum, from Lubemba, southern
Mozambique. Collected by M. Cooper. Aptian I-II.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 237

Derivation of name

The species is named for Michael Cooper, of Wolfson College, Oxford, who
collected the type material.

Material

Apart from the holotype; SAM-PCM5313, SAM-PCM5215, from
Lubemba, and SAS 54/41/1 and SAS 54/41/2 from locality 170, Mlambo-
ngwenya, Zululand, Aptian I-II. ©

Description

Coiling is tight, resulting in a small planispirally coiled section. The whorls,
however, are not impressed. The very early whorls are unknown, but at a
diameter of c. 20 mm the section is already essentially circular, and only a little
wider than high. The dorsum is slightly flattened, but with no trace of the
impressed zone. Ornament initially consists of radial, single, broad, low ribs,
each bearing three rows of low and rounded tubercles. The tubercles are of
approximately equal size, and appear to be the bases of septate spines. The
tuberculate ribs are separated by one or two narrow intermediaries. The latter
are thin and separated by interspaces of equal width. They sweep backward over
the umbilical wall, and are straight and radiai to rursiradiate across the flanks
and pass straight across the venter. On the dorsum, the ribs curve forward and
show a notable tendency towards duplication. With increasing diameter the
major ribs tend to become wider and the lateral and ventral tubercles disappear
at a whorl breadth of around 23 mm. At this stage, ribs now arise in two’s,
three’s and four’s from the remaining, but weakened, umbilical tubercles. On the
holotype, umbilical tubercles are still visible at a whorl breadth of 32 mm, but
on another specimen, SAS 54/41/2 (Fig. 14), no sign of tubercles is visible at a
whorl breadth of 21 mm. Two very slight constrictions are present on
SAM-PCM5313.

Dimensions

Specimen D Wb Wh Wb/Wh U
SAM-PCM 5317 89,2 37(41,4) 40,2(45) 0,92 33, 5(51.))
Discussion

The round whorl section clearly distinguishes A. (Ad.) cooperi from
A. (Ad.) mozambiquense. Furthermore, the initial spire is much smaller in the
former. The holotype bears a superficial resemblance to some of the species
redescribed by Thomel (1964), e.g. A. (Ad.) aff. collignoni Sarkar, 1955 (Thomel
1964: 58, pl. 9 (fig. 1), text-fig. 7A), A. (Ad.) kaliae Sarkar 1955 (Thomel 1964:
57, pl. 10 (fig. 1), text-fig. 7B), and A. (Ad.) adouli Astier (Thomel 1964: 56, pl. 9
(fig. 2), text-fig. 7C) as far as the size of the planispirally coiled section is
concerned. A. (Ad.) adouli has a whorl section similar to that of the present
species, but lacks the regular tuberculation (see Thomel 1964: 58, fig. 7) and has

238 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 12. A. Ancyloceras (Adouliceras) mozambiquense Krenkel, 1910. SAM—PCM5505 from

Lubemba, Mozambique, Lower Aptian. x 1,3. B. Ancyloceras (Adouliceras) cooperi sp. nov.

SAM-PCM5S215 from same locality and horizon as above. x 1,4. C. Ancyloceras (Adouliceras)

cooperi sp. nov. SAS 54/42/2 from locality 170, Zululand, Aptian I-II. Typical rounded form.

x 1,4. D. Ancyloceras (Adouliceras) mozambiquense Krenkel, 1910. SAM-—PCMS5351 from
Mozambique, Lower Aptian. x 1,3.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 239

Fig. 13. A-C. Ancyloceras (Adouliceras) cooperi sp. nov. SAM-PCMS5317, holotype from
Lubemba, Mozambique, Lower Aptian. x 1,05.

240 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 14. A-C. Ancyloceras (Adouliceras) cooperi sp. nov. SAS 54/41/2 from locality 170,
Zululand, Aptian I-II. x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 241

numerous intermediary ribs. A. (Ad.) kaliae Sarkar from the Upper Barremian
is very similar as far as ornamentation is concerned, but has an ovoid whorl
section. A. (Ad.) aff. collignoni has irregular ornamentation.

Occurrence
Lower Aptian of Zululand and Mozambique.

Ancyloceras (Ad.) gr. ex. mozambiquense—cooperi
(Description of shafts and croziers)
Form 1
Figs 15, 16, 17, 18A, 19, 20A—B
Material
SAM-—PCM5337, SAM-—PCM5395, SAM-PCM5368, SAM-PCM5449,

SAM-PCM5324, from Lubemba, southern Mozambique. BMNH C7884 from
Locality 170, Zululand.

Description

The shaft is of variable length with a subtrigonal section, having a narrower
venter than dorsum and weakly inflated flanks. Towards the crozier the dorsum
becomes more flattened and the dorso-ventral width increases. In the bend of
the crozier and on the descending limb the whorl section is dome-shaped with a
flat dorsum and broadly rounded flanks and venter. There is no distinct inflation
of the body chamber.

At the proximal end of the shaft, ornament consists of narrow, low, slightly
prorsiradiate ribs only, separated by wider interspaces. Average rib density is
about sixteen to eighteen per dorso-ventral diameter. On the distal half to third
of the shaft, stronger ribs start appearing on the flanks, bearing umbilical and
lateral tubercles. These ribs and the tubercles become progressively more
prominent towards the bend in the crozier. At the bend the intermediaries
become fainter and eventually disappear. In the bend itself, one or two of the
major costae now acquire ventral tubercles and cross the venter with a
forward curvature. At this stage the lateral tubercles are generally strongest. In
the descending limb the ribs become narrower and increasingly flared, accom-
panied by loss of tuberculation. Three flared ribs are present in the most
complete specimen.

The last septum occurs at a point coinciding approximately with the onset
of major ribbing.

Dimensions

Specimen SAM-PCM5324 SAM-—PCM5368
Wb. at proximal end of shaft . .. 91 78
Mvh-at-proximalend of shaft .~. .. . 105 87

WD. in crozier : : ‘ : ; ; 117 99

242 ANNALS OF THE SOUTH AFRICAN MUSEUM

Specimen SAM-PCM5324 SAM-PCM5368
Wh. incrozier. , ae: 109 7 93
Wb. near aperture : : 110 97
Wh. near aperture oe: 100 90
Width of crozier = venter of shaft to
venter of descending limb . ; 295 230
Discussion

This is the most common form present, and it is likely that it belongs to the
planispiral whorls identified as A. (Ad.) mozambiquense, which are also the most
abundant, as is suggested by the subtrigonal whorl. Until more complete
material becomes available to substantiate or discredit this assumption, it is
considered advisable to refer the shafts and croziers in open nomenclature to
A. (Ad.) gr. ex. mozambiquense-cooperi.

A number of shafts and croziers from various unrelated species with quite
distinct early whorls are virtually identical to, or impossible to distinguish from,
the group of specimens available. These include A. (Ad.?) durelli Anderson
(1938: 210, pl. 65 (figs 1-2), pl. 68 (fig. 1)), Ancyloceras matheronianum @ Orbigny
in Drushchitz & Kudryavtsev (1960, pl. 34 (fig. 2)), Ancyloceras urbani Neumayr
& Uhlig, 1881, and Ancyloceras rochi Dimitrova (the latter two in Dimitrova
1967, pls 23, 24 respectively), and it is apparent that species based on croziers
and shafts are of little value in at least some ancyloceratid genera.

Form 2
Fig. 21B
Description and discussion
Crozier SAS 54/45/5 is virtually identical to the previous form as far as
ornamentation is concerned, but is only half the size. These differences probably
reflect sexual dimorphism, although this has not been widely recognized in
ancyloceratids.

Dimensions
Wb. at proximal end of shaft . . 54
Wh. at proximal end of shaft . . 66
WIN GROZICr = vo os Sa ee
Wh. incrozier_. : te
Wb. near aperture. : : eal
Wh: near aperture — 229 75-2 on
Width ofeenozier 4) 20
Form 3

Figs 11A, 23
Description and discussion
Crozier SAS 54/45/3 has only one lateral row of irregularly developed
tubercles and does not develop flared ribbing immediately before the aperture.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 243

Dimensions
Wb. at proximal end of shaft . . —
Wh. at proximal end of shaft . . 78
Wontascrozier oh...) Sa 91
WhowinterOzien $2 5. Bie 1) 8d
Wb. at aperture . re Se en es Se
Wh. at aperture . 77 86
MIGtIMOnerO7ich.. so ea ts 21S

Fig. 15. Ancyloceras (Adouliceras) gr. ex mozambiquense-cooperi. SAM-PCMS5324, crozier
from Lubemba, Mozambique. Lower Aptian. Most common form, Form 1. x 0,48.

244 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 16. Ancyloceras (Adouliceras) gr. ex mozambiquense-cooperi. SAM-PCMS5324, ventral
view of crozier from Lubemba, Mozambique, Lower Aptian. Most common form, Form 1.
<OLD7:

CRETACEOUS FAUNAS FROM SOUTH AFRICA 245

Fig. 17. Ancyloceras (Adouliceras) gr. ex mozambiquense-cooperi. SAM—PCM 5368, ventral view
of crozier from Lubemba, Mozambique, Lower Aptian. Most common form, Form
ix 0167:

246 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 18. Ancyloceras (Adouliceras) gr. ex mozambiquense-cooperi. SAM-PCM5449, lateral

view of shaft from Lubemba, Mozambique, Lower Aptian. Most common form, Form 1.

x 0,77. B. Ancyloceras (Ancyloceras?) sp. indet. SAS B11 from locality 170, Zululand,
Aptian I. x 1,55.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 247

Fig. 19. Ancyloceras (Adouliceras) gr. ex mozambiquense-cooperi. SAM-PCMS5337 from
Lubemba, Mozambique, Lower Aptian. Most complete shaft of common form, Form 1. x 0,68.

248 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 20. A-B. Ancyloceras (Adouliceras) gr. ex mozambiquense-cooperi. BMNH C78884 from
locality 170, Zululand, Aptian I-II. Specimen with finer ribbing than previous examples.
x 0,67.

Form 4

Fig. 21A
Description and discussion

Crozier SAS 54/45/6 completely lacks major ribbing on the ascending shaft.
This crozier may be compared with A. (Ad.) ewaldi Dames (1880: 690, pl. 25,
pl. 26 (fig. 1)). The latter species, however, has stronger and more rounded
ribbing on the shaft. This specimen is crushed, hence no measurements are given.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 249

Fig. 21. A-B. Ancyloceras (Adouliceras) gr. ex mozambiquense-cooperi. A. SAS 54/45/6 from
locality 170, Zululand, Aptian I-II. Form 4. x 0,57. B. SAS 54/45/5 from same locality and
horizon as above. Form 2. x 0,53.

250 ANNALS OF THE SOUTH AFRICAN MUSEUM

Form 5

Fig. 22
Description and discussion

Crozier SAS 54/45/4 is a micromorph with an ovoid whorl section on the
shaft and with plain, simple ribbing throughout, apart from a smooth zone on
the descending shaft and a slightly flared rib just before the aperture. A few
paired ribs are more prominent than the others.

The rounded whorl section may indicate that this type of crozier may
belong to the inner whorls referred to A. (Ad.) cooperi sp. nov. The absence of
major costae on the crozier is reminiscent of that of Colchidites colchicus
Djanélidzé (1926, pl. 1 (fig. 1)). The smooth part of the descending limb (which
may be a malformation) is reminiscent of Anderson’s (1938) genus Shastoceras.
The body chamber of Shastoceras, however, is inflated and internal moulds are
reportedly smooth.

Dimensions
Wb. at proximal end of shaft . . 54
Wh. at proximal end of shaft . . 57

Wb. incrozier . ‘ i : ego 7/
Wh. incrozier_ . : ; » 6 16x66
Wb. at aperture: : - 4 “+ 3 eo
Whataperture,. . .‘°-.. eS
Width of crozier . y : . eS

Length of shaft to bend in crozier . 170

Form 6
Figs 9A, 24
Description and discussion

A single specimen, SAM-PCM5328 represents that part of a specimen at
the point where the shaft begins curving into a crozier. Three sets of major ribs
are present, separated by two to three low, rounded ribs. On the first major rib
three rows of tubercles are present, of which the lateral one is best developed.
At the distal end, however, only umbilical and lateral tubercles remain, but the
lateral tubercle is enormous.

Viewed laterally there is resemblance to Murphy’s Ancyloceras thomeli
(1975: 24, pl. 3 (figs 1, 5), pl. 11 (figs 1-2)). In the Californian species, however,
the ventral row of tubercles only appears towards the bend in the crozier,
whereas the reverse holds true for the Zululand specimen.

Genus Lithancylus Casey, 1960
Type species
Hamites grandis J. de C. Sowerby, 1828 from the Lower Aptian of southern
England by original designation of Casey (1960: 16).

CRETACEOUS FAUNAS FROM SOUTH AFRICA 251

Diagnosis

Coiling probably ancyloceratid, with a long, curved or straight shaft,
ending in a recurved hook. Section of shaft circular to oval, sub-octagonal on
crozier. Ornament consists of low, rounded oblique ribs on the shaft; towards

the crozier trituberculate ribs occur.

Fig. 22. Ancyloceras (Adouliceras) gr. ex mozambiquense-cooperi. SAS 54/45/4 from locality
170, Zululand, Aptian I-II. Form 5. x 0,59.

Ze ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 23. Ancyloceras (Adouliceras) gr. ex mozambiquense-cooperi. SAS 54/45/3 from locality
170, Zululand, Aptian I-II. Form 3. x 0,79.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 253

Fig. 24. Ancyloceras (Adouliceras) gr. ex mozambiquense-cooperi. SAM-PCM5328 from
Lubemba, Mozambique, Lower Aptian. Form 6. x 0,8.

Discussion

This is a relatively rare genus, characterized by the long, plainly ribbed shaft
ending in a crozier with trituberculate ribs. The presence of an initial planispirally
coiled part has yet to be demonstrated.

Apart from the type species, the genus is represented by L. fustis Casey
(1961: 75, pl. 21 (fig. 4a-d)) from the Lower Aptian of southern England,
L. tirolensis Casey (1961 : 74, text-fig. 29) from the Upper Barremian of Puez Alp,
Tyrol, L. mirabilis Anderson (1938: 220, pl. 78 (figs 1, 2a, 3)), L. nauplius
Anderson (1938: 221, pl. 66 (fig. 2—2a)), L. neleus Anderson (1938: 221, pi. 59
(fig. 2—2a)), and L. cyclopius Anderson (1938: 221, pl. 58 (fig. 3), pl. 66 (fig. 3—-3a)),
all from the Horsetown Beds of California, and probably of Barremian to
Aptian age. L. australe Day (1967: 19, pl. 2 (figs 1-5), text-fig. 2) is from the
Aptian of the Eromanga Basin of Queensland, Australia, and L. guanacoensis

254 ANNALS OF THE SOUTH AFRICAN MUSEUM

Leanza (1970: 204, fig. III, 1-4) is reported from Patagonia. Thomson (1974: 15,
pl. 2 (fig. 1) tentatively referred a specimen from Alexander Island (Falkland
Island Dependencies) to Lithancylus. Reference of the Bulgarian specimen of
Ancyloceras elephas Anderson, 1938 to Lithancylus by Dimitrova (1967: 61,
pl. 29 (fig. 1)) is questionable. The Bulgarian specimen shows an inflated body
chamber which is a characteristic of Ancyloceras (Adouliceras) rather than
Lithancylus. |

Differentiation of species is based mainly on the whorl section of the shaft
and the density of ribbing thereupon.

Occurrence

Upper Barremian and Lower Aptian of England, Tyrol, California,
Falkland Island Dependencies?, Patagonia, Mozambique and Australia.

Lithancylus sp.
Figs 25A-B, 83A, 84A—C
Material
SAM-PCM 5436 from Lubemba, southern Mozambique.

Description

A straight, septate piece of shaft is probably referable to this genus. At the
proximal end the whorl section is subtrigonal, with a broadly rounded dorsum
and little-inflated flanks converging to a narrower, rounded venter. At the distal.
end the whorl section is virtually equidimensional, and octagonal. Ornament at
the proximal end consists of low, rounded prorsiradiate ribs, separated by
interspaces wider than themselves. It appears as if the ribs are weaker on the
dorsum than on the flanks. At the distal end, three stronger trituberculate ribs
occur, separated by two intermediary ribs. One pair of tubercles is situated
ventrally, the other on the ventral third of the flanks and the third, which is
smallest and virtually imperceptible, on the dorsal third of the flanks.

The suture line is very incised with trifid L, U and I lobes and bifid saddles.

Dimensions
Wb. Ae)
(at distal end)
Wh. 52,9.

Discussion

This specimen corresponds approximately to Casey’s Lithancylus grandis
(Sowerby) specimen figured in his plate 20, fig. 1b, where ornament on the shaft
changes from normal to trituberculate ribbing. L. grandis, however, has an
almost circular whorl section on the shaft and is not conspecific. L. fustis has a
depressed whorl section; L. neleus (Anderson), L. mirabilis (Anderson) and
L. australe Day have ovoid whorl sections. L. cyclopius (Anderson) has a
subtrigonai section as the present specimen, but finer ribbing, as has L. tirolensis
Casey.

CRETACEOUS FAUNAS FROM SOUTH AFRICA DSS

Thomson’s (1974) Lithancylus sp. is poorly preserved, but appears to have
finer ribbing at the proximal end.

Associated shafts of Ancyloceras (Adouliceras) spp. are all more massive
and have finer ribbing.

Occurrence
Lower Aptian of southern Mozambique.

Fig. 25. Lithancylus sp. SAM-—PCM5436 from Lubemba, Mozambique, Lower Aptian. x 1,2.

256 ANNALS OF THE SOUTH AFRICAN MUSEUM

Genus Tropaeum J. de C. Sowerby, 1837
Type species
Tropaeum bowerbanki J. de C. Sowerby, 1837 from the Lower Aptian of
England, by monotypy.

Diagnosis

Generally large; coiling predominantly crioceratitid, but in early species
the body chamber may be distinctly uncoiled, giving rise to ancyloceratid
or to aspinoceratid forms. Ornament on phragmocone consists of plain ribbing
with occasional umbilical or ventral tubercles in early or late stages. On the body
chamber a rapid change in ornament may take place, with the development of
heavy, distantly spaced ribs. Aperture may be contracted. Dorsum ornamented
by forward directed striae only.

Discussion

For details on the somewhat unusual manner in which the genus was
introduced, and the accompanying nomenclatorial implications, see Casey
(1960: 24), who also provides the most complete discussion of the genus.
Additional information is provided by Day (1974: 5) who emended the diagnosis
to accommodate loosely coiled forms and those in which no abrupt change in
ornament takes place on the body chamber.

The best documented occurrences of Tropaeum are in England, the Caucasus ~
and transcaspian regions of the U.S.S.R., and Australia. Recent descriptions of
the English and Australian faunas are available, but for descriptions of the
Russian species we are still heavily dependent on the older works of Sinzow
(1872, 1905).

Together with the puzosiids, e.g. P. seppenradensis (Landois) and
baculitids, e.g. Baculites rex Anderson and Eubaculites latecarinatus Brunn-
schweiler (unpublished data on Zululand specimens), Tropaeum are amongst the
largest Cretaceous ammonites. FGérster 1975a: 153) recently described a specimen
of Tropaeum subsimbirskense compressum (Sinzow) from the Aptian of Mozam-
bique with a diameter of over 900 mm. Tropaeum imperator Howchin &
Whitehouse (1928: 487, figs 144-145) is reported to be over 770 mm in diameter
(Casey 1960: 41). A body chamber fragment from Zululand to be described
below measures c. 730 mm, and a virtually complete specimen 700 mm. The
large size, and loose crioceratitid or even ancyloceratitid coiling of the genus
suggests that Tropaeum may have been a poor swimmer and led a rather
sluggish life.

With the exception of some species which may possess tubercles in the early
or late stages, e.g. Tropaeum imperator Howchin & Whitehouse, the genus is
easily distinguished from the other ancyloceratids by the general lack of
tubercles.

Within the genus, three species groups, based on the coiling of the body
chamber, may be recognized:

CRETACEOUS FAUNAS FROM SOUTH AFRICA 2)

(i) Tropaeum gr. ex. hillsi J. de C. Sowerby
(ii) Tropaeum gr. ex. bowerbanki J. de C. Sowerby
(iii) Tropaeum gr. ex. subarcticum Casey

In each of these groups, successive species show parallel evolution of the
outer whorls and body chamber with ancyloceratid, aspinoceratid and crio-
ceratitid coiling respectively. This is a parallel development to that encountered
in Australiceras, to be discussed below, and, as demonstrated by Casey (1960:
20, text-fig. 5), is one of the several examples of recoiling seen in ancyloceratids.

Specific differentiation amongst Tropaeum species is rather unsatisfactory,
but is based primarily on the coiling of the outer whorls and body chamber,
whorl section and ornamentation of the adult body chamber. Specific identifi-
cation of fragments is usually virtually impossible. Species to be referred and
possibly belonging to Tropaeum are as follows:

1. Tropaeum? aegoceras (von Koenen) (1902: 328, pl. 36 (fig. la—c)).

2. Tropaeum arcticum (Stolley) (1912: 16, pl. 2 (fig. 1)).

3. Tropaeum australe (Moore) (1870: 257, pl. 15 (fig. 3)).

4. Tropaeum benstedti Casey (1960: 37, pl. 3 (fig. 3), text-fig. 11g).

5. Tropaeum bowerbanki J. de C. Sowerby (1837: 535).

6. Tropaeum bowerbanki Sow. var. densistriatum Casey (1960: 30, pl. 6 (fig. 1)).
7. Tropaeum caseyi Collignon (1962: 18, pl. 222 (fig. 965)).

8. Tropaeum drewi Casey (1960: 35, pl. 8 (figs 1—2)).

9. Tropaeum hillsi (J. de C. Sowerby) (1836: 128).

10. Tropaeum imperator Howchin & Whitehouse (1928: 487, figs 144-145).

11. Tropaeum keepingi Casey (1960: 41, text-figs lle, 13).

12. Tropaeum? lamprum (Etheridge Jun.) (1909: 157, pl. 48 (figs 1-2)).
(According to Day (1974: 12) the holotype of this species is distinctly
trituberculate and thus an Australiceras.)

13. Tropaeum leptum (Etheridge Jun.) (1909: 143, pl. 30 (figs 1-3), pl. 34 (fig. 2)).

14. Tropaeum? multicingulatum (Von Koenen) (1902: 314, pl. 34 (fig. 2)).

15. Tropaeum rossicum Casey (1960: 25).

16. Tropaeum simbirskense (Sinzow) (1872: 33, pl. 5 (figs 2-5), pl. 6 (figs 1-4)).

17. Tropaeum subarcticum Casey (1960: 40, pl. 8 (fig. 3a-c), pl. 10 (fig. 1),
text-figs 11d, 12).

18. Tropaeum subsimbirskense subsimbirskense (Sinzow) (1905: 320, pl. 22
(figs 3—-5)).

19. Tropaeum subsimbirskense compressum (Sinzow) (1905: 230, pl. 22 (figs 1—2)).

20. Tropaeum undatum Whitehouse (1926: 215).

Occurrence

Tropaeum occurs in the Aptian of England, northern Germany, the
caucasian and transcaspian regions of the U.S.S.R., Bulgaria, Spitzbergen,
eastern Greenland, arctic Canada, California, Japan, Patagonia, Falkland
Island Dependencies?, Australia, Madagascar, Mozambique and Zululand.

258 ANNALS OF THE SOUTH AFRICAN MUSEUM

Tropaeum sp. aff. subsimbirskense subsimbirskense (Sinzow, 1905)
Figs 26, 27A-B
Compare:
Crioceras subsimeirskense (sic) var. inflata Sinzow, 1905: 320, 330, pl. 22
(figs 3-5).
? Tropaeum subsimbirskense compressa: Dimitrova, 1967: 63, pl. 12 (fig. 1).

Material

SAS EM 109 from Manyola Drift, northern Zululand (26°50’20’S
3273.0" E):

Description

Coiling is very involute, with an umbilical diameter of 33,5 per cent. The
whorl section is rounded subtriangular with a flat dorsum, sharp umbilical edge
and very little-inflated flanks converging to a sharp, narrow venter. The body
chamber remains in contact with the phragmocone.

Ornament on the phragmocone consists of fine, sinusoidal ribbing. The ribs
are faint on the dorsum and curve broadly forward. At the umbilical edge they
curve sharply backward, forming a slight elevation, and then cross the flanks in
sinusoidal fashion. Bifurcations and intercalations occur quite frequently at
midflank or ventral thereof. On the nucleus, there are approximately seventy
ribs per whorl. Towards the body chamber ornament gradually becomes
coarser. On the earliest part of the body chamber virtually every rib bifurcates
on the ventral quarter of the flanks, thus forming loops over the venter. There-
after intercalated ribs arising on the ventral half of the body chamber replace the
bifurcating ribs. On the last part of the body chamber, only strong, relatively
high-crested ribs are present, separated by much wider interspaces. Each of these
ribs has a prominent forward curvature over the venter. The suture line is very
incised, with a huge lateral lobe.

Dimensions

Specimen D Wb Wh Wb/Wh U
SAS EM 109 305 95(31,6) 112(37,2) 0,85 102(33,5)

Discussion

Casey (1960: 39 footnote) remarked that one of Sinzow’s varietal names,
inflata or compressa was superfluous, and subsequently regarded the variety
inflata as representing 7. subsimbirskense s.s. with Sinzow’s (1905, pl. 22
(figs 4-5)) specimen as lectotype.

It should be pointed out here that the original spelling in the Russian text
(Sinzow 1905: 320) is subsimeirskense, and subsimbirskense in the German
résumé. The species is named after the Russian district of Simbirsk, and the
name in the Russian text is obviously a typographical error. The valid name is
thus subsimbirskense.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 259

T. subsimbirskense subsimbirskense differs from the subspecies compressum
in having a more inflated and depressed whorl section and a smaller umbilical
diameter. Ribbing on the nominal subspecies also appears to be more sinusoidal
when compared to the rursiradiate to rectiradiate ribbing of the subspecies
compressum.

Dimitrova’s (1967: 63, pl. 12 (fig. 1)) specimen is more suitably placed in
the nominal subspecies rather than in 7. subsimbirskense compressum.

Fig. 26. Tropaeum aff. subsimbirskense subsimbirskense (Sinzow, 1905). SAS EM 109 from
Manyola Drift, Zululand, Aptian III. x 0,3.

260 ANNALS OF THE SOUTH AFRICAN MUSEUM

~ AO ii,

om

Fig. 27. A-B. Tropaeum aff. subsimbirskense subsimbirskense (Sinzow, 1905). SAS EM109
from Manyola Drift, Zululand, Aptian III. x 0,34 and 0,55 respectively.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 261

The Zululand specimen differs from 7. subsimbirskense subsimbirskense
mainly in not being quite as inflated and in lacking the strong major costae with
intermediaries on the body chamber, and is hence referred to the Russian species
in open nomenclature. The umbilical diameter of the Zululand specimen is
intermediate between that of Sinzow’s two figured specimens.

In the lack of a sudden change of ornament on the body chamber, the
present species is similar to the Australian Tropaeum undatum Whitehouse 1926
(see Day 1974: 6, pl. 3 (fig. 2a—b)). The latter species has, however, a subquadrate
whorl section and much stronger ribbing.

Occurrence
Aptian III of Zululand.

Tropaeum subsimbirskense compressum (Sinzow, 1905)
Figs 28, 29A—-B, 87

Crioceras subsimeirskense var. compressa Sinzow, 1905: 320, pl. 22 (figs 1-2) (error).

Tropaeum mozambiquense: Da Silva, 1962: 24, pl. 9 (figs 1-2), pl. 12, non pl. 10, pl. 11, pls 13-14.

? Australiceras cf. gigas: Wachendorf 1967: 283 pl. 34 (fig. 4).

? Ammonitoceras tovilense: Wachendorf, 1967: 282, pl. 36 (fig. 1).

Tropaeum subsimbirskense compressum: Forster, 1975a: 152, pl. 2 (figs 3-6), text-fig. 29.
Non Dimitrova, 1967: 63, pl. 12 (fig. 1).

Lectotype

Sinzow’s (1905, pl. 22 (fig. 1)) specimen is herein designated lectotype. So
far as can be determined, Sinzow did not designate a holotype. Dimitrova
(1967: 63) stated that Sinzow’s plate 22, figure 1 specimen is the type, but this
cannot be taken as a lectotype designation.

Material
SAS EM 110 from Manyola Drift, Zululand. Aptian III.

Description

Coiling is closely crioceratitid with no sign of uncoiling of the body
chamber. The innermost whorls, up to a diameter of 75 mm, are not preserved.
At a diameter of 160 mm the whorl section is subtriangular with a flat to concave
dorsum, a sharp umbilical edge, very little-inflated flanks and a narrow venter.
Apart from the fact that the whorl height increases more rapidly than the whorl
breadth, the whorl section remains basically subtriangular up to the body
chamber. On the body chamber the umbilical wall becomes more rounded, the
umbilical edge less acute and the flanks more inflated. The whorl section
becomes subrectangular at this stage.

On the inner whorls, ornament consists of single, low, rounded rursiradiate
to radial ribs separated by interspaces of the same width. At a diameter of
approximately 300 mm occasional stronger ribs occur, separated by three to five

262 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 28. Tropaeum subsimbirskense compressum (Sinzow, 1905). SAS EM110 from Manyola
Drift, Zululand, Aptian III. x 0,25.

normal ribs which may bi- or trifurcate at the umbilical edge. Towards the end
of the phragmocone the major ribs become stronger. The beginning of the body
chamber is marked by a very strong, low broad rib. From here onward, as far as
the aperture, ornament consists solely of ten widely spaced, initially low, broad
ribs; the intercalatories having disappeared completely. Towards the aperture,
the last four or five major ribs become progressively flared and narrower on the
flanks, and also closer spaced. The first three to four major ribs on the body
chamber are restricted to the flanks, but towards the aperture, as they become
increasingly flared, they bifurcate at the ventral edge and cross the venter as

CRETACEOUS FAUNAS FROM SOUTH AFRICA 263

B

Fig. 29. A-B. Tropaeum subsimbirskense compressum (Sinzow, 1905): SAS EM110 from
Manyola Drift, Zululand, Aptian III. x 0,34.

264 ANNALS OF THE SOUTH AFRICAN MUSEUM

very low, broad looped folds. The last three or four ribs before the aperture are
the most prominent, being thin and flared on the flanks and broad and high over
the venter.

Part of the aperture is preserved. It appears to have been simple, and
somewhat constricted.

The suture is very incised and dentritic, consisting of a trifid L, U and I.
The lateral lobe is largest and occupies virtually the whole of the flank (see
Fig. 87).

Dimensions
Specimen D Wb Wh Wb/Wh U
SAS EM110 510 140(27) 151(29) 0,93 222(43)
390 109(27) 139(35) 0,80 163(44)
Discussion

The main characteristic features of the phragmocone of this species are the
whorl section, with little-inflated flanks, converging to a narrow dorsum (see
Forster 1975a, text-fig. 29), and the more or less straight, rursiradiate ribbing.
On the body chamber, the low, major costae, separated by wide interspaces and
devoid of intercalatories are equally characteristic.

The lectotype (Sinzow 1905, pl. 22 (fig. 1)) specimen does not have quite as
prominent major costae as the Zululand or Mozambique material, but it appears
that only a part of the body chamber is preserved. The last part of the outer
whorl on the lectotype also is slightly detached. Illustrations of 7. bowerbanki
in Casey 1960 (compare pl. 4 (fig. 1) and pl. 5 (fig. 1)), however, show that there
is some degree of variation in the tightness of coiling of the outer whorl in
Tropaeum, and that it can be included in the range of intraspecific variation.

The closest European ally to this species is Tropaeum bowerbanki J. de C.
Sowerby, from which the present form appears to have been derived. The major
differences are that the flanks are more convergent on the phragmocone in
T. subsimbirskense compressum than in T. bowerbanki, whilst the body chamber
of T. bowerbanki is also more robust; the ribs are flared and high-crested,
compared to the low, broad ribs in T. subsimbirskense compressum.

As yet no Malagasy equivalent of this species has been described.

Forster (1975a) has shown that specimens occur with a diameter in excess
of 900 mm, surpassing all other records of large Tropaeum.

Occurrence
Upper Aptian of Mangyschlak, southern Russia, Bulgaria?, Mozambique
and Zululand.
Tropaeum rossicum Casey, 1960
Figs 30, 31B

Ancyloceras gracilis, Sinzow, 1872: 35, pl. 6 (figs 5S—6, 10-11).
Crioceras gracile Sinzow, 1905: 306, pl. 17 (figs 1-4), pl. 18 (fig 6), pl. 19 (fig. 1).
Tropaeum rossicum Casey, 1960: 43.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 265

Material

SAM-PCZ5681 from an unknown locality in Zululand.
Holotype

Sinzow’s (1905, pl. 18 (fig. 1)) specimen from Stepnaja, Simbirsk.
Description

The available specimen consists of just over one whorl. Coiling is loose,
with the whorls just touching but not impressed. The whorl section is ovoid,

"eae j
+ (a ea

Fig. 30. Tropaeum rossicum Casey, 1960. SAM-—PCZ5681 from an unknown locality in
Zululand. x 0,57.

266 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 31. A. Australiceras wandalina (Boshoff MS) sp. nov. SAS EM106 from Manyola Drift,
Zululand, Aptian III. x 0,69. B. Tropaeum rossicum Casey, 1960. SAM-—PCZ5681 from an
unknown locality in Zululand. x 0,6.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 267

higher than wide on the early part of the whorl, but becomes more rounded on
the outer part. Ornament consists of approximately 100 strong, rursiradiate,
rounded ribs, separated by interspaces of equal width. Initially, ribbing is quite
prominent on the umbilical wall, but eventually becomes quite faint at the
largest diameter preserved. The ribs cross the flanks and venter without
interruption.

Discussion

Casey (1960: 43) drew attention to the fact that Ancyloceras gracile Sinzow,
1872 was a homonym of Ancyloceras gracile Oppel in Zittel 1868 and that a
new name was required. Being uncertain as to whether Sinzow’s 1872 (pl. 6
(figs 5—6, 10-11)) immature specimens of Ancyloceras gracile were conspecific
with his (Sinzow’s) 1905 specimens of Crioceras gracile, Casey designated
Sinzow’s (1905 pl. 18 (fig. 1)) specimen as holotype of Tropaeum rossicum.

With only one specimen available, it is impossible to comment on the
relationship of Sinzow’s 1872 and 1905 specimens, and the species is here
interpreted on the basis of the 1905 specimens. Sinzow (1905, pl. 19 (fig. 1))
illustrated a specimen with finer ribbing with which the Zululand specimen
compares well. The whorl section provided by Sinzow (1905: 308) also corre-
sponds to that of the present specimen. No sign of ventral tuberculation is
present, but according to Sinzow this only is present in the early and late
stages.

T. rossicum is easily distinguished by the ovoid whorl section on the
phragmocone, the strong, rounded and relatively straight, rursiradiate ribbing.
T. sp. aff. undatum Whitehouse, to be described below, has similar ornament on
the inner whorls, but shows a rapid increase of whorls and a square whorl
section. T. drewi Casey (1960: 35, pl. 8 (figs 1-2), pl. 9 (fig. 1), text-figs 10, 11f)
has a similar rib density (c. 90 at 150 mm diameter), but ribbing is more or less
radial and the whorl section compressed.

Tropaeum caseyi Collignon (1962: 18, pl. 222 (fig. 965)) has a similar, but
much coarser style of ribbing.

By the possession of tuberculation in the early and late stages, 7. rossicum
occupies an intermediate position between Tropaeum and Australiceras. Casey
(1960: 25) refers to 7. rossicum as being intermediate between T. hillsi and
A. gigas, both forms with a body chamber terminating in a straight shaft and
hook. It is not quite clear if this is meant to imply a hook-shaped body chamber
in T. rossicum. Sinzow’s figures and his comparisons with T. bowerbanki indicate,
rather, that the coiling was aspinoceratid or crioceratitid.

Occurrence

T. rossicum is only known from the Aptian of Simbirsk and Zululand.
Casey recorded Tropaeum cf. rossicum from the Upper Aptian martini Zone of
Maidstone, Kent.

268 ANNALS OF THE SOUTH AFRICAN MUSEUM

Tropaeum dayi sp. nov.
Figs 32-34
Tropaeum cf. australe: Day, 1974: 5, pl. 1 (fig. la—b), pl. 2 (fig. 2a—b).

Holotype

SAS EM109b from Manyola Drift, northern Zululand, Aptian III.
Collected by the late Mr E. Meyer.

Derivation of name
Named after Dr R. Day, Geological Survey of Queensland, Brisbane.

Description

The innermost whorls are not preserved. Coiling is open, crioceratitid, with
the whorls not touching. The body chamber becomes a little detached. The
whorl section is compressed with a rounded dorsum lacking a zone of impression,
a broadly rounded umbilical edge, weakly inflated flanks and a broadly rounded
venter. Ornament on the early whorls consists of delicate, thin ribs, separated
by wider interspaces. The ribs are straight to sinuous on the flanks and cross the
dorsum and venter with a forward curvature. Rib density is c. 100 at a diameter
of 150 mm. With increasing diameter ribbing becomes coarser, but there is no
abrupt change in ornament on the body chamber. On the body chamber of the
holotype, of which half a whorl is preserved, twenty-six ribs occur per half
whorl. Ribbing on the body chamber is broad and low, with very faint traces of
tuberculation on the last few ribs. The aperture is unknown.

The suture is minutely frilled. The lateral lobe is largest, asymmetrical and
occupies virtually the whole of the flanks (see Fig. 34).

Dimensions
Specimen D Wb Wh Wb/Wh U
SAS EM 109b 285 110(28,5) 140(36,3) 0,78 157(40,7)
Discussion

Apart from slight differences in relative proportions, the Zululand specimen
and the Australian specimen figured by Day (1974: 5, pl. 1 (fig. la—b), pl. 2
(fig. 2a—b)) as Tropaeum cf. australe are virtually identical. Ornament on the
inner and outer whorls is the same and both lack a sudden change in ornament
at the beginning of the body chamber. The loose coiling is conspicuous in both
forms and serves to distinguish it from T. australe (Moore). T. australe (Moore)
is a poorly-known species, but from the available illustrations (Moore 1870,
pl. 15 (fig. 3); Etheridge Jun. 1892, pl. 31 (fig. 1), pl. 32 (figs 3-4)) it can be seen
that ornamentation and whorl section are similar to those of the present species,
but that the coiling is closer with the whorls touching.

Australiceras rabenjanaharyi Collignon (1962: 26, pl. 226 (fig. 969)) from
Madagascar, has similar ornament after the initial trituberculate stage, but is
also tightly coiled with the whorls touching.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 269

Tropaeum rossicum Casey, 1960 has a similar whorl shape but ribbing is
much coarser and ornament on the body chamber differs from that of T. dayi
sp. Nov.

The presence of faint tubercles on the inner whorl of this species (Day,
1974: 5) indicates that it is somewhat intermediate between Tropaeum and
Australiceras, as is the case in T. rossicum.

Occurrence

The Australian specimens are from the Aptian of the Eromanga and Surat
basins. The Zululand specimen is from the authors’ Aptian III.

Fig. 32. Tropaeum dayi sp. nov. SAS EM109b, holotype from Manyola Drift, Zululand.
Aptian III. x 0,32.

270 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 33. A-B. Tropaeum dayi sp. nov. SAS EM109b, holotype from Manyola Drift, Zululand,
Aptian III. x 0,42.

CRETACEOUS FAUNAS FROM SOUTH AFRICA Din

Fig. 34. Tropaeum dayi sp. nov. SAS EM109b. Suture line of holotype. x 1.

Tropaeum obesum sp. nov.

Figs 35-37, 38D, 39-42, 65E
Holotype
SAM-PCZ5681 from locality 34, Zululand. Aptian IV. Housed in the
South African Museum. Collector unknown.

Material

The holotype and SAS A4000 from locality 34, Aptian IV, and numerous
fragments from locality 168, Aptian IV, including BMNH C7885. BMNH
C78886—C8887 are from the same horizon at locality 171.

Description

Very large, coiling is crioceratitid with the whorls touching and with a
dorsal zone of impression. The body chamber remains in contact with the
phragmocone. The whorl section is initially subtrigonal, slightly higher than
wide with rounded umbilical edges, moderately inflated flanks and a narrow
rounded venter. With increasing diameter the whorl section becomes more

272 ANNALS OF THE SOUTH AFRICAN MUSEUM

inflated and rounded. On the body chamber, which consists of about half a
whorl in the holotype, the section becomes depressed.

Ornament is variable, but consists mostly of slightly sinusoidal ribs on the
phragmocone. They are generally inclined backward, but may on occasion be
radial. Over the dorsum the ribs are reduced to striae with a strong forward
curvature. With increasing diameter ribbing becomes coarser, and the body

Fig. 35. Tropaeum obesum sp. nov. SAM-PCZ5681, holotype from locality 34, Zululand,
Aptian IV. Area between X—X restored. x 0,22.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 21S

Fig. 36. Tropaeum obesum sp. nov. SAM-PCZ5681 holotype from locality 34, Zululand,
Aptian IV. Restored part removed. x 0,26.

274 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 37. Tropaeum obesum sp. nov. SAS A4000 from locality 34, Zululand, Aptian IV. Inner
whorls up to X sculpted in matrix. x c. 0,5.

CRETACEOUS FAUNAS FROM SOUTH AFRICA Df)

Fig. 38. A-B. Australiceras sp. indet. A. SAS L3 from locality 170, Zululand, Aptian I-II. x 1.

C. Tonohamites aequicingulatus (von Koenen, 1902). SAS Z8/Tal from locality 168, Aptian

IWI-IV. x 1,5. D. Tropaeum obesum sp. nov. SAS A4000 from locality 34, Zululand, Aptian
We x c310:S;

276 ANNALS OF THE SOUTH AFRICAN MUSEUM

A B

Fig. 39. A-B. Tropaeum obesum sp. nov. BMNH C78886, fragment from locality 171, Zululand,
Aptian IV. x 1.

207

CRETACEOUS FAUNAS FROM SOUTH AFRICA

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puelninZ “TLT Aypeoo] wory yuowsely ‘73898879 HNWA ‘Aou ‘ds uinsaqgo Unavdo.], *-V ‘Op ‘314

278

Fig. 41. Tropaeum

ANNALS OF THE SOUTH AFRICAN MUSEUM

Be

obesum sp. nov. BMNH C78885 from locality 168, Zululand, Aptian IV.
x 0,9.

sy

CRETACEOUS FAUNAS FROM SOUTH AFRICA 279

Fig. 42. Tropaeum obesum sp. nov. BMNH C788835 from locality 168, Zululand, Aptian IV.
x 0,9.

pee nae =" & “Fe ££ fe 'Y

280 ANNALS OF THE SOUTH AFRICAN MUSEUM

chamber bears about twelve heavy, laterally flared, sharp-crested ribs. Over the
venter, these ribs are broad and low. Major ribs are also absent on the dorsum
~ and only striae occur.

The suture line is characterized by thin-stemmed saddles and a large
asymmetrically trifid lateral lobe.

Dimensions

Specimen D Wb Wh Wb/Wh U
SAM-PCZ5681 630 250(39,9) 210(33,4) 1,18 260(41)

Discussion

Numerous fragments of Tropaeum are tentatively referred to this species
which is characterized by its rounded adult whorl section and generally rursi-
radiate sinusoidal ribs.

Tropaeum imperator Howchin & Whitehouse (1928: 487, figs 144-145) is a
similar large species with comparable ornament on the phragmocone. The whorl
section on the final whorl, however, is subquadrate, compared to rounded
depressed in the present species. Apparently, coiling in 7. imperator is loosely
crioceratitid, whereas that of 7. obesum sp. nov. is tight, with the inner whorls
touching.

Tropaeum percostatum (Gabb) (1864: 77, pl. 16 (fig. 26), pl. 17 (fig. 26a));
(see also Anderson, 1938: 212, pl. 71 (fig. 1), pl. 72 (fig. 1) is a related species
with denser, lower ribbing on the body chamber. The whorl section of the latter
species is subquadrate, compared to rounded-depressed in T. obesum. The body
chamber of 7. percostatum is said to become slightly detached.

Crioceras aegoceras von Koenen (1902: 328, pl. 36 (figs la-c, 2-3)) has
similar ornament on the outer whorl, but coarser costation on the inner whorls
than T. obesum sp. nov., and is only doubtfully referred to Tropaeum.

Occurrence
Upper Aptian of Zululand.

Tropaeum sp. aff. undatum Whitehouse, 1926
Figs 43-44

Compare:
Tropaeum undatum: Jeletzky, 1964: 68, pl. 20 (fig. 2). Hill, Playford & Woods,

1968: K16, pl. K7 (fig. 3). Day 1974: 6, pl. 3 (figs 2a—b).
Tropaeum rarum Whitehouse, 1926: 216, pl. 36 (fig. 1a—b).
Material

SAS UMS1 from locality 34, Mzinene River. Aptian IV.

Description

In the Zululand specimen parts of three whorls are still in contact. Coiling
is Crioceratitid with the whorls just touching. On the innermost whorl the section

CRETACEOUS FAUNAS FROM SOUTH AFRICA 281

is square, but with increasing diameter soon becomes subtrigonal and higher
than wide. The dorsum is flat, with a slight dorsal zone of impression on the last
whorl, rather sharp umbilical edges and little inflated flanks converging to a
broadly rounded venter.

On the innermost whorl still preserved, ornament consists of very strong,
broad, slightly rursiradiate single ribs; approximately five in a distance equal to
the whorl height. On the succeeding whorl ornament becomes finer, the ribs are
weak over the dorsum, crossing it with a slight forward flexure, then passing
backwards over the umbilical wall and flexing backwards over the flanks. A few
bifurcations occur on the dorsal third of the flanks. Eight ribs occur in a distance
equal to the whorl height on the second preserved whorl, and on the last whorl
ribbing is very coarse and widely spaced, with five to six ribs per whorl height.

Fig. 43. Tropaeum sp. aff. undatum Whitehouse, 1926. SAS UMS1 from locality 34, Mzinene
River, Zululand, Aptian IV. x 0,52.

amnmm a t= 6 ST

282 ANNALS OF THE SOUTH AFRICAN MUSEUM

Discussion

Tropaeum rarum Whitehouse, 1926 was included in the synonomy of
T. undatum by Day (1974: 6, 7). The Zululand specimen is tentatively referred to
the Australian species because of the similar coarse ribbing on the innermost
whorls, followed by slightly sinuous ribbing, together with a rapid increase in
whorl diameter. The only obvious difference is that the whorl section of the
Australian material is square to depressed, compared to the compressed, higher
than wide outer whorl of the Zululand specimen.

Crioceras australe Waagen, 1875 non Moore, is superficially similar as

Fig. 44. Tropaeum sp. aff. undatum Whitehouse, 1926. SAS UMS1 from locality 34, Mzinene
River, Zululand, Aptian IV. x 0,58.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 283

regards the rapid increase in whorl diameter, but is, of course, generically
distinct, belonging to Australiceras.

The rapid increase in whorl diameter and the coarse ribbing of inner and
outer whorls, together with finer ribbing during middle growth stages, clearly
distinguish this species from the other Zululand Tropaeum.

Occurrence
Upper Aptian, Aptian IV of Zululand.

Tropaeum sp. gr. ex bowerbanki (J. de C. Sowerby) var.
densistriatum Casey—T. hillsi (J. de C. Sowerby)

Fig. 45
Compare:

Tropaeum bowerbanki (J. de C. Sowerby) var. densistriatum Casey, 1960: 30,

pl. 6 (fig. 1).
Tropaeum hillsi: Casey, 1960: 30, pl. 7, text-figs 8-9, 11c.

Material

SAS L3 from locality 170 Mlambongwenya Creek, Zululand. Lower Aptian,
Aptian I or II.

Description

One specimen consisting of parts of two embracing whorls is available.
Coiling is open, crioceratitid, and the whorls are not in contact. The specimen is
slightly crushed, but the whorl section appears to have been rounded-
subrectangular, with a rounded venter narrower than the dorsum. The flanks
and venter are ornamented by approximately fifty radial ribs per half whorl.
The dorsum is smooth and devoid of ribbing.

Discussion

The specimen is from the horizon of abundant Ancyloceras (Adouliceras)
spp. The absence of tuberculation on the inner whorl, however, clearly shows
that it is a Tropaeum.

The whorl section and density of ribbing are the same as those of Tropaeum
bowerbanki var. densistriatum (Casey 1960: 31, pl. 6 (fig. 1)) and Tropaeum hillsi
(J. de C. Sowerby) (see Casey 1960: 31, pl. 7, text-figs 8-9, llc). The inner
whorls of these two species are indistinguishable (Casey 1960: 33). For the
present it is therefore only possible to refer to this specimen as Tropaeum sp. gr.
ex. T. bowerbanki var. densistriatum—T. hillsi. As Casey has shown (1960: 33)
the two species are very close and connected by transitional forms. Because of
differences in age, however, he maintained the two species apart.

Tropaeum subsimbirskense (Sinzow) (1905: 320, pl. 22 (figs 1-5)) has
similar inner whorls, but the flanks are strongly convergent to the venter.

284 ANNALS OF THE SOUTH AFRICAN MUSEUM

Occurrence

T. hillsi occurs in the Lower Aptian, top of the deshayesi Zone in England.
T. bowerbanki var. densistriatum is ubiquitous in the bowerbanki Zone of Kent,
England.

Fig. 45. Tropaeum sp. gr. ex bowerbanki var. densistriatum-hillsi. SAS L3 from locality 170,
Zululand, Aptian I or II. x 1,1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 285

Tropaeum sp. indet.

Fig. 46
Material

SAS Z804 from locality 151, Mkuze Game Reserve, Aptian IV.

Description and discussion

A massive body chamber fragment, approximately 730 mm long, is of
interest in that it is filled with abundant small tonohamitids (to be described
below).

The whorl section is wider than high with a slight dorsal zone of impression.
Ornament consists of about fifteen heavy, sharp ribs. Ribbing is restricted to the
flanks and venter. The dorsum is ornamented by forwardly directed striae, not
corresponding in number to the lateral ribs.

Although the inner whorls are not preserved the stratigraphic horizon is
one that is characterized by Tropaeum with crioceratitid coiling, and the body
chamber more or less in contact with the rest of the phragmocone.

The present specimen may be compared with Tropaeum subarcticum Casey
(1960: 40, pl. 8 (fig. 3a—c), pl. 10 (fig. 1), text-figs 11d, 12) which also grows to
great size. Ribbing on 7. subarcticum is denser, however, and some of the ribs
on the body chamber illustrated by Casey (1960: 40, text-fig. 12) are looped,
bifurcating near the venter.

Tropaeum imperator Howchin & Whitehouse (1928: 487, figs 144-145),
another gigantic species from the Aptian of Australia, has much sharper, and
more distantly spaced ribbing on the body chamber. Tropaeum obesum sp. nov.
has a more inflated whorl section and more distantly spaced ribs on the body
chamber.

Genus Australiceras Whitehouse, 1926
(= Colombiaticeras Royo y Gomez, 1945)
Type species
Crioceras jacki Ethridge Jun., 1880 from the Aptian of eastern Australia
by original designation Whitehouse (1926: 208).

Diagnosis

Coiling ancyloceratid or aspinoceratid in early Aptian species, but
crioceratitid in later ones. Early whorls not always in one plane; ornamented by
trituberculate ribs separated by a variable number of intermediaries. Tuber-
culation ceases at variable diameters in middle stages of growth and may or
may not reappear on the body chamber.

Discussion

Whitehouse originally introduced Australiceras for crioceratitid forms only,
a view followed by Wright (1957: L211). Latterly, the genus has been taken to

286 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 46. Tropaeum sp. indet. SAS Z804 from locality 151, Zululand, Aptian IV. Large body
chamber fragment with last septum preserved and filled with numerous specimens of Tono-
hamites? caseyi sp. nov. Photograph by courtesy of Tony Harris (Salisbury). xX c. 0,24.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 287

include the early Aptian representatives of the lineage with ancyloceratid
coiling. The extent and variability of the early Aptian representatives of the
genus in the English Lower Greensand was exhaustively reviewed by Casey
(1960, 1961), as was the scope of the Australian late Aptian representatives by
Day (1974). Unfortunately, however, little or no work has been done on the
Central European or Soviet material since the beginning of the century.

Spath (1931: 656) advocated that the name Australiceras be abandoned in
favour of Tropaeum, due to the difficulty encountered in deciding whether or not
tuberculation was present, especially in fragmentary material. Transitional
forms do occur, for example Australiceras pingue Casey, T. rossicum Casey, and
T. imperator Howchin & Whitehouse, but both genera are accepted as valid taxa.
For a further discussion on the differences between Tropaeum and Australiceras,
see Casey (1960: 40, 1961: 45). Day’s (1974: 8) assertion that the saddles of the
suture lines of the Australian representatives of Australiceras tend to be broader
than those in Tropaeum is also true to a certain extent in the Zululand material,
and it appears that in some late Aptian species the saddles tend to become
broader, accompanied by a general simplification of the suture line.

Differences between the homoeomorphic Ancyloceras (Adouliceras) and the
early uncoiled forms of Australiceras are discussed elsewhere (p. 226).

As in Tropaeum, three species groups are recognized, based on the coiling.

(i) Australiceras gr. ex. gigas (J. de C. Sowerby)
(ii) Australiceras gr. ex. tuberculatum (Sinzow)
(iii) Australiceras gr. ex. jacki (Etheridge)

Specific differentiation is a moot point, as it is in Tropaeum. Virtually all the
species described from the Eurasian, Australian, North American, South
American and Madagascan regions appear to be endemic to these regions, and
either based on unique type specimens, or, if based on large collections, are
interpreted so widely that differences between the various species are difficult to
formulate, all of which suggests a superfluity of specific names, and high
intraspecific variability.

The diameter at which tuberculation ceases is variable, as has been
illustrated in A. gigas (see Casey 1961: 47) and A. jacki (see Day 1974: 9).
Rib density and whorl section in the early stages seem to be equally unreliable
specific characters in A. jacki as shown by Day (1974: 9) and also here in
Figures 47-49, based on Australian material housed in the British Museum
(Natural History).

The only apparent differences of use in specific separation are the coiling
of the body chamber, whorl section, and rib density in the adult stage together
with, to a lesser degree, strength of tuberculation and the number of inter-
mediaries in the early stages.

Species to be referred to Australiceras are as follows:

1. Australiceras argus Anderson (1938: 211, pl. 70 (figs 1—2)).
2. Australiceras australe (Waagen non Moore) (1875: 246, pl. 60 (fig. 1)).

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288 ANNALS OF THE SOUTH AFRICAN MUSEUM

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Downs, Queensland, Australia. x 1.

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CRETACEOUS FAUNAS FROM SOUTH AFRICA 289

. 48. Australiceras jacki (Etheridge Jun., 1880). BMNH C40760 from the Aptian of Rolling
Downs, Queensland, Australia. Note the pathological body chamber. x 1.

Australiceras bolivari (Royo y Gomez) (1945: 470, pl. 75 (fig. 2)).
Australiceras ? bulgaricum Dimitrova (1967: 62, pl. 30 (fig. 1)).

? Crioceras deeckei Favre (1908: 636, pl. 36 (fig. 4), pl. 37 (fig. 1), text-fig. 7).
? Crioceras carinato-verrucosum Sinzow (1905: 316, pl. 21 (figs 1-2)).
Australiceras gigas (J. de C. Sowerby) (1828: 188, pl. 593 (fig. 2)).
Australiceras gigas (Sow.) var. anguimanum Casey (1961: 52, pl. 13).
Australiceras gigas (Sow.) var. arcuatum Casey (1961: 52, pl. 12 (fig. la—c)).
Australiceras gigas (Sow.) var. inscriptum Casey (1961: 52, text-fig. 17).
Australiceras ? hirtzi Collignon (1962: 19, pl. 223 (fig. 966), pl. 244 (fig. 967),
pl. 225 (fig. 968)). (According to F6rster 1975a: 57 these are actually
specimens of Ammonitoceras pavlowi Wassiliewski, 1908.)

Australiceras irregulare (Tenison Woods) (1883: 151, pl. 8 (fig. 2)).

asa

290 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 49. Australiceras robustum Whitehouse, 1926. BMNH C25357, paratype from the Upper
Aptian of Flinders River, North Queensland, Australia. x 1. (See Whitehouse 1926: 211.)
According to Day (1974: 9-10) A. robustum may be included in the synonomy of A. jacki.

13. Australiceras jacki (Etheridge Jun.) (1880: 305, pl. 17 (figs 55—58)).

14. Australiceras lamprum (Etheridge Jun.) (1909: 157, pl. 48 (figs 1—2)).

15. Australiceras laticeps (Sinzow) (1905: 314, pl. 19 (figs 2-4)).

16. Australiceras pingue Casey (1961: 55, pl. 14 (fig. 1), pl. 15 (fig. 1), text-
fig. 18b).

17. Australiceras rabenjanaharyi Collignon (1962: 26, pl. 226 (fig. 969)).

18. Australiceras? ramboulai Collignon (1962: 30, pl. 228 (fig. 970)).
(According to Forster 1975a: 158-9 this is an Ammonitoceras.)

CRETACEOUS FAUNAS FROM SOUTH AFRICA 291

19. Australiceras ramososeptatum (Anthula) (1899: 127, pl. 14 (fig. 4)).

20. ? Crioceras sarasini Favre (1908: 638, pl. 36 (figs 1-3), pl. 37 (fig. 2)).

21. Australiceras tuberculatum (Sinzow) (1905: 309, pl. 19 (figs 5-6), pl. 20
(figs 4—5)).

22. Australiceras tuberculatum var. graciloides (Sinzow) (1905: 312, pl. 20
(figs 1-3)).

23. ? Ancyloceras urbani Neumayr & Uhlig (1881: 190, pl. 49 (fig. 3), pl. 50
(fig. 1)). .

Occurrence

Australiceras occurs in the Aptian of England, France, northern Germany,
Bulgaria?, the caucasian and transcaspian regions of the U.S.S.R., India,
Columbia, California, Patagonia?, Australia, Madagascar, Mozambique and
Zululand. Neocomian reports from Japan (Matsumoto 1947) have been dis-
proved by Day (1969: 158). However, since then Obata et al. (1975) recorded
Australiceras aff. gigas from the Chosi Group of Japan. Dimitrova’s (1967: 62,
pl. 30 (fig. 1)) Australiceras bulgaricum cannot be referred to the genus with
certainty.

Australiceras ramososeptatum (Anthula, 1899)

Figs 50-52

Crioceras ramososeptatum Anthula, 1899: 127, pl. 14 (fig. 4). Non Sinzow 1905: 249, pl. 1
(figs 1-3).
Ancyloceras sp. Krenkel, 1910: 154.
Crioceras aff. ramososeptatum: Kasansky, 1914: 40, pl. 1 (fig. 10).
Ancyloceras ramososeptatum: Rouchadzé, 1933: 220, pl. 9 (fig. 1).
? Ammonitoceras ramososeptatum: Drushchitz & Kudryavtsev, 1960: 249, pl. 39 (fig. 1).
Toxoceratoides royerianus: Wachendorf, 1967: 290, pl. 35 (Ags 1, 4, non fig. 5).
Australiceras ramososeptatum: Forster, 1975a: 153, pl. 3 (figs 1-2), text-fig. 30.
Holotype
The specimen figured by Anthula (1899, pl. 14 (fig. 4a)), here refigured as
Figure 51. Original at Palaeontological Institute Uppsala University (Sjorgen

Collection).

Material

SAS L7 (3) from the upper part of the section at locality 170, Mlambong-
wenya Creek, Zululand. Aptian I.

Description

The available specimen consists of one and a third whorls, but is sufficiently
well preserved to illustrate the australiceratid characteristics. Coiling is criocera-
titid with the whorls touching, but not impressed. The whorl section is sub-
quadrate to rounded in the early stages of growth, but soon becomes triangular
with a flattened dorsum, a steep umbilical wall and gently inflated flanks,
converging to a rounded venter.

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292 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 50. Australiceras ramososeptatum (Anthula, 1899). SAS L7(3) from the upper part of the
section at locality 170, Zululand, Aptian I. x 1,1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 293

Fig. 51. Australiceras ramososeptatum (Anthula, 1899). The original of Anthula’s (1899, pl. 14
(fig. 4a)) Crioceras ramososeptatum. Housed in the Palaeontological Institute, Uppsala Univer-
sity (Sjorgen Coll.). Photograph supplied by Dr P. Bengtson (Uppsala). Slightly reduced.

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ANNALS OF THE SOUTH AFRICAN MUSEUM

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CRETACEOUS FAUNAS FROM SOUTH AFRICA 295

At a whorl height of approximately 20 mm, ornament consists of alternate
trituberculate ribs and non-tuberculate intermediaries. The tuberculate ribs are
much stronger than the intermediaries, being about twice as wide. The ventral
tubercles are largest, and situated a little distance from the siphonal line. The
lateral tubercles are on the ventral third of the flanks. The umbilical tubercles
are smallest and either pointed or elongated in the direction of ribbing.

After about a third of a whorl tuberculation disappears rather suddenly,
and ornament consists of slightly rursiradiate ribs only, separated by interspaces
of similar width. Occasional bifurcations occur at midflank.

The outer whorl is abraded and encrusted by epizoans, but appears to have
borne similar ribs. The suture line is very incised, with a large asymmetrical
trifid lateral lobe which covers virtually the whole of the flanks. The saddles are
very narrow-stemmed.

Dimensions
Specimen D Wb Wh Wb/Wh U
SAS L7 (3) 87,5 36(41,1) 36(41,1) 1 39(44,5)
155 73(47,1) 66(42,5) let c. 65(41,9)
Discussion

The strong tuberculation on the inner whorl, combined with the steep
umbilical wall on the outer whorl are characteristic of the species. Anthula
(1899: 127) mentioned the presence of intermediary, non-tuberculate ribs which
may appear regularly or sometimes be absent. This, however, does not feature
prominently in Anthula’s illustration or in the holotype here refigured as
Figure 51.

Some subsequent references of material to Anthula’s species appear to be
incorrect. Sinzow’s (1915, pl. 11 (figs 1-3a)) Crioceras ramososeptatum (especially
fig. 1) has distinctly looped and ventrally bifurcating ribs, and should probably
be referred to Ammonitoceras. The specimen figured by Drushchitz &
Kudryavtsev (1960: 294, pl. 39 (fig. 1)) as Ammonitoceras ramososeptatum has
finer ribbing than Anthula’s species, and indeed appears to be Ammonitoceras.

Crioceras jackii Etheridge Jun. (see Figs 47-49), especially Etheridge’s
specimen (1909, pl. 35 (fig. 1)), is close to the present species, but differs mainly in
having finer ribbing, more intermediaries during the tuberculate stage, and more
bifurcating ribs in the non-tuberculate stage.

Crioceras australe Waagen non Moore (1875: 246, pl. 60 (fig. la—c)) is
slightly similar, but has a rounded whorl section and a dorsal zone of impression.

Australiceras argus Anderson (1938: 211, pl. 70 (figs 1-2)) is similar in having
alternate tuberculate and non-tuberculate ribs on the inner whorls, but has
coarser ribbing and an ovoid whorl section on the outer whorl.

Australiceras lampros (Etheridge Jun.) has a similar triangular whorl
section (see Whitehouse 1926, pl. 35 (fig. 1b)), but is only weakly trituberculate
in early stages of growth.

296 ANNALS OF THE SOUTH AFRICAN MUSEUM

Occurrence
Lower Aptian of southern Russia, Mozambique and Zululand.

Australiceras sp. aff. A. irregulare (Tenison Woods, 1883)
Figs 65C—D, 68F, 80A

Compare:

Crioceras irregulare Tenison Woods, 1883: 151, pl. 8 (fig. 2). Etheridge Jun.
1892: 501, pl. 33 (fig. 1), pl. 42 (fig. 16).

Crioceras jackii Etheridge Jun., 1909: 145 (pars), pl. 35 (fig. 1), pl. 36 (fig. 1),
pl 37 tig. 2):

Australiceras irregulare: Whitehouse, 1926: 210, pl. 37 (fig. la—b). Day, 1974: 10,
pl. 2 (fig. 3), pl. 6 (figs 1-2a-c), text-figs 3A, C. (With complete synonomy.)

Material

SAS 63/2, SAS LJE183; SAS Z8/2 from locality 168 and BMNH C79714
from locality 167, Mfongozi Creek, Aptian III-IV.

Description

A small Australiceras, with slightly elliptical, crioceratitid coiling. The
whorls touch, but are not impressed. The whorl section is initially rounded, but
progressively becomes higher than wide. The initial whorls, at a whorl height of
approximately 5,5 mm, are ornamented by narrow widely spaced prorsiradiate
ribs, apparently lacking tubercles. At a whorl height of approximately 7 mm,
every second rib is strengthened and tubercles appear, and by a whorl height of
10 mm tuberculation is already quite prominent. The ventral tubercles are
largest, and the umbilical ones smallest. The distance between the umbilical and
lateral tubercles is greater than that between the latter and the ventral tubercles.
The shape of tubercles is variable, the ventral ones are usually round and
bullate, whereas the lateral and umbilical ones are elongated in the direction of
ribbing in the early stages, but eventually also become slightly rounded. Non-
tuberculate intermediaries vary from one in the initial stages to up to three,
although there are normally only two in later growth stages. Ribbing is
continuous, though weakened over the dorsum.

Dimensions

Specimen D Wb Wh Wb/Wh U
SAS 63/2 64,6 6. 20(6.-31) 20,5(32) c. 0,86 32(50)

Discussion

The scope of Australiceras irregulare (Tenison Woods) has been extended
so far by Day (1974: 10) that the Zululand material may be tentatively referred
to that species. A. irregulare has initial whorls with either weak or no tuber-
culation, and, as Day (1974, pl. 2 (fig. 3)) has shown, they are not always coiled

CRETACEOUS FAUNAS FROM SOUTH AFRICA 297

in one plane. On the basis of the material available, a definite identification is not
possible. The specimens are of some importance though, for they may point to
the origin of Helicancyloceras gen. nov., to be described below.

Crioceras (Ancyloceras) matheroni in Pavlow (1890: 4, pl. 6 (fig. 2a—b)) bears
superficial resemblance to the present species, but is too poorly known for
further comment.

Occurrence
Upper Aptian of Zululand.

Australiceras wandalina (Boshoff MS.) sp. nov.
Figs 6B, 53-57, 61D
Holotype
UPG-B8 (Boshoff Collection), Department of Geology, University of
Pretoria, from the Upper Aptian of Manyola Drift, northern Zululand,
Aptian III. Collected by J. C. Boshoff. (Unpublished thesis 1945.)

Material

Paratype SAS EM106 is from Manyola Drift, Aptian III, Paratype
BMNH C78888 is from the Upper Aptian of Mlambongwenya Spruit.

Description

The specimens are still septate at the largest diameters preserved, but
judging by the rather tight coiling, it appears unlikely that the body chamber
becomes detached. The initial whorl section is suboctagonal, slightly higher than
wide, but eventually becomes subrectangular with a rounded venter and a slight
dorsal zone of impression (Fig. 6B).

At the smallest preserved diameter (Wh = 25 mm), ornament consists of
Straight, radial ribs arising at the umbilical edge, each bearing three rows of
pointed tubercles. The umbilical and lateral ones are situated on the dorsal and
ventral thirds of the flanks respectively; the ventral ones are a little distance
away from the siphonal line. All the tubercles are pointed and elongated in the
direction of ribbing to a certain extent. With increasing diameter ribbing
becomes blunter and the tubercles more bullate. The umbilical tubercles
disappear at a diameter of c. 225 mm, whereas in some specimens the lateral
ones, which have now migrated closer to the venter, become indistinct at a
diameter of 240 mm. The ventral tubercles persist as swellings on the ventral
part of the ribs to much greater a diameter.

On the inner whorl, the rib density is forty-nine per whorl and intermediaries
seem to be absent. On the outer whorl, rib density is fifty-one per whorl and two
intermediaries occur.

The suture line, despite decortication, appears very simple, with large and
asymmetrically trifid lobes. The lateral lobe is largest and occupies virtually the
whole of the flanks.

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ANNALS OF THE SOUTH AFRICAN MUSEUM

298

Fig. 53. Australiceras wandalina (Boshoff MS) sp. nov. UPG B8, holotype from Manyola

Drift, Zululand, Aptian III. x 0,44.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 299

Fig, 54, Australiceras wandalina (Boshoff MS) sp. nov. UPG B8, holotype from Manyola
Drift, Zululand, Aptian III, =< 0,44,

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300 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 55. Australiceras wandalina (Boshoff MS) sp. nov. SAS EM106 from Manyola Drift,
Zululand, Aptian III. x 0,59.

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CRETACEOUS FAUNAS FROM SOUTH AFRICA

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302 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 57. Australiceras wandalina (Boshoff MS) sp. nov. BMNH C78888 from locality 171,
Zululand, Found loose. Specimen with much finer ribbing on inner whorls, probably repre-
senting a new subspecies. < 0,67, |

CRETACEOUS FAUNAS FROM SOUTH AFRICA 303

Dimensions
Specimen D Wb Wh Wb/Wh U
UPG-B8 a73 134(36) 144(38,6) 0,93 160(43)
Discussion

The strong ornamentation on the inner whorls, combined with the
persistence of ventral tubercles to such a large diameter, characterize the species.

A single specimen (BMNH C78888 from locality 171 found loose on the
surface (Fig. 57)) and consisting of less than half of two successive whorls has
fine ribbing on the inner whorls, c. eighty per whorl, but ornament comparable
to that of A. wandalina on the outer whorl, and may possibly represent a new
subspecies.

Amongst Madagascan species, Australiceras hirtzi Collignon (1962: 19,
pl. 223 (fig. 966), pl. 224 (fig. 967), pl. 225 (fig. 968)), has comparable inner
whorls and relative proportions. It differs from A. wandalina, however, in
losing its ventral tubercles at a relatively early stage, and in having finer ribbing
on the outer whorl. (See Collignon 1962, pl. 225 (fig. 968).)

Australiceras rabenjanaharyi Collignon (1962: 26, pl. 226 (fig. 969)) has
finer ornament both on the inner and outer whorls. Australiceras? ramboulai
Collignon (1962: 30, pl. 228 (fig. 970)) has whorls increasing very slowly in size,
and a curious ornament of ribs bifurcating from the umbilical tubercles in the
later stages. As Forster (1975a: 158) has shown, this species is probably
synonymous with Ammonitoceras pavlowi (Wassiliewski 1908: 46, pl. 3
(fig. la—c)).

Retention of tubercles to such a large diameter is reminiscent of ‘Crioceras’
deeckei Favre (1908: 636, pl. 36 (fig. 4), pl. 37 (fig. 1), text-fig. 1) and ‘Crioceras’
sarasini Favre (1908: 638, pl. 36 (figs 1-3), pl. 37 (fig. 2)) and the Zululand
specimen to be described below as Australiceras? sp. cf. ‘Crioceras’ sarasini
Favre. The Patagonian species, whatever their generic identity, however, have
much stronger tuberculation than the present species.

Occurrence
Upper Aptian of Zululand.

Australiceras ? sp. cf. ‘Crioceras’ sarasini Favre, 1908

Fig. 10A
Compare:

Crioceras sarasini Favre, 1908: 638, pl. 36 (figs 1-3), pl. 37 (fig. 2).
Material
SAS H71D/18 from Nhlohlela Pan, Mkuze Game Reserve, locality 150,
Aptian ITI-IV.
Description |

A small septate fragment, 130 mm long, is comparable with the larger end
of the outer whorl of the original of Favre’s pl. 36 (figs 1-2). The intercostal

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304 ANNALS OF THE SOUTH AFRICAN MUSEUM

section is rounded subtriangular with a flat dorsum and moderately inflated
flanks converging to a narrow venter. Maximum width is at the dorsal third of
the flanks. Ornament consists of five strong, rounded, radial ribs, bearing three
rows of tubercles on either side. The dorsum is smooth and devoid of ribbing.
There is a distinct depression along the venter between the tubercles where
ribbing is interrupted.

Discussion

As mentioned above, there is striking similarity between the Zululand
specimen and the larger end of Favre’s “Crioceras’ sarasini. The only apparent
difference is that the umbilical tubercles in the Patagonian species are larger,
and that the ribs are wider spaced. However, on the basis of the limited material,
a definite specific allocation is impossible. Both ‘Crioceras’ sarasini Favre and
‘Crioceras’ deeckei Favre are of enigmatic generic affinity. Spath erected the
genus Peltocrioceras for ‘Crioceras’ deeckei which ‘requires a new generic
designation’ (Spath 1924: 85). Neither then, nor later, was any generic diagnosis
given by Spath, and whether “Crioceras’ sarasini was to be included in Pelto-
crioceras or not was never specified. Riccardi (1971: 277) regards both species
as being representatives of Peltocrioceras, but as being close to both Tropaeum
and Australiceras. Leanza (1963: 221) and Day (1974: 6) (the latter in comparing
‘Crioceras’ deeckei with Tropaeum imperator Howchin & Whitehouse) both
indirectly suggested that Peltocrioceras might be a junior synonym of Tropaeum.
Wiedmann (1962: 112) regarded Peltocrioceras as a synonym of Crioceratites.
‘Crioceras’ sarasini differs from ‘Crioceras’ deeckei mainly in having weaker
lateral tuberculation on the inner whorls, and in having interrupted ribbing
over the venter, ending in strong nodes. Both species belong to the same genus,
which appears to bear superficial similarity to both Tropaeum and Australiceras.

Australiceras wandalina sp. nov. is superficially similar in retaining tuber-
culation to large diameters, but has denser ribbing and the tubercles are not as
strong.

Australiceras ramboulai Collignon (1962: 30, pl. 227 (fig. 970)) has a
similar whorl section, but lacks the strong ornament.

Occurrence

‘Crioceras’ sarasini is of Aptian age, and not Barremian as Favre (1908) and
Piatnitzky (1938) had incorrectly assumed. The Zululand specimen is of Upper
Aptian age, Aptian III-IV.

Australiceras ? sp. indet. A.
Fig. 38A—B
Description and discussion

A septate fragment, SAS L3 from locality 170, and of Lower Aptian age,
has an octagonal whorl section and four trituberculate ribs, and is referred to

CRETACEOUS FAUNAS FROM SOUTH AFRICA 305

Australiceras with doubt. Ribbing is radial on the flanks, interrupted over the
venter and absent on the dorsum. The fragment is possibly allied to
A. wandalina sp. nov. and A. sp. cf. ‘Crioceras’ sarasini Favre in being strongly
tuberculate at large diameters.

Ammonitoceras pavlowi (Wassiliewski, 1908), described and figured from
Mozambique by Forster (1975a: 156, pl. 3 (figs 5-7)), has a similar whorl
section, but differs in having regular ventrally bifurcating or intercalatory ribs.

Australiceras ? sp. indet. B.

Fig. 58A—B
Description and discussion

A curved body chamber fragment, SAS 54/45 from the Lower Aptian at
locality 170 below the horizon of abundant Ancyloceras (Adouliceras) with
flared, trituberculate ribs, may possibly belong to the uncoiled forms of genus
Australiceras. The whorl section is ovoid, depressed, with a little-rounded
dorsum and strongly rounded flanks and venter. Five flared ribs with distinct
ventral but indistinct lateral and umbilical tubercles are present. The dorsum is
ornamented by faint striae.

The strongly depressed whorl section recalls Sinzow’s Crioceras laticeps
(1905: 314, text-fig. 3, pl. 19 (figs 2-4)). In Sinzow’s species, however, rib density
is less than in the present specimen. Specimens of Australiceras gigas (J. de C.
Sowerby) sometimes also show such flared ribs in the crozier, but again, the rib
density is lower.

Subfamily Helicancylinae Hyatt, 1894

Hyatt’s family Helicancylidae was revived by Casey (1961: 76, as Helican-
cylinae) ‘as a useful subfamily term for those diminutive ancyloceratids in
which the sculpture is simplified on the terminal hook’.

The subfamily, as interpreted by Casey, is here accepted with reservation as
a useful grouping of three genera only: Tonohamites Spath, 1924, Toxoceratoides
Spath, 1924, and Helicancylus Gabb, 1869, although Acrioceras Hyatt, 1900,
Lytocrioceras Spath, 1924, and Leptoceras Uhlig 1883, have also tentatively been
referred to the sub-family.

Apart from the fact that the type genus is difficult to interpret (see Casey
1961: 77), transitional forms occur, linking the genera. Classification is based
mainly on the mode of ribbing on the body chamber and, to a lesser extent, the
presence or absence of (tri-)tuberculation on the phragmocone.

Genus Toxoceratoides Spath, 1924
Type species
Toxoceras royerianum d’Orbigny, 1842 from the Lower Aptian of France.

ANNALS OF THE SOUTH AFRICAN MUSEUM

306

54/45 from locality 170, Zululand, Aptian J. x 1.

SAS

? sp. ind. B.

Fig. 58. Australiceras

CRETACEOUS FAUNAS FROM SOUTH AFRICA 307

Diagnosis

Small forms with ancyloceratid or leptoceratid coiling, probably with an
initial helix. Trituberculate ribs occur on the phragmocone but disappear on the
recurved hook.

Discussion

The genus was discussed extensively by Casey (1961: 79) and little can be
added. Differences between Toxoceratoides and Tonohamites are slight, and
trituberculate forms such as Tonohamites decurrens connect the two genera.

The following species were referred to genus Toxoceratoides by Casey
(1961):

1. Toxoceratoides biplex (von Koenen) (1902: 381, pl. 41 (figs 3, 10a—b, 1 la—b).
2. Toxoceratoides caucasicus (Kasansky) (1914, pl. 1 (fig. 8a—c)).
3. Toxoceratoides fustiformis (von Koenen) (1902: 384, pl. 41 (figs 4-5, 7a-c,
9a—b), pl. 53 (figs 8a—b, 9a—b)).
4. Toxoceratoides obliquatus (Young & Bird) (1828: 278, pl. 18 (fig. 11).
Toxoceratoides proteus (Spath) (1930: 461, pl. 16 (fig. 7)).
6. Toxoceratoides rochi Casey, 1961 (=Ancyloceras royerianum in Roch 1927:
30, pl. 1 (fig. 4)).
7. Toxoceratoides rotundus (Phillips) (1875: 264, pl. 1 (fig. 24)).
8. Toxoceratoides royerianum (d’Orbigny) (1842: 481, pl. 118 (figs 7-11)).
9. Toxoceratoides seminodosus (Roemer) 1841.
10. Toxoceratoides sheperdi (Spath) (1942: 173, figs 5-6).
To this list may be added:
11. Toxoceratoides biplicatum (von Koenen) (1902: 379, pl. 41 (figs 2a—b, 8a—b)).
12. Toxoceratoides krenkeli Forster (1975a: 160, pl. 4 (figs 1-2), text-fig. 33a—b).
13. Toxoceratoides saulae Murphy (1975: 31, pl. 4 (figs 4, 6)).
14. Toxoceratoides starrkingi (Anderson) (See Murphy 1975: 32, pl. 4 (figs 3, 5)).
15. Toxoceratoides corae Murphy (1975: 33, pl. 5 (figs 1, 5)).
16. ? Toxoceratoides greeni Murphy (1975: 33, pl. 5 (figs 2-3, 6)).
17. Toxoceratoides spp. indet. described by Murphy as Toxoceratoides sp. |
(1975: 35, pl. 6 (figs 1-2, 11) and ?Toxoceratoides sp. (1975: 35, pl. 3 (fig. 6),
pl. 6 (figs 5-6)).
18. Toxoceratoides sp. nov. Thomson (1974: 16, pl. 3a, d).

Sol

Occurrence

Toxoceratoides ranges from the Upper Barremian to Upper Aptian, and is
known from Europe, California and south-east Africa. Reports from Australia
are not accepted here. ‘Ancyloceras’ taylori Etheridge, referred to Toxoceratoides
by Whitehouse (1926: 216) is a doubtful contender, as are the five specimens
described by Day (1974: 13) as Toxoceratoides ? spp.

308 ANNALS OF THE SOUTH AFRICAN MUSEUM

Toxoceratoides royerianus (d’Orbigny, 1842)

Fig. 59E

Toxoceras royerianus d@’Orbigny, 1842: 481, pl. 118 (figs 7-11).

? Toxoceras emericianus d’Orbigny, 1842: 487, pl. 120 (figs 5-9).

Toxoceratoides royerianus: Casey, 1961: 78, pl. 6 (fig. 2a—b), pl. 17 (fig. 3a—b), text-fig. 30a—h)
(cum synon.).

Neotype

The specimen figured in Casey (1961, text-fig. 30a—c) from the Lower Aptian
of Haute Marne, France, and in the collections of the Sorbonne, Paris.

Material
SAM-PCM5313 from Lubemba, southern Mozambique.

Description

A recurved crozier with part of the phragmocone is referred to the species.

The part of the phragmocone preserved on the recurved crozier is heavily
abraded and only umbilical tubercles are preserved. At the apertural end of the
crozier the whorl section is ovoid, with a flattened dorsum, sharp umbilical edge
and broadly rounded venter. Ornament consists of relatively narrow, high ribs,
separated by deep interspaces which are wider than the ribs. The ribs arise in
groups of two or three from the umbilical tubercles. They are prorsiradiate at
the proximal end of the crozier, but become radial on the bend of the crozier
and at the distal end. At the apertural end a pair of single ribs arises from the
umbilical tubercles.

Discussion

The recurved crozier compares well with the illustrations and figures
provided by Casey (1961), who also discussed the species extensively.

Toxoceratoides krenkeli Forster, 1975a differs from T. royerianus mainly in
having hardly any intermediary ribs on the shaft, and in having coarser ribbing
on the crozier.

Occurrence
Lower Aptian of western and central Europe, Mozambique and Zululand.

Toxoceratoides krenkeli Forster, 1975

Fig. 59F

Hamites royerianus: Kilian, 1902: 465.

Ancyloceras royerianum: Krenkel, 1910: 151, pl. 17 (figs 12-13).

Tonohamites royerianum: Haughton & Boshoff, 1956: 13, pl. 2 (fig. 3).
Acrioceras dissimilis: Wachendorf, 1967: 281, pl. 35 (figs 2-3), text-fig. 5.
Toxoceratoides royerianus: Wachendorf, 1967: 290, pl. 35 (fig. 5, non figs 1, 4).
Toxoceratoides krenkeli Forster, 1975a: 160, pl. 4 (figs 1-2) text-fig. 33a—b.

Holotype
Wachendorf’s (1967, pl. 35 (fig. 2)) specimen from Chalala, Mozambique.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 309

Fig. 59. A-D. Toxoceratoides ? haughtoni sp. nov. BMNH C79701 from locality 166, Zululand,

Aptian III. x 2. E. Toxoceratoides royerianus (d’Orbigny, 1842). SAM-PCM5313 from

Lubemba, Mozambique, Lower Aptian. x 1,7. F. Toxoceratoides krenkeli Forster, 1975a.
SAM-PCM5342 from Lubemba, Mozambique, Lower Aptian. x 1,5.

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310 ANNALS OF THE SOUTH AFRICAN MUSEUM

Material
SAM-PCM 5342 from Lubemba, southern Mozambique, Lower Aptian.

Description and discussion

A small, slightly curved part of the phragmocone is referred to Forster’s
species. |

The whorl section is nearly circular on the earliest part preserved. Up to a
diameter of 5,5 mm ornament consists of strongly prorsiradiate ribs, separated
by interspaces of similar width. Rib density is about four per whorl height. The
ribs weaken markedly on, and pass straight across the dorsum. Tuber-
culation commences at a diameter of 5,5 mm; at first only ventral tubercles are
present, but four ribs further on, distinct, though small lateral and umbilical
tubercles are present, and there are faint indications of duplication of ribs over
the dorsum.

Forster (1975a: 160 et seq.) discussed the affinities of the species in detail;
its main characteristic is the sparsity or absence of non-tuberculate intercalatories
on the shaft. The present specimen adds to our knowledge of the species in that
it illustrates that the early parts of the curved shaft lack tubercles; in this
respect it resembles the later Toxoceratoides ? haughtoni sp. nov. of Late Aptian
age, described below.

Occurrence

Lower Aptian of southern Mozambique.

Toxoceratoides? haughtoni sp. nov.

Figs 59A-D, 60A-I, 61A—C, 62A-D, 63, 64A-C, 65A—B, 66B, 79A—B

Holotype

SAS 64/T1 from locality 168, Mfongozi Creek, northern Zululand.
Aptian IIJ-IV. South African Geological Survey Collection, Pretoria. Collected
by H. Klinger 1970.

Derivation of name

The species is named in honour of Dr S. H. Haughton, discoverer of many
Cretaceous localities in northern Zululand.

Material

Apart from the holotype, SAS L65/T2, SAS Z8/T1, SAS Z8/T4, SAS Z8/T5,
SAS LJE 186, all from locality 168, Mfongozi Creek, Zululand, and BMNH
C79690, C79191, C79694-79701 from locality 166, Aptian III, Aptian I-IV,
and SAS N2 from locality 50.

CRETACEOUS FAUNAS FROM SOUTH AFRICA Sel

t a oes Pigg: fs
eign Sp
wae ee nae

ee

Fig. 60. Toxoceratoides ? haughtoni sp. nov. A-C. BMNH C79695 from locality 166, Zululand,
Aptian III. D-F. BMNH C79696, locality and age as above. G-I. BMNH C79712, locality
and age as above. All x 2. |

312 ANNALS OF THE SOUTH AFRICAN MUSEUM ,

Fig. 61. A-C. Toxoceratoides ? haughtoni sp. nov. BMNH C79694 from locality 166, Zululand,
Aptian III. x 2. D. Australiceras wandalina (Boshoff MS) sp. nov. BMNH C7888 from locality
171, Zululand. Aptian. x 0,67.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 313

B

Fig. 62 A-D. Toxoceratoides ? haughtoni sp. nov. BMNH C79694 from locality 166, Zululand,
Aptian III. x 2.

ANNALS OF THE SOUTH AFRICAN MUSEUM

314

we naw ee”

i sp. nov.

truction of Toxoceratoides ? haughton

1c recons

t

lagramma

D

63:

1g

P

CRETACEOUS FAUNAS FROM SOUTH AFRICA 315

Fig. 64. Toxoceratoides ? haughtoni sp. nov. A. SAS Nd from locality 50, Upper Aptian. x 1,6.
B-C. SAS 64/T1, holotype from locality 168, Zululand, Aptian III-IV. x 1,4.

316 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 65. A~B. Toxoceratoides ? haughtoni sp. nov. A. SAS LJE 186 from locality 168, Zululand,
Aptian III-IV. x 1,5. B. SAS Z8/T5 from locality 168, Zululand, Aptian III-IV. x 1,5.
C-D. Australiceras sp. aff. irregulare. C. SAS Z8/TS5a from locality 168, Aptian III-IV. x 1,5.
D. SAS L65/2 from locality 168, Zululand, Aptian IIJ-IV. x 1. E. Tropaeum obesum sp. nov.
SAM-PCZ5681. Fragment of inner whorl! of holotype. x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 317

Fig. 66. A. Suture line of Tonohamites koeneni Casey, 1961. SAS Z1115 at diameter c. 8 mm.

x 6. B. Suture line of Toxoceratoides ? haughtoni sp. nov. at diameter c. 8 mm. x 6. C-D.

Suture lines of Tonohamites koeneni Casey, 1961. C. SAS Z1115 at diameter c. 15 mm. x 3.
D. SAS H71D/19 at diameter c. 16 mm. x 3.

318 ANNALS OF THE SOUTH AFRICAN MUSEUM

Description |

The coiling is toxoceratoid, open with a recurved crozier. The very early
stage of growth is unknown. The whorl section is initially rounded, but becomes
laterally compressed as size increases. Ornamentation is best shown by the
holotype. At the smallest diameter, 6,1 mm, ornament consists of single, simple
ribs, slightly prorsiradiate and separated by interspaces of similar width. At this
stage there are approximately six ribs in a distance equal to the whorl diameter. ~
At a slightly larger diameter the ribs become increasingly differentiated over the
venter. Initially this appears as a mere flattening along the siphonal line, but at
larger diameters the ribs are distinctly interrupted over the venter, terminating
in small tubercles on either side of the siphonal line. Initially, all the tubercles
are of equal size, but at a diameter of 8,5 mm some appear stronger than the
others. With increasing diameter, these tubercles appear with greater frequency
and become greatly enlarged, whilst the ribs on which they are situated become
bolder on the flanks, and small ventrolateral and even smaller umbilicai
tubercles appear on the ventral and dorsal thirds of the flanks respectively. One
(or rarely two) intermediate non-tuberculate ribs are present. At large diameters,
the ribs connecting the ventral and ventro-lateral tubercles may become
flattened, or even show signs of looping, in a manner reminiscent of some
Protanisoceras species.

Ribbing weakens on the dorsum, and the ribs pass straight across or have a
slight forward curvature. Some of the major ribs show distinct looping or
duplication on the dorsum.

Ornament remains more or less constant up to the sharply recurved
crozier. In the bend of the crozier more intercalated, thin ribs occur, and on the
recurving limb all ornament, with the possible exception of the ventral tubercle,
weakens.

The suture is relatively symmetrical with a trifid lateral, umbilical and
internal lobe.

Discussion

All the available specimens show toxoceratoid coiling. If this species is a
true Toxoceratoides, it extends the range of the genus to the Upper Aptian. The
peculiar ornament in the early whorls and the lack of regular simple ribbing in
the crozier is different from that of the normal Toxoceratoides ornament, hence
the species is referred to this genus with doubt. This also serves to distinguish it
from all the other known species of Toxoceratoides.

The non-tuberculate ribbed stage is comparable to that of T. krenkeli. That
species, however, is tuberculate at much smaller diameters and has a depressed
whorl section, different from the compressed whorl of 7. ? haughtoni.

Ancyloceras patagonicum Stolley (1912: 11, pl. 1 (figs 2a—b, 3a—b)) may also
be compared with the present species. The Patagonian species has, however,
stronger ribbing in the early stages of growth, whilst the major costae are not as
strongly developed as in the present species.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 319

One of Von Koenen’s specimens of Ancyloceras fustiforme (1902, pl. 53
(fig. 9)) bears some resemblance to the early parts of 7. ? haughtoni in having
ventrally differentiated ribbing which becomes tuberculate with increasing
diameter. In later stages, however, this species is not as strongly tuberculate as
the Zululand material, as is illustrated by the original of Von Koenen’s (pl. 41,
fig. 9) specimen.

Occurrence
Fairly common in the Upper Aptian of Zululand.

Genus Tonohamites Spath, 1924
Type species
Tonohamites decurrens Spath, 1924 from the Lower Aptian of Ahaus,
Germany, by original designation.

Diagnosis
Coiling labeceratid. Tuberculation weaker than in Toxoceratoides and

mostly confined to the venter. Ribbing in early stages hamitid, but towards body
chamber broad and flat in some.

Discussion

Like Toxoceratoides, Tonohamites was doubtfully regarded as a synonym of
Hamiticeras by Wright (1957: L212). Casey (1961: 84), however, maintains that
it is quite a valid and useful genus for small labeceratid-like forms with reduced
tuberculation on the phragmocone and rounded ribbing on the venter.
Tonohamites decurrens Spath with strong trituberculation on the phragmocone
connects Tonohamites with Toxoceratoides, thus illustrating the close relationship
between the two genera.

The genus was discussed at length by Casey (1961: 84) and, apart from
extending the stratigraphic and geographical range of the genus, no further
comments are necessary.

Species referred to Tonohamites by Casey are as follows:

1. Tonohamites decurrens Spath, 1924: 461. (Lectotype is Von Koenen’s 1902,
pl. 33 (fig. 2) and lower part of pl. 33 (fig. 3)).

2. Tonohamites aequicingulatus (von Koenen) (1902: 394, pl. 37 (figs 5a-c,
6a-e)).

3. Tonohamites ? eichwaldi (Jasykow) in Sinzow (1872: 36, pl. 6 (figs 7—9)).

4. Tonohamites ? hunstantonensis Casey (1961: 90, pl. 21 (fig. la—d)).

5. Tonohamites koeneni Casey: (1961: 89). Holotype is Von Koenen’s (1902,
pl. 33 (fig. 3a)) specimen, upper part only.

6. Tonohamites limbatus Casey (1961: 89, pl. 22 (figs 3a—c, 4), pl. 21 (fig. 3a—b)).

7. Tonohamites ? undosus (von Koenen) (1902: 393, pl. 35 (fig. 13a-f)).

Occurrence

Tonohamites occurs in the Lower Aptian of Europe and the Upper Aptian
of Madagascar and Zululand.

320 ANNALS OF THE SOUTH AFRICAN MUSEUM

Tonohamites koeneni Casey, 1961
Figs 66A, C—D, 67C—D, 73A, 81F

Hamites decurrens (non Roemer) von Koenen, 1902: 392, pl. 33 (fig. 3a upper part only).
Tonohamites koeneni: Casey, 1961: 89, text-fig. 35g—h.

Holotype

The top part only of the specimen figured by Von Koenen (1902, pl. 33 ©
(fig. 3)) as Hamites decurrens by original designation (Casey 1961: 89).

Material

SAS Z1115, SAS Z1118, SAS H71D/19 all from locality 152, Mkuze Game
Reserve. Aptian IV.

Description

The most complete specimen, SAS Z1115 (Fig. 67C—D), shows the coiling
to be labeceratid, with a moderately long shaft. Coiling of the very early whorls
is unknown.

The whorl section is rounded in the early whorls, but becomes progressively
flattened dorsoventrally with a nearly flat dorsum. Maximum width is at midflank
or just dorsal to it. Ornament on the curved shaft consists of simple, prorsiradiate
ribs, separated by narrower interspaces. The ribs are narrow on the dorsum and
nearly radial, then are prorsiradiate and cross the flank with increasing strength.
Towards the venter the ribs become more rounded and wider, a tendency which
is especially pronounced on the body chamber. There are four to four and a half
ribs in a distance equal to the whorl diameter on the shaft. At the bend on the
crozier, the ribs become extremely flattened on the venter and the interspaces on
the flanks become wider. The aperture appears to be quite simple, with neither
lappets nor a constriction.

Discussion

As Casey (1961: 84) has outlined, Von Koenen’s illustration of Hamites
decurrens (1902, pl. 33 (figs 2, 3a—b)) is a synthetograph. The lower part of the
synthetograph was chosen by Casey as lectotype of Tonohamites decurrens Spath,
whilst the upper part was regarded as a new species, Tonohamites koeneni. The
Zululand material fits the upper part of Von Koenen’s figure so well that the
authors have no hesitation in referring it to 7. koeneni, despite differences in age.
T. koeneni is believed to be of Lower Aptian age (as is the whole genus in the
sense used by Casey (1961: 85)). The Zululand material is of Upper Aptian age,
occurring just below the line of hiatus concretions (Kennedy & Klinger 1972)
marking the Aptian/Albian boundary.

Casey’s Tonohamites cf. koeneni (1961: 89, pl. 20 (fig. 5a—b)) from the
Lower Aptian of Kent does not show as pronounced broad flattened ribs on the
body chamber as do Von Koenen’s or the Zululand specimens.

Tonohamites decurrens Spath has a distinct trituberculate phragmocone,

CRETACEOUS FAUNAS FROM SOUTH AFRICA 321

Fig. 67. A-B. Helicancyloceras (Helicancyloceras) densecostatum sp. nov. SAM-—PCZ5682,
holotype from locality 168, Zululand, Aptian III-IV. x 1,3. C-D. Tonohamites koeneni
Casey, 1961. SAS Z1115 from locality 152, Zululand, Aptian IV. x 1,4.

322 ANNALS OF THE SOUTH AFRICAN MUSEUM

and is easily distinguished. 7. aequicingulatus (von Koenen) has a somewhat
similar body chamber, but has ventral tubercles on the shaft. 7. limbatus Casey
has a long, descending limb and small tubercles on the phragmocone.

Occurrence

Upper Aptian of Zululand. The English specimen tentatively referred to the
species by Casey is from the Lower Aptian bowerbanki Zone. In Germany the -
species occurs in the Lower Aptian of Ahaus.

Tonohamites aequicingulatus (von Koenen, 1902)
Figs 68A-E, 38C, 88D

Hamites aequicingulatus von Koenen, 1902: 394, pl. 37 (figs 5-6).
Tonohamites aequicingulatus Casey, 1961: 87, pl. 9 (figs 2a—b, 3a—b, 4), text-fig. 32.
Tonohamites sp. aff. aequicingulatus Collignon, 1962: 14, pl. 221, (fig. 970).

Lectotype

The original of Von Koenen (1902, pl. 37 (fig. 5a—c)) by subsequent
designation Casey 1961: 87.

Material

SAS Z8Tal from locality 168, BMNH C79690, C79692 from locality 166,
Mfongozi Creek, Zululand. Aptian III-IV.

Description

The available specimens are fragmentary and consist of slightly curved
shafts. The whorl section is depressed, with rounded flanks and a flattened
dorsum and venter.

Ornament consists of prominent, obliquely prorsiradiate ribs, separated by
wider interspaces. The ribs are strongest on the flanks, thickening towards the
ventrolateral region. In the siphonal area the ribs are interrupted or weakened
at the larger diameter. At the smaller ends of the shaft there is no distinct
weakening of ribbing over the venter. The ribs pass radially or with a slight
forward curvature over the dorsum.

Discussion

T. aequicingulatus presents difficulties in interpretation, as it is only known
from fragments, and, as Casey (1961: 88) noted, it ‘seems practicable to treat
these fragments as belonging to a single, variable species’.

The Madagascan specimen tentatively referred to this species by Collignon
has a rib density of four or five per diameter, the general appearance being
similar to that of the Zululand material, and it is probably conspecific. The only
apparent difference between the figured European specimens in Casey (1961)
and Von Koenen (1902) and those from Zululand and Madagascar, is that the
ribbing in the latter specimens is stouter near the venter. The subsymmetrical

CRETACEOUS FAUNAS EROM SOUTH AFRICA 323

Fig. 68. A-E. Tonohamites aequicingulatus (von Koenen, 1902). A-B. BMNH C79692 from

locality 166, Zululand, Aptian III. x 2. C-E. BMNH C79690 locality and age as above. x 2.

F. Australiceras sp. aff. A. irregulare (Tenison Woods). BMNH C79714 from locality 167,

Zululand, Aptian II-IV. x 1. G—H. Helicancyloceras (Nonyaniceras) sp. indet. BMNH 79717
from locality 167, Zululand, Aptian III-IV. x 2.

324 ANNALS OF THE SOUTH AFRICAN MUSEUM

lateral lobe mentioned by Casey (1961: 88) is noticeable in both the Zululand
and the Madagascan material.

Occurrence

Tonohamites aequicingulatus occurs in the Lower Aptian bowerbanki
subzone in England. Von Koenen (1902: 394) referred his specimens to the
Aptian and with a question mark to the Zone of Hoplites deshayesi. The
Madagascan specimen is from the Upper Aptian Zone of Aconeceras nisus and
Melchiorites melchioris.

Tonohamites? caseyi sp. nov.

Fig. 69 (see also Fig. 46)
Holotype

SAS Z804, the specimen marked A from locality 150 or 151, Mkuze Game
Reserve. Aptian IV. Geological Survey Collection. Collected by the late
Dr E. C. N. van Hoepen.

Derivation of name

Named after Dr R. Casey, Institute of Geological Sciences, London.

Description

Eleven fragments of a heteromorph with labeceratid coiling were partially
exposed by preparation from the body chamber of the large Tropaeum described
above (p. 285).

The holotype shows that coiling is very similar to that of Tonohamites
koeneni with the early part protruding beyond the venter of the aperture.
Ribbing on the early parts is only partially known, and appears to have been
strongly rounded and annular. Towards the crozier ribbing becomes more
prominent, and the ribs appear to arise at the umbilical edge, are slightly
sinusoidal across the flanks, and become progressively wider towards the
venter. Over the venter the ribs are broad and elevated, with high-walled
interspaces. Occasional bifurcations occur. The suture line is unknown.

Discussion

Initially only part of one specimen was exposed at the venter of the body
chamber of the Tropaeum sp. After preparation, parts of ten more were found,
but due to the hardness of the matrix much of the detail of the specimens has
been destroyed.

The labeceratid coiling, and the apparent absence of tuberculation are the
most obvious Tonohamites-like characteristics. The very striking broad, high-
crested ribs on the body chamber are very much like those found in hamitids of
the Hamites gibbosus type, and point to allocation to Helicancylus in the sense
of Casey (1961: 85), confirming once more the close relationship between the

CRETACEOUS FAUNAS FROM SOUTH AFRICA 325

various helicancylid genera, and the perhaps doubtful validity of retaining
Toxoceratoides, Tonohamites and Helicancylus separate.

None of the other Aptian heteromorphs from Zululand has the high-crested
crozier ribbing of 7. ? caseyi; neither has any of the Tonohamites or toxoceratoid
species described by Casey (1960).

Occurrence
Upper Aptian of Zululand.

Subfamily Heteroceratinae Spath, 1922

Heteromorphs with an initial helix or open spire, followed by planispirally
coiled whorls, a curved or straight shaft and recurved crozier are referred to the
Heteroceratinae.

Apart from the fact that the group is in need of restudy in the classical
European localities, there are some nomenclatorial difficulties, one of which
will be discussed below.

A perusal of the literature shows that authorship of the Heteroceratinae
(or -dae) is ascribed to either Hyatt 1900 (see Wright 1957: L212; Drushchitz &
Kudryavtzev 1960: 296; Dimitrova 1967: 64; Thomel 1964: 62; VaSicec 1972:
57, etc.) or to Spath 1922 (see Luppov & Drushchitz 1958: 105; Egojan 1965:
117; Dimitrova 1970: 94; Kakabadze 197la: 109, 19716: 39, etc.). In the
English version of Zittel’s Textbook of Palaeontology published in 1900, Hyatt,
however, never mentioned Heteroceratidae. He included Hamites Parkinson,
1811, Hemibaculites Hyatt, 1900, Dirrymoceras Hyatt, 1900 and Torneutoceras
Hyatt, 1900, in the Hamitidae, and added ‘here should also be placed Helicoceras
and Heteroceras d’Orb., and Lindigia, Karstens .. .’. Spath stated (1922: 148)
that ‘Heteroceras, Hemibaculites, Dirrymoceras and Lindigia may be united as
Heteroceratinae’, and from this it should be clear that authorship is referable to
Spath (1922) and not to Hyatt (1900).

Some of the most recent comprehensive publications on Heteroceratinae,
especially the colchiditids, are by Kotetitchvili (1970) and Kakabadze (1971a,
19715).

To the Heteroceratinae may be added Helicancyloceras gen. nov. (type
species Heteroceras (Argvethites ?) vohimaranitraensis Collignon, 1962) from the
Upper Aptian of Madagascar and possibly Kutatissites Kakabadze, 1970. The
subfamily Heteroceratinae is otherwise represented in the Barremian of Zululand
by Heteroceras sp. Colchidites gr. ex. colchicus spp., Colchidites sp. indet. and
Paraimerites sp. These species, and a detailed discussion on the Heteroceratinae
are given elsewhere (Klinger 1976).

Helicancyloceras gen. nov.
Type species
Heteroceras (Argvethites?) vohimaranitraensis Collignon (1962, pl. 14,
pl. 221 (fig. 962), from the Upper Aptian of Madagascar.

om

326 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 69. Tonohamites ? caseyi sp. nov. SAS Z804 from locality 151, Zululand, Aptian IV.
Photograph of cast of original, arrow indicates holotype. x 0,65.

Diagnosis

In typical forms, early whorls coiled in a shallow helix; later whorls may be
coiled planispirally in same plane as helix. Size varies considerably, and early
species may possibly be more loosely coiled. Ornament consists of ribbing with
or without two to three rows of tubercles on each side. Tubercles, if present,

become weaker or disappear at varying stages. Lateral, umbilical and internal
lobe trifid. Saddles bifid. Age: Upper Aptian.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 327

Helicancyloceras (Helicancyloceras) subgen. nov.
Diagnosis
Ribbing interrupted over venter in early whorls and ribs may end in small
tubercles on either side of the siphonal line. Adult ornament consists of non-
tuberculate ribs. Ribs may be flattened over venter with slight forward curvature
nearly touching each other, creating ‘paternoster’ beaded effect.

Helicancyloceras (Nonyaniceras) subgen. nov.
Type species
Helicancyloceras (Nonyaniceras) nonyani sp. nov.

Diagnosis

Planispiral coiling is attained rapidly. Early whorls with either ventral or
ventral and lateral rows of tubercles on every rib. With increasing diameter
intermediary non-tuberculate ribs occur. Tuberculate ribs may be stronger than
intermediaries. Umbilical tubercles, if present, very weak. Tuberculation may
disappear or become modified at various stages.

Discussion

Abundant fragments of this heteromorph genus occur in the Upper Aptian
of Zululand in the Mkuze Game Reserve and along the Mfongozi Creek. Due to
the general fragmentary nature of the material, only a few well-defined species
may be recognized. The rest of the material is either mentioned or described
without specific reference.

The majority of specimens consists of small helices only. Fortunately one
specimen (Figs 70-71) shows remains of helical inner whorls, followed by a
massive planispiral phragmocone and parts of the body chamber. The authors
were at first inclined to regard the helically coiled forms as the ancestors of the
larger forms with planispiral outer whorls. Detailed collecting, however, showed
that both forms can occur at the same stratigraphic level. Whether the two size
groups represent juvenile and adult stages respectively or sexual dimorphs is not
known. The predominance of small helices may be due to selective postmortal
transport, and may not reflect the true character of the population. In general,
however, specimens with well-developed helices dominate at lower stratigraphic
levels whereas forms with very shallow helices appear to be younger.

The affinities of the genus are enigmatic. At first glance the helical coils
recall the Middle Albian hamitids once referred to d’Orbigny’s genus Helicoceras.
These, however, are true hamitids with a bifid lateral lobe. The helical coiling in
Helicancyloceras points strongly to the Heteroceratinae, whilst the tuberculate
Species show ancyloceratine or perhaps helicancyline affinities. As has been
pointed out earlier, the subfamilies are primarily accepted because they feature
a certain degree of morphological homogeneity. Reference of Helicancyloceras
to the Heteroceratinae rather than the Ancyloceratinae or Helicancylinae is thus
based entirely on morphological criteria.

328 ANNALS OF THE SOUTH AFRICAN MUSEUM

None of the heteroceratids appears suitable as an ancestral form, mainly
because of the time gap between them and Helicancyloceras. The heteroceratids
with the possible exception of the dubious Hemibaculites Hyatt, 1900, are
restricted to the Barremian. Nevertheless, the relationship of H. (Helicancylo-
ceras) to H. (Nonyaniceras) is similar to that of H. (Heteroceras) to H. (Argve-
thites) or of Colchidites to Imerites.

The very enigmatic genus Kutatissites Kakabadze, 1970 (type species
K. bifurcatus Kakabadze, 1970), thus far known only from the Late Barremian
of Georgia (U.S.S.R.) and France is another helically coiled form which may be
considered ancestral. Thieuloy (1976) provides an extensive description of the
French representative of this species, K. edwardsi (Reynes, 1876) and tentatively
refers the genus to the Helicancylinae. Again the difference in age (late
Barremian vs late Aptian) seems to stand in the way of deriving Helicancyloceras
directly from Kutatissites.

Representatives of the Ancyloceratinae may provide the answer. Helicancy-
loceras could possibly be derived from an australiceratid such as Australiceras
irregulare (Tenison Woods) with irregular early whorls not lying in one plane
(see Day 1974, pl. 2 (Fig. 3)). Australiceras loses tuberculation in the middle
stages of life. In the hypothetical ancestor of Helicancyloceras (Helicancyloceras),
reduction of tuberculation could have affected the early whorls already, whereas
the reduction in H. (Nonyaniceras) could have been retarded. In this case the
relationship of H. (Helicancyloceras) to H. (Nonyaniceras) would be somewhat
analogous to that of Tropaeum to Australiceras as far as the presence or absence
of tubercles is concerned.

Whatever the origin of Helicancyloceras may be, the ornamentation and
mode of coiling combines features of Ancyloceratinae, Helicancylinae and
Heteroceratinae, again pointing to the rather artificial status of these groups.
Reference of Helicancyloceras to the Heteroceratinae is done for pure morpho-
logical reasons, in full cognizance that this might imply a diphyletic origin for
the Heteroceratinae (if they are to be retained as a separate group).

Furthermore, Helicancyloceras illustrates another attempt towards
recoiling, and thus, (presumably) a more active mode of life for the animals
concerned.

Helicancyloceras (Helicancyloceras) vohimaranitraensis
(Collignon, 1962)
Figs 70, 71A—B, 73B, D, 74A—F, 75B—-C, E-G, 76A-B?, 77D-E, ? F—G, 78C,
88A—C

Heteroceras (Argvethites ?) vohimaranitraensis Collignon 1962: 14, pl. 221 (fig. 962). Klinger
1976: 14, pl. 1 (nos. 3-5), figs 5d—e, 6b-e.

Holotype

Collignon’s (1962, pl. 221 (fig. 962)) specimen from the Upper Aptian of
Betioky, Madagascar.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 329

Fig. 70. Helicancyloceras (Helicancyloceras) vohimaranitraensis (Collignon, 1962). SAS

H71D from locality 152, Mkuze Game Reserve, Zululand, Aptian IV. Largest available speci-

men with helicoid. inner whorl, but completely symmetrical planispirally-coiled outer whorl.
x 0,86. (See also Fig. 71.) Possible sexual dimorph?

330 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 71. A-B. Helicancyloceras (Helicancyloceras) vohimaranitraensis (Collignon, 1962).
SAS H71D from locality 152, Mkuze Game Reserve, Zululand, Aptian IV. Note the helical
inner whorl. x 0,75.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 335)!

Fig. 72. A-E. Helicancyloceras (Nonyaniceras) circulare sp. nov. BMNH C79718 from locality
166, Zululand, Aptian III. x 2.

332 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 73. A. Tonohamites koeneni Casey, 1961. SAS Z1115 from locality 152, Zululand, Aptian

IV. x 1,3. B, D. Helicancyloceras (Helicancyloceras) vohimaranitarensis (Collignon, 1962).

SAS Z9(5) from locality 168, Zululand, Aptian III-IV. This is an extremely coarsely ribbed
specimen. x 1,6. C. Helicancyloceras (Helicancyloceras) sp. indet. x 1,5.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 333

B's E F

Fig. 74. A-F. Helicancyloceras (Helicancyloceras) vohimaranitraensis (Collignon, 1962).
A-C. BMNH C74709 from locality 166, Zululand, Aptian III. x 2. D-F. BMNH C79711,
same locality and age as above. x 2.

334 ANNALS OF THE SOUTH AFRICAN MUSEUM

Material

SAS Z8(6), SAS Z8/3, SAS 871D/12, SAS H71D, SAS Z1153, all from
locality 168 and 152. Aptian III-IV: BMNH C79703-—C79712, all from
locality 166, Aptian III.

Description

Early stages of growth are helical, either dextral or sinistral, consisting of —
about one and a half to two whorls. The distance between the successive whorls
is extremely variable. After the helical stage coiling becomes planispiral in some
specimens.

Ornament in the helix consists of prominent ribs, narrowest and least
prominent on the dorsum, wedge-shaped on the flanks and broadest over the
venter. Interspaces on the flanks are about as wide as the ribs. Ribbing over the
venter is very variable. At small diameters the ribs are interrupted or weakened
on the venter, in some cases forming small pimple-like tubercles on either side
of the siphonal line. At large diameters the ribs cross the venter with a slight
forward curvature and a thickening in the interspaces, thus forming a ‘pater-
noster’ effect. In the planispiral part of the shell the ribbing becomes finer and
wider-spaced and somewhat rursiradiate. Ribbing over the dorsum is very weak
at this stage. Occasional bifurcations occur at the umbilical edge. On the body
chamber the whorl section becomes dome-shaped with a flattened dorsum and
broadly inflated flanks and venter. The suture line is relatively simple with bifid
saddles and trifid L, U, I lobes. L is largest and U smallest.

Dimensions
Specimen D Wb Wh Wb/Wh U H
(height of
spire)
H71D 175 64(36,7) 76(44,7) 0,84 64(36,5) —
SAS Z9(5) 46 14(30,4) 12(26) 117 22(47,8) 23
Discussion

As can be seen from the figures, the species is interpreted rather widely, and
is intimately connected to H. (H) densecostatum sp. nov. by finer-ribbed
specimens such as BMNH C79713 (Fig. 75B-C), SAS Z8/Hd (Fig. 76B) and
SAS A1294 (Fig. 79C). It is similarly connected to H. (N.) circulare from which
it differs in becoming planispiral at later stages and in lacking regular tuber-
culation in the early stages of growth.

The occurrence of both large and small forms at the same stratigraphic level
(e.g. SAS H71D and SAS H71D/12, Figs 70, 71A—B, 76A) is of interest; sexual
dimorphism seems the most likely explanation. The helical inner whorls of the
large specimen SAS H71D are indistinguishable from isolated helices or
fragments of helices, indicating conspecificity, and, that in some cases at least,
the shell changes to a planispiral mode of coiling after an initial helical stage.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 335

The inner whorls are easily distinguished from other heteromorphs
occurring at this stratigraphic level by the helical coiling and curious mode of
ornament over the venter. Small fragments bear similarity to the inner whorls of
Tropaeum, but are distinguished by the oblique ribbing. Small pieces of
Tonohamites koeneni, which occur at the same level, are distinguished by the
same character.

The outer, planispiral whorls, with symmetrical ribbing are easily mistaken
for Tropaeum or Australiceras, and; unless associated with the inner whorls,
may be indistinguishable.

Occurrence

The species occurs in the Upper Aptian zone of Epicheloniceras tscherni-
schewi of Madagascar, and Aptian IIJ-IV of Zululand.

Helicancyloceras (Helicancyloceras) densecostatum sp. nov.
Figs 67A-B, 75B-C?, I-J, 76B?, 79C, 81D
Heteroceras (Argvethites ?) vohimaranitraensis Klinger, 1976: 14 (pars), pl. 2 (no. 5) only.
Holotype

SAM-PCZ5682 from locality 168 Mfongozi Creek, Zululand. Aptian
I-IV. South African Museum Collection. Collected by H. Klinger.

Material

Apart from the holotype, SAS LJE138, SAS A294, A1294, SAS Z8/6 from
locality 168 and BMNH C79713 from locality 166 Mfongozi Creek, Zululand.
Aptian IJIJ-IV.

Description

The initial helix is very low and consists of about one whorl. Coiling
becomes planispiral at an early stage. The whorl section is rounded throughout.

Ornament consists of very fine ribs, approximately twenty-five per half
whorl, separated by interspaces of comparable width. On the earliest whorls
preserved, the ribbing is distinctly interrupted over the venter, each rib ending
in a minute tubercle on either side of the siphonal line. With increasing diameter,
the ventral interruption of ribbing disappears and is present as a mere swelling
on each rib along the siphonal line, again creating a ‘paternoster’ effect. On
specimen SAS A1294 (Fig. 79C) ventral tubercles occur on the body chamber
on two ribs, separated by five intermediaries.

Dimensions
Specimen D Wb Wh Wb/Wh U H
SAM PCZ5682 AD? £8(42,8) ~16(8,1) lei 23(50) 14

Discussion

The diagnostic characteristics of the species are the fine ribbing and the
very shallow helix. Whereas H. (H) vohimaranitraensis has about 15 ribs per

336 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 75. A, D, H. Helicancyloceras (Nonyaniceras) sp. indet. BMNH C79698 from locality

166, Zululand, Aptian III. x 2. B-C. Helicancyloceras (Helicancyloceras) vohimaranitraensis

(Collignon, 1962). BMNH C79704 from locality 166, Zululand, Aptian III. x 2. Transitional

to H. (H.) densecostatum. E-G. Helicancyloceras (Helicancyloceras) vohimaranitraensis

(Collignon, 1962). BMNH C79705 from locality 166, Zululand, Aptian III. I-J. Helicancylo-

ceras (Helicancylo ceras) densecostatum sp. nov. BMNH C79706 from locality 166, Zululand,
Aptian III. x 2.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 337

Fig. 76. Helicancyloceras (Helicancyloceras) vohimaranitraensis (Collignon, 1962). A. SAS
H71D/12 from locality 152, Zululand, Aptian IV. x 1,8. B. SAS Z8/Hd x 1,5 from locality
168, Aptian III-IV. Transitional to H. (H.) densecostatum sp. nov. X 1,7.

q
a.

338 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 77. A-C. Helicancyloceras (Nonyaniceras) circulare sp. nov. BMNH C79708 from locality

166, Zululand, Aptian HI. x 2. D-E. Helicancyloceras (Helicancyloceras) vohimaranitraensis

(Collignon, 1962). BMNH C79710 from locality 166, Zululand, Aptian III. x 2. F—G. Helican-

cyloceras (Helicancyloceras) sp. indet. BMNH C79693 from locality 166, Zululand, Aptian III.
<2,

CRETACEOUS FAUNAS FROM SOUTH AFRICA 339 I

Fig. 78. A-B. Helicancyloceras (Nonyaniceras) circulare sp. nov. SAS H71/2, holotype from N

locality 151, Zululand, Aptian IV. x 1,5. C. Helicancyloceras (Helicancyloceras) vohimaranitra-
ensis. A1924 from locality 168, Aptian III-IV.. x 1,5. D. Helicancyloceras (Nonyaniceras)
sp. indet. SAM—PCZ5504 from unknown locality. southern part Mkuze Game Reserve. x 1,5.

340 ANNALS OF THE SOUTH AFRICAN MUSEUM

half whorl, H. (H) densecostatum has about 25. Transitional forms like SAS
Z8/Hd (Fig. 76B) with 21 ribs per half whorl connect the species and may be
referred to either. The presence of tubercles on the body chamber of some
specimens points to affinities with H. (Nonyaniceras). The presence of tubercles
in the very early and late stages with a non-tuberculate middle stage is a situation
comparable to that found in Australiceras. It is not known if large forms also
occur in this species, but the outer whorl is virtually bilateral symmetrical.

Occurrence

Upper Aptian of Zululand.

Helicancyloceras (Helicancyloceras) spp. indet.

Several small fragments of heteromorphs with MHelicancyloceras-like
ornament are too poorly preserved for specific identification. A specimen in the
collection of the Geological Survey, Z8/21, here figured as Figure 81E is a
slightly twisted fragment with the typical ornament over the venter, but does not
appear to have been helically coiled. BMNH C79693 is also only slightly
curved, indicating looser coiling in, as yet, unknown species of the genus.

Helicancyloceras (Nonyaniceras) nonyani sp. nov.
Figs 82A-B, 83B
Holotype

SAS LJE182 from locality 168, Mfongozi Creek, Zululand. Aptian III-IV.
Geological Survey Collection. Collected by L. J. Engelbrecht.

Derivation of name

The species is named for Mr Johannes Nonyane, Klinger’s field assistant
and laboratory preparator from 1967 to 1975.

Material
Only the holotype.

Description

The early whorl forms a very shallow helix, and on the second, and last
whorl, coiling is perfectly bilaterally symmetrical at the aperture. The body
chamber occupies slightly less than one half of a whorl of the holotype.

The whorl section on the early whorl is round, but becomes ovoid, higher
than wide in the body chamber. Initial ornament consists of narrow, rounded
radial ribs, separated by interspaces of similar width, each bearing a pair of
ventral tubercles. At a diameter of approximately 8 mm, ornament becomes
irregular, lateral tubercles appear and the ribs develop a prorsiradiate curvature.
On the body chamber the ribs pass over the dorsum with a slight forward
curvature, but much weakened. The ribs are narrowest at the umbilical edge,

CRETACEOUS FAUNAS FROM SOUTH AFRICA 341

becoming broader towards the venter. Five ribs on the body chamber have
prominent ventral tubercles, separated by one to three intermediaries. Lateral
tubercles situated on the ventral third of the flanks are developed only on one
side, due to injury. Ventral tubercles are variably developed on the inter-
mediaries. They may appear as distinct tubercles or merely as slight edges on
the ventrolateral sides of the whorl. The major ribs are slightly thickened and
flattened between the lateral and ventral tubercles, and also over the venter,
forming incipient loops. Slight thickenings occur between the ribs along the
siphonal line, creating a ‘paternoster’ effect.

Discussion

The distinct ventral tuberculation and prominent ribbing on the body
chamber clearly separates this species from other Helicancyloceras and deserves
subgeneric rank. The angular costal whorl section on the body chamber is
superficially similar to that of Pedioceras cundinamarcaea Karsten, 1886, from
the Barremian of Colombia (see Wright, 1957: L202, fig. 237: 5a—c). That
however, is as far as the similarity goes. Pedioceras cundinamarcaea has a
distinct dorsal zone of impression and is coiled planispirally.

Occurrence
Upper Aptian, Aptian HI-IV of Zululand.

Helicancyloceras (Nonyaniceras) circulare sp. nov.

Figs 72A-E, 77A-C, 78A—B
Holotype
SAS H71/2 from locality 151, Mkuze Game Reserve. Aptian IV. Geological
Survey Collection. Collected by H. C. Klinger.

Material

The holotype and BMNH C79698, BMNH C79708, BMNH 79718 from
locality 166, Aptian III.

Description

The initial helix is very shallow as can be seen in Figure 77A-C, and
bilateral symmetry is attained rapidly. The whorl section is circular. Ornament
on the inner whorl consists of fine, slightly prorsiradiate ribs, each bearing weak
ventral and lateral tubercles. On the outer whorl the ribs are stronger,
prominently rounded and separated by narrow interspaces. Two quadri-
tuberculate ribs, separated by three non-tuberculate ribs, are present at the
smaller end of the outer whorl of the holotype. The tubercles, lateral and ventral,
are very small and pointed.

342 ANNALS OF THE SOUTH AFRICAN MUSEUM

Discussion

The presence of the two rows of tubercles on all ribs of the inner whorl,
and the strong, rounded ribbing on the outer whorl, combines features of both
subgenera, showing the close relationship between the two. The outer whorl
might easily be mistaken for a juvenile Tropaeum, as coiling is already planispiral
and, unless accompanied by the inner whorls, distinction from Tropaeum would
be difficult. The very low helix plus the presence of tubercles on the inner whorls
and at larger diameters serves to distinguish the species from H. (H.) vohimarani-
traensis.

Occurrence
Upper Aptian of Zululand, Aptian IV.

Helicancyloceras (Nonyaniceras) crassetuberculatum sp. nov.
Fig. 79E-F
Holotype

SAS L65/13 from locality 168, Mfongozi Creek, Zululand. Aptian IHI-IV.
Geological Survey Collection. Collected by H. Klinger, 1970.

Material
Apart from the holotype, SAS L65/14 from the same locality.

Description

The holotype consists of half a whorl. The whorl section is rounded. The
original helix must have been very shallow, but the helical coiling is nevertheless
rendered distinct by the asymmetry of ornament over the venter. Ornament
consists of major trituberculate ribs, separated by one to three narrower
intermediaries. The umbilical tubercles are smallest, and virtually absent at the
smallest diameters visible. The ventrolateral and ventral tubercles are conical,
very prominent, and situated close together. On the flanks the trituberculate
ribs show signs of flattening or duplication. Ribbing is weakened on the dorsum,
and the major ribs duplicate at larger diameters. The intermediary ribs are
weakly tuberculate over the venter at small diameters. With increasing diameter
the tuberculation becomes less noticeable.

Discussion

H. (N.) crassetuberculatum is distinguished from other species of the
subgenus by the very prominent tuberculation at small diameters. Fragments
bear similarity to Ancyloceras, but the asymmetry of ornament over the venter
reveals the generic affinity.

Anderson’s Helicancylus gabbi (1938: 222, pl. 79 (figs 4-5)) is a similar
helical form of uncertain generic affinity. The ribbing, however, is much
stronger and there are fewer intercalatories. As yet the systematic position of

CRETACEOUS FAUNAS FROM SOUTH AFRICA 343

Fig. 79. A-B. Toxoceratoides? haughtoni sp. nov. SAS Z8/Ti from locality 168, Zululand,

Aptian III-IV. x 1,6. C. Helicancyloceras (Helicancyloceras) densecostatum sp. nov. SAS

A1294 from locality 168, Zululand, Aptian III-IV. x 1,55. D. Ancyloceras (Ancyloceras ?)

sp. indet. SAS H54/39 from locality 170, Zululand, Aptian I. x 1,5. E-F. Helicancyloceras

(Nonyaniceras) crassetuberculatum sp. nov. SAS L65/13, holotype from locality 168, Zululand,
Aptian ITI-IV. x 1,5.

344 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 80. A. Suture line of Australiceras sp. aff. irregulare (Tenison Woods, 1883). SAM-—
PCZ5506. x 3. B. Suture line of Z8n, Insertae sedis. x 1.

this species is not known, and it is not certain if the specimen is congeneric with
the hamulinid Helicancylus aequicostatus (Gabb).

Ancyloceras helicoides Rouchadzé (1933: 17, pl. 8 (fig. 3) is another
ancyloceratid form, but again the tuberculation is much stronger and is referred
to Kutatissites of Late Barremian age.

Occurrence

Upper Aptian of Zululand.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 345

Fig. 81. A, C. Helicancyloceras (Nonyaniceras) sp. indet. SAS Z8(7) from locality 168, Aptian

IlI-IV. x 1. B. Helicancyloceras (Nonyaniceras) sp. indet. SAS H71D/15 from locality 152,

Zululand, Aptian IV. x 1. D. Helicancyloceras (Helicancyloceras) densecostatum sp. nov.

SAS LJE 138 from locality 168, Zululand, Aptian III-IV. x 1. E. Helicancyloceras (Helican-

cyloceras) sp. indet. Unnumbered specimen SAS collection. x 1. F. Tonohamites koeneni
Casey, 1961. SAS Z193 from locality 152, Zululand, Aptian IV. x 1,5.

m1

346 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 82. A-B. Helicancyloceras (Nonyaniceras) nonyani sp. nov. SAS LJE182, holotype from
locality 168, Zululand, Aptian IIJ-IV. x 1,6 (See also Fig. 83B.)

ie.
CRETACEOUS FAUNAS FROM SOUTH AFRICA 347 ‘y

Fig. 83. A. Lithancylus sp. SAM—PCM5436 from Lubemba, Mozambique, Lower Aptian. F
Dorsal view. x 1,2. B. Helicancyloceras (Nonyaniceras) nonyani sp. nov. SAS LJE182, N
holotype from locality 168, Zululand, Aptian IIJ-IV. Tuberculation absent on body chamber
due to pathological disturbance. x 1. C—D. Helicancyloceras (Nonyaniceras) sp. indet.
SAM-PCZ5504 from unknown locality in southern part of Mkuze Game Reserve. x 1,3.

348 ANNALS OF THE SOUTH AFRICAN MUSEUM

B G

Fig. 84. Lithancylus sp. SAM-—PCMS5436. Parts of the suture line. x 1,5.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 349

Helicancyloceras (Nonyaniceras) spp. fragments

Several fragments are definitely referable to the genus, but are too frag-
mentary or poorly preserved to merit specific status. SAS Z8(7) (Fig. 81A, C)
shows that coiling may have been very loose in some species.

SAS H71D/15 (Fig. 81B) seems to represent the aperture of a small helix.

Two very worn specimens, SAM-PCZ5504 (Figs 83C-D, 78D) and
BMNH C79162 (Fig. 68G—H) have ribbing very similar to that of H. (N.)
circulare, but have very strongly developed trituberculate ribs. The ventral
tubercles are largest and the umbilical ones smallest and pointed.

Incertae sedis

Two specimens from the Upper Aptian of the Mfongozi Creek cannot be
referred with certainty to any ancyloceratid genus. Z8(1) (Fig. 85A—B) is part of
a phragmocone with definite impressions of tubercles on the dorsum. Ornament
consists of slightly sinusoidal ribs separated by narrow interspaces. At the
smaller end there is a distinct interruption of ribbing over the venter, but at the
larger there is no trace of this. The suture line is partially exposed, showing a
large, bifid E/L saddle, a long, narrow trifid L and a very small trifid U.

The impression of tubercles on the dorsum and the ribbing in general is
reminiscent of specimens of A. (Adouliceras) mozambiquense, but the former is a
Lower Aptian species. It could possibly be an australiceratid.

Z8n (Fig. 86) is a J-shaped fragment of an uncoiled ancyloceratid ammonite.
Ornament is very weak, consisting of very weak ribs which are radial over the
flanks and curve broadly forward over the dorsum. Ribbing is absent on part
of the internal mould. The whorl section is more or less oval throughout. Again
affinities with the Early Aptian uncoiled forms of A. (Adouliceras), Tropaeum
and Australiceras have to be ruled out because of its occurrence in the Upper
Aptian. Egojan (1974) recently described Upper Aptian uncoiled ammonites
from the Caucasus, which he referred to a new family, Epacrioceratidae. The
main characteristic of the family is the presence of a bipartite lateral lobe (L)
and a smaller trifid U and I. In the present specimen the lateral lobe is distinctly
asymmetric (Fig. 80B) but is more trifid than bifid. On the basis of the limited
material further speculation on the affinity of the Zululand specimen with the
Caucasian material serves little purpose.

PALAEOBIOGEOGRAPHY

The distribution of the Aptian ammonoid genera Ancyloceras (Adouliceras),
Tropaeum, Australiceras, Lithancylus, Toxoceratoides and Tonohamites is
summarized in Figure 89.

The sources of these data are as follows:

Lithancylus Casey 1961 (England & Tyrol); Anderson 1938 (California);

Day 1974 (Australia); Leanza 1970 (Patagonia); Thomson 1974 (Alexander

Island).

350 ANNALS OF THE SOUTH AFRICAN MUSEUM

oe

A ee

Fig. 85. Insertae sedis. SAS Z8(1) from locality 186, Zululand, Aptian III-IV. x 1,3.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 351

Fig. 86. Insertae sedis. SAS Z8n from locality 186, Aptian III-IV. x 0,5.

Tropaeum Casey 1960 (England); Sinzow 1872, 1905 (Caucasus); Von Koenen
1902 (Northern Germany); Dimitrova 1967 (Bulgaria); Stolley 1912,
Frebold 1935 (Spitzbergen); Spath 1946 (Greenland); Jeletzky 1964
(Arctic Canada); Nakai 1968, Obata et al 1975 (Japan); Wilckens 1947
(south Georgia, Antarctica); Leanza 1963 (Patagonia?); Da Silva 1962
(Mozambique); Haughton 1936 (Zululand); Thomson 1974 (Alexander
Island ?); Etheridge 1909; Whitehouse 1926; Day 1974 (Australia); Royo y
Gomez 1945 (Columbia); Spath 1931 (India); Forster 1975a (Mozambique).

352 ANNALS OF THE SOUTH AFRICAN MUSEUM

Australiceras Casey 1960, 1961 (England); Sinzow 1905 (Caucasus); Dimitrova
1967 (Bulgaria ?); Royo y Gomez 1945 (Columbia); Waagen 1875 (India);
Anderson 1938 (California); Favre 1908 (Patagonia?); Thomson 1974
(Alexander Island); Haughton 1936; Kennedy & Klinger 1975 (Zululand);
Forster 1975a (Mozambique); Obata et al. 1975 (Japan).

Tonohamites Casey 1961 (England); Von Koenen 1902 (northern Germany);
Collignon 1962 (Madagascar); Kennedy & Klinger 1975 and herein
(Zululand).

Toxoceratoides Casey 1961 (Western Europe); Forster 1975a (Mozambique);
Thomson 1974 (Alexander Islands); Murphy 1975 (California).

Ancyloceras (Adouliceras) Thomel 1964 (France); Anderson 1938 (California
and Oregon?); Dames 1880 (Germany); Zululand (herein).

Interpretation of data

The main picture that emerges, not only from the map, but from the
systematic descriptions, is the close relationship between the faunas of the
southern part of European U.S.S.R. and Western Europe, and Zululand and
Mozambique. Open marine connections clearly existed in both Aptian and
Barremian times between Tethys and the eastern parts of Africa and India, to
facilitate free migration.

The most primitive forms of Tropaeum, Australiceras and Ancyloceras, are

eo.

Fig. 87. Suture line of Tropaeum subsimbirsteense compressum. X 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 353

Fig. 88. A-C. Suture lines of Helicancyloceras (Helicancyloceras) vohimaranitraensis (Collignon,
1962) from unnumbered fragments. x 6. D. Suture line of Tonohamites aequicingulatus
(von Koenen, 1902). SAS Z8/Tal. x 6.

ANNALS OF THE SOUTH AFRICAN MUSEUM

354

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CRETACEOUS FAUNAS FROM SOUTH AFRICA 355

to be found in the region ranging east-west from Great Britain to east of the
Caspian Sea, and current evidence suggests this as the source area of the group,
genera migrating from there to other parts of the world during early Cretaceous
time. There is great similarity between the faunas of Zululand and Mozambique
and those of Madagascar, India, Australia and to a lesser extent south-eastern
South America and the Antarctic regions. There is little similarity, however,
with the west coast faunas of the Americas.

Part of the dispersal pattern shown in Figure 89 may be explained by taking
ocean currents prevailing during the Early Cretaceous into account. A study of
this kind was recently undertaken by Gordon (1973), the results of which are
here briefly summarized and illustrated in the insert in Figure 89. Due to the
absence of the Panama Isthmus, a westward-flowing circumglobal tropical
current existed. Apart from assisting in distributing heat around the globe, the
equatorial current was forced northwards into the Tethyan region by the bulge
of Africa—Arabia. By this mechanism the Tethyan fauna was distributed around
the globe. Water from this current was probably deflected southwards along the
eastern coast of Africa by the ‘horn’ of Africa, past Mozambique and Zululand.
Due to the higher temperature of this current, the Tethyan faunas could survive
at these high latitudes. The absence of comparable faunas on the west coast of
South America seems to indicate the presence of a cold current flowing
equatorialward. Due to west-wind drift, an eastward-flowing current must have
existed at high latitudes in the Southern hemisphere. Due to the mixing of the
Antarctic current and the southward-flowing water from the equator, a low
temperature differential existed, allowing part of the Tethyan fauna to be
distributed eastwards. This may explain the similarities between the faunas of
the southern parts of South America, the Antarctic regions, south-east Africa,
Madagascar, India and Australia. The similarity between the south-east African
and Australasian fauna becomes even more obvious during the Albian as shown
by the distribution of the labeceratids, or during the Maastrichtian by the
eubaculitids.

It is also interesting to note that, apart from the Columbian occurrence of
Australiceras, the majority of ancyloceratids appear to have lived in temperate
regions between latitudes 20 to 60 degrees.

ACKNOWLEDGEMENTS

The authors are grateful to many of their colleagues for advice, stimulating
discussions and constructive criticisms, and for providing comparative material,
literature and access to their collections: Professor Dr J. Wiedmann (Tiibingen),
Dr R. Forster (Munich), Professor Dr D. H. Wachendorf (Braunschweig),
Professor J.-P. Thieuloy (Grenoble), General M. Collignon (Moirans), Professor
G. Thomel (Nice), Dr R. Casey (London), Dr M. K. Howarth and Mr D.
Phillips (British Museum (Natural History)), Mr C. W. Wright (London),
Dr R. Day (Brisbane), Professor J. Visser (Pretoria) and Dr M. R. Cooper
(Oxford).

356 ANNALS OF THE SOUTH AFRICAN MUSEUM

Both authors express their thanks to the Director of the Geological Survey
of South Africa (Pretoria) for putting some of the material at their disposal.

Cordial thanks are due to Dr P. Bengtson (Uppsala) for providing a
photograph of the holotype of Australiceras ramososeptatum.

Assistance with the photography by Messrs N. Eden (South African
Museum, Cape Town) and W. Wetzel (Tiibingen) is gratefully acknowledged.
Mr C. Hunter and Mr V. Branco (South African Museum) kindly assisted with
some of the illustrations.

Part of this study was undertaken during the tenure of a D.A.A.D. grant
during 1972/3 to Klinger. Part of Kennedy’s expenses during the 1970 field
season was defrayed by a grant from the Sir Henry Strackosch Bequest; the
assistance of the Natural Environment Research Council is also gratefully
acknowledged.

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~wa:-

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:

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

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

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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.
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“Ca”

wi Ue

HERBERT CHRISTIAN KLINGER

&
WILLIAM JAMES KENNEDY

CRETACEOUS FAUNAS FROM ZULULAND,
SOUTH AFRICA AND SOUTHERN MOZAMBIQUE
THE APTIAN ANCYLOCERATIDAE (AMMONOIDEA)

YF THE SOUTH AFRICAN |

CAPETOWN

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

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

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

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

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

THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
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 N
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM f

Volume 73 #£4Band |
November 1977 November
Part 10 #£Deel

A NEW SPECIES OF HALICYCLOPS
(COPEPODA, CYCLOPOIDA)
FROM ESTUARIES IN TRANSKEI, SOUTHERN AFRICA

By

T. WOOLDRIDGE

Cape Town Kaapstad in

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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

A NEW SPECIES OF HALICYCLOPS (COPEPODA, CYCLOPOIDA)
FROM ESTUARIES IN TRANSKEI, SOUTHERN AFRICA

By
T. WOOLDRIDGE
Department of Zoology, University of Port Elizabeth

(With 3 figures and | table)

[MS. accepted 28 April 1977]

ABSTRACT

A new species of the genus Halicyclops Norman, 1903 is described and illustrated from
estuaries in Transkei, southern Africa. The species, Halicyclops pondoensis, is closely allied to
H. neglectus, H. rotundipes and H. higoensis. \t is distinguished from these three forms particu-
larly in regard to the fourth and fifth pair of legs. In H. pondoensis hair-like structures are
present in the proximal part of the seta of the first endopodite segment of leg 4. These hair-like
structures are absent in other species. Two spines and three setae are present on leg 5 of the
male of H. pondoensis. In H. neglectus, H. rotundipes and H. higoensis the male Sth leg bears
three spines and two setae. In the present species the second segment of the Sth leg in both the
male and female is elongated.

A key to the species recorded from the African continent south of the Tropic of Cancer,
and from the islands of Madagascar and Réunion, is given.

CONTENTS

PAGE
Introduction. . “. . . = 361
Description of material . . . 362
BISGUSSION.-¢ 3520 ees. 5 368
Acknowledgements . . . . 370
Ieterenices- 9 wk TO

INTRODUCTION

A new species of the cyclopoid copepod genus Halicyclops was found in
plankton samples collected from estuaries on the east coast of southern Africa.
The genus has a world-wide distribution and is recorded from marine, brackish
and fresh-water habitats. Although most species are free-living, a number are
reported from the interstitial fauna of marine beaches. Species of Halicyclops
are also found on the body surface or in the tubes of polychaete worms.
Halicyclops caridophilus inhabits the gill chamber of the mudlobster, Thalassina
anomala.

The genus is so far known from 47 described forms. The present species is
the first member of the genus described from estuaries in southern Africa,
although Hill (1966) tentatively recorded H. pilifer from the Umlalazi estuary
(28°57’S), Zululand. The new species of Halicyclops reported here is recorded

361

Ann S. Afr. Mus. 73(10), 1977: 361-371, 3 figs, 1 table.

e~e
rr

ove
Sr

a

mae oP Se

362 ANNALS OF THE SOUTH AFRICAN MUSEUM

from Msikaba estuary (31°19’S) and Mbotyi estuary (31°28’S) on the Pondoland
coast, Transkei. Specimens were taken at all seasons from May 1972 to March
1973 in plankton tows collected from just below the water surface in salinities
ranging from 1-33%,.

DESCRIPTION OF MATERIAL

Halicyclops pondoensis sp. nov.
Figs 1-3
HOLOTYPE

SAM-—A15614 in the South African Museum, Cape Town. Adult female from
Mbotyi estuary on the Pondoland coast (31°28’S), collected by T. Wooldridge,
29 November 1972.

PARATYPES

SAM-A15614 in the South African Museum, Cape Town. Numerous adult
males and adult females from Mbotyi estuary on the Pondoland coast (31°28’S),
collected by T. Wooldridge, 29 November 1972.

Description

Female. Total length 0,61-0,68 mm (mean of five specimens 0,65 mm). Meta-.
some about twice the length of the urosome. Rostrum small, bluntly triangular,
curved ventrally and posteriorly, and hence not visible from the dorsal side.

Cephalic segment bluntly rounded anteriorly, wider than long (1,25—1,33: 1),
the greatest width in the posterior region (Fig. 1A). Second thoracic segment
longer than third and fourth respectively, which are equal in length. The posterior
lateral edges of second and third segments free, angular; those of the fourth
segment free but more rounded. Fifth thoracic segment short, its posterior
lateral margins bluntly pointed and bearing the fifth pair of legs.

Genital segment (Fig. 2A) with small protrusions on the anterior lateral
margin. Length subequal to the width. Urosomal segments posterior to genital
segment progressively shorter and narrower, the anal segment with a deep medial
incision. Posterior border of the first three segments of urosome denticulate.
Anal segment serrated on the posterior ventral border only. A row of fine
spinules on anterior dorsal side of the anal segment.

Furcal rami (Fig. 2A) slightly longer than wide and about equal in length to
the anal segment. All furcal setae usually found in the genus present, the proximal
dorso-lateral seta reaching slightly beyond furca. Dorsal seta about twice the
length of the dorso-lateral seta, arising from the inner posterior region.
Ornamented basally with minute spinules. Apical setae long and stout, the inner
as long or slightly longer than the length of the metasome, the outer less than
half as long. Proximal half of shorter and less stout seta spinose on its outer
margin only, distal half plumose, ornamented basally on dorsal surface by a row

A NEW SPECIES OF HALICYCLOPS FROM ESTUARIES IN TRANSKEI 363 NI

wre
ldo)

ere
Gnhy

"AIL

E

Fig. 1. Female. A. 0,62 mm female. B. Antennule. C. Antenna. D. Mandible. . i
E. Maxilla 1. F, Maxilliped. 3

oh

364 ANNALS OF THE SOUTH AFRICAN MUSEUM

of short spinules. Inner apical seta spinose on lateral margins in proximal half,
the spines becoming less sparse posteriorly. Terminal half plumose. Innermost
furcal seta short, about one-third the length of ramus. Outermost seta non-
plumose, about one and a half times length of ramus.

Antennule six segmented (Fig. 1B), almost half as long as cephalic segment.
Ultimate and penultimate segment combined slightly longer than anti-penultimate
segment. Setation and relative lengths illustrated in Figure 1B. Aesthetask
present on distal margin of ultimate segment.

Antenna three segmented (Fig. 1C), with two setae on anterior lateral
border of first segment. Inner lateral margin with a number of short spines.
Second segment with one seta. Third segment about three and a half times as
long as broad, bearing five lateral setae. Two lateral setae, of which one is
relatively stout, born on raised portion midway along length of segment.
Terminal group consisting of three slender and four stout setae, the longest
placed sub-terminally. A row of fine spinules along inner lateral border of
ultimate segment.

Mandible slender (Fig. 1D), with seven irregular teeth terminally. Outer
basal area with a long and a shorter plumose seta.

Maxilla | bilobed (Fig. 1E), with broad basal segment. Inner lobe with four
apical teeth, the posterior lateral tooth longer and more robust. Dorsal surface
bearing four short and two large robust teeth. A plumose seta on inner, proximal
border. Outer lobe or palp with four lateral, non-plumose setae, the three distal
teeth borne on a slightly produced lobe. Apex of palp bearing a barber spine and
two slender setae.

Maxilla 2 (Fig. 2B), largest of oral appendages. First and second segment
bearing two plumose, and a single non-plumose seta respectively. Third segment
modified, expanded and produced into two stout serrated claws and a single
non-plumose seta Attenuated lacinia between second and third segments,
bearing a smooth spine in mid-region, and three more on distal half. Terminal
segment slightly wider than long, with two stout, serrated claws, two hair-like
setae and a smooth spine.

Maxilliped slender (Fig. 1F), basal segment with three attenuated spines
along inner lateral margin. Terminal segment less than half as long as basal, with
two attenuated spines on inner lateral margin. More proximal spine robust and
long. Distal margin bears a claw-like attenuated spine and two subterminal setae
the longer equal in length to the terminal spine and about two and a half times
as long as apical segment.

Swimming legs 1-4 biramous (Figs 2C—D, 3A), first basipodite segment
bearing seta on inner margin, the second with seta on outer margin. Distal
margin of second basipodite segment of first four pairs of legs with fine spinules.
Exopodite and endopodite three segmented, the second podomere shorter than
the first and third, more pronounced on exopodite. Outer lateral margins of
endopodite segments of legs 1-4 with fine, hair-like setae.

First pair of legs (Fig. 2C) smaller than succeeding pairs, the second basi-

A NEW SPECIES OF HALICYCLOPS FROM ESTUARIES IN TRANSKEI 365 N

== == =
us = Cl he

0.1mm

6 tLIL>>

Fig. 2. Female. A. Abdomen. B. Maxilla 2. C. Leg 1. D. Leg 2.

366 ANNALS OF THE SOUTH AFRICAN MUSEUM

podite segment with a stout, serrated spine on inner distal margin, reaching to
about the mid-point of the endopodite. Proximal region of spine bearing a
number of smooth setae, the number and relative lengths showing individual
variation. First exopodite segment with one serrated spine and one short plumose
seta. Second exopodite segment with one serrated spine and one long plumose
setae. First and second endopodite segments each with a single plumose seta.
Third segment with sharp, spine-like distal corners and bearing two serrate
spines and four plumose setae.

Second and third pairs of legs similar, the ornamentation as follows: first
and second exopodite segment each with a serrated spine and one plumose seta
(Fig. 2D). Apical segment with three lateral spines, one long terminal spine and
five plumose setae. Outer lateral border of first exopodite of second leg with fine
spinules. These spinules present on first and second exopodite segment of third
pair of legs. First endopodite segment with one plumose seta, second segment
with two plumose setae and apical segment with three plumose setae, two
terminal spines and one lateral spine. Exopodite and endopodite spines all
serrated. Terminal segment with sharp distal corners.

Fourth pair of legs similar to preceding two pairs, but showing differences
in ornamentation of the terminal segments and in the structure of the seta of
first endopodite segment (Fig. 3A). Exopodite three with five plumose setae,
two lateral spines and one terminal spine. Outer lateral border with minute
spinules. Terminal segment of endopodite with one lateral spine, two terminal |
spines and two inner plumose setae. Seta of first endopodite segment with about
eight long, slender hairs in proximal region.

Leg 5 (Fig. 3B) with first segment fused with thoracic segment, bearing a
long, smooth seta on a basal protuberance. Protuberance ornamented with a
row of fine spinules. Second segment about one and three-quarters as long as
wide. Inner lateral margin straight, with one subterminal, finely serrated spine.
Length of this spine distinctly less than length of second segment. Terminal seta
long and slender, about twice the length of inner spine. Outer margin with two
finely serrated spines, the more proximal always the longest and about equal in
length to the second segment. All spines relatively slender. Origin of terminal
seta almost equidistant between two subterminal spines. Second segment with a
number of rows of fine spinules, their relative positions illustrated in Figure 3B.
Male. General body form similar to female, but smaller in size (Fig. 3C). Total
length 0,49-0,53 mm (mean of eight specimens 0,51 mm). Urosome slender, five
segmented. Anal segment and caudal furcae as in female.

Antennule of about thirteen segments (Fig. 3D), prehensile. Segments
difficult to define. Structure and arrangement of setae illustrated in Figure 3D.
Remaining cephalic appendages and legs 1-4 as in female.

Leg 5 first segment and ornamentation similar to female (Fig. 3E). Second
segment about twice as long as wide, inner margin straight, with two slender
plumose setae in distal half. Setae slightly longer than second segment. Terminal
setae distinctly plumose and almost twice as long as second segment. Rounded

AB D-F

Fig. 3. A-B. Female. A. Leg 4. B. Leg 5. C—F. Male. C. 0,49 mm male.
Paves 5) bY Ees/6:

A NEW SPECIES OF HALICYCLOPS FROM ESTUARIES IN TRANSKEI 367

D. Antennule.

a

368 ANNALS OF THE SOUTH AFRICAN MUSEUM

outer lateral margin with two short, finely serrated, slender spines. Length of
spines about half the length of lateral setae on the inner border of segment.
Margin of segment with a number of fine spinules.

Leg 6 (Fig. 3F) with a plumose outer seta almost equal in length to terminal
seta of leg 5, borne on a produced lobe. Inner lobe broad, bearing a sharp
serrated spine and a short spine-like seta.

DISCUSSION

The genus Halicyclops Norman, 1903 is widely scattered geographically and
is known from forty-seven described species. A key to the species by Lindberg
(1957) and a paper by Wilson (1958) were valuable aids to workers and did much to
emphasize important diagnostic characteristics. At the time of Lindberg’s (1957)
publication, thirty-three species of the genus had been described. A further ten
species were noted by Kiefer (1967). To this list must be added Halicyclops
ryukyuensis Ito, 1962, H. latus Chia-Jui & Ai-Yun, 1964, H. ambiguus Kiefer,
1967, and Halicyclops longispinosus Monchenko, 1974.

The present species differs from those already described, but shows similari-
ties to H. neglectus Kiefer, 1935 (redescription: Kiefer 1936; Candeias 1964);
to H. rotundipes Kiefer, 1935 (redescription: Kiefer 1936; Plesa 1956; Petkovski
1955 as H. neglectus rotundipes); and to H. higoensis Ito, 1958. H. neglectus and
H. rotundipes are no doubt closely allied and Wilson (1958) suggests that the .
two forms may be synonymous.

Halicyclops pondoensis differs from H. neglectus, H. rotundipes and H. higo-
ensis particularly in regard to the fourth and fifth pair of legs. In H. pondoensis
the seta on the first endopodite segment of the fourth pair of legs in the male and
female bears hair-like structures in the proximal region. These hair-like structures
are absent in other species.

The inner proximal seta on the third endopodite segment of the fourth pair
of legs differs in H. higoensis. In H. pondoensis, H. neglectus and H. rotundipes
a typical seta is present. H. higoensis is distinguished in having this seta in the
form of a spine.

Important differences between the species are also found on the fifth pair
of legs. In H. neglectus and H. rotundipes the second segment in both the male
and female is rounded. In H. higoensis this segment is slightly elongated. In
H. pondoensis the second segment in the female is about one and three-quarters
as long as wide and about twice as long as wide in the male. The armature on
this segment in the male is also characteristic. In H. pondoensis there are two
spines and three setae. In H. neglectus, H. rotundipes and H. higoensis three
spines and two setae are present.

The armature of the sixth pair of legs in the male also shows important
differences between the species. In H. neglectus, H. rotundipes and H. pondoensis
the spine on leg six is relatively long and slender. In H. higoensis this spine is
stout. Two setae are also present on leg six and in H. higoensis, H. neglectus and

A NEW SPECIES OF HALICYCLOPS FROM ESTUARIES IN TRANSKEI 369

H. rotundipes the outermost seta is slightly longer than the inner seta. In
H. pondoensis the inner seta is short and spine-like.
Small differences between the species are also apparent in the form of the
genital segment. In H. neglectus and H. rotundipes the lateral protrusions are
well marked, while in H. pondoensis and H. higoensis these protrusions are small.
The main differences separating these four species are summarized in

Table 1.

H. pondoensis is the fourth member of the genus to be described from the
continent of Africa south of the Tropic of Cancer. A further four species are
known from the islands of Madagascar and Réunion. A key to these eight species
with their geographical distribution is given.

TABLE |

Summary of important differences separating H. pondoensis from three closely allied forms.

Seta on first endo-
podite segment of
leg four, male and
female

Inner proximal
seta on third
endopodite seg-
ment of leg four,
male and female

Second segment

of fifth pair of legs
in male and female

Armature of leg
five in male

Armature of leg
six in male

Genital segment

Total body length:
Male

Female

Halicyclops
neglectus

Typical seta
present

Typical seta
present

Rounded

Three spines and
two setae present

Spine long and
slender

Outermost seta
slightly longer
than inner seta

Lateral protru-
sions well marked

390-440 ppm

600 »m

Halicyclops
rotundipes

Typical seta
present

Typical seta
present

Rounded

Three spines and
two setae present

Spine long and
slender

Outermost seta
slightly longer
than inner seta

Lateral protru-
sions well marked

460 »m

500-560 pm

Halicyclops
higoensis

Typical seta
present

Spine present

Slightly longer
than round

Three spines and
two setae present

Spine short and
stout

Outermost seta
slightly longer
than inner seta

Lateral protru-
sions small

460 pm

520 pm

Halicyclops
pondoensis

Seta with hair-like
structures in proxi-
mal region

Typical seta
present

One and three-
quarters as long as
wide in female and
twice as long as
wide in male

Two spines and
three setae present

Spine long and
slender

Outermost seta
much longer than
inner seta which is
spine-like

Lateral protru-
sions small

490-530 pm

610-680 pm

Mr?

<FILILSS

370 ANNALS OF THE SOUTH AFRICAN MUSEUM

KEY TO THE SPECIES OF HALICYCLOPS FROM THE CONTINENT OF AFRICA
SOUTH OF THE TROPIC OF CANCER AND FROM THE ISLANDS OF
MADAGASCAR AND REUNION WITH A NOTE ON THEIR DISTRIBUTION

1. Inner seta absent on endopod segment 3 of leg 4 H. korodiensis Onabamiro, 1952
Distribution: Nigeria.

— Inner seta present on endopod segment 3 of leg 4 x Pe

2. Spines along posterior dorsal edge of penultimate abdominal ‘care longer in the
middle region than on the sides... : : 3

— Spines along posterior dorsal edge of acu ulenaic Aptana wenn equa in een 6

3. Spine formula on terminal exopod segments of legs 1-4, 2.3.3.3. Furcal rami about as
long as broad . ~ i .. H. pusillus Kiefer, 1954
Interstitial, Menoantsena room Madasasear.

— Spine formula on terminal exopod segments of 1-4, 3.4.4.3. Furcal rami longer

than broad _... Be E bbe se ie a
4. Chitinous spine present on cach ane of Benita cats H. thermophilus Kiefer, 1929
In a limestone cave, Madagascar (Lindberg 1952).
— No chitinous spine present on each side of genital segment .... 5

5. Furcal rami at least 2 times as long as broad. Spines along the posterior dara oe
of penultimate abdominal segment extremely long in the middle region, reaching beyond
midlength of anal segment .. Si? ke H. denticulatus Kiefer, 1960
Brackish water near Manambato, Madagascar.

— Furcal rami about 1,5 times as long as broad. Spines along the posterior dorsal edge
of penultimate abdominal segment slightly longer in the middle region than on the

sides, not reaching beyond midlength of anal segment .. H. reunionis Kiefer, 1960
Island of Réunion.

6. Genital segment broader than long + Si .. A. gauldii Plesa, 1961
Interstitial: Lighthouse Beach, Accra, Ghana.
Genital segment about as long as broad .. an eee)

7. Genital segment with small lateral protrusions. ace rami coat as ie as ea
Hair-like structures on distal part of seta of Ist endopodite segment of leg 4. Three
spines and a single seta on second segment on leg 5 of female. Two spines and three
setae present on leg 5 in male bp a = .. H. pondoensis sp. nov.
Pondoland estuaries, Transkei. Planktonic.

— Genital segment without lateral protrusions, or at most, a weak projection present.
Furcal rami about 1,5 times as long as broad. No hair-like structures on seta of Ist
endopodite segment of leg 4. Three spines and a single seta on second segment of
leg 5 of female. Male undescribed .. ee - H. orae eburnensis Lindberg, 1957
Ebrie lagoon, Ivory Coast.

ACKNOWLEDGEMENTS

I am indebted to the South African National Council for Oceanographic
Research and to the Department of Planning and the Environment who funded
this project and to Dr J. R. Grindley under whose directorship this work was
initiated. I thank Dr J. P. Furstenberg who critically read drafts of this
manuscript.

REFERENCES

CANDEIAS, A. 1964. Contribution to the knowledge of the development of Halicyclops neglectus
Kiefer. Revta Biol., Lisb. 4: 171-188.

Cuta-Jur, S. & Al-YuN, T. 1964. Descriptions of new species of fresh-water copepods from
Kwangtung Province, South China. Acta Zootaxon. sin. 1: 367-396.

A NEW SPECIES OF HALICYCLOPS FROM ESTUARIES IN TRANSKEI 37]

Hitt, B. J. 1966. A contribution to the ecology of the Umlalazi estuary. Zool. afr. 2: 1-24.

Ito, T. 1958. Groundwater copepods from south-western Japan. Hydrobiologia 11: 1-28.

Ito, T. 1962. Groundwater copepods from the Ryu-Kyu Island. Jap. J. Zool. 13: 275-292.

KieFerR, F. 1929. Neue Ruderfusskrebse von den Sunda-Inseln. (1. Mitteilung tiber die Cope-
poden der Sunda-Expedition Rensch-Hebener.) Zool. Anz. 84: 46-49.

KIEFER, F. 1935. Zur Kenntnis der Halicyclopen (Crustacea Copepoda). Zoo/. Anz. 110: 10-13.

KIEFER, F. 1936. Freilebende Stiss- und Salzwassercopepoden von der Inseln Haiti. Mit einer
Revision der Gattung Halicyclops Norman. Arch. Hydrobiol. 30: 263-317.

Krererr, F. 1954. Neue Cyclopoida Gnathostoma (Crust. Cop.) aus Madagascar 1. Cyclopiniae
und Halicyclopinae. Zool. Anz. 153: 308-313.

KIEFER, F. 1960. Neue Cyclopoida Gnathostoma (Crust. Cop.) von den Inseln Madagascar
und Réunion. Zool. Anz. 165: 226-232.

KIEFER, F. 1967. Cyclopiden aus salzhaltigen Binnengewdssern Australiens (Copepoda).
Crustaceana 12: 292-302.

LINDBERG, K. 1952. Cyclopides (Crustacés copepodes) de Madagascar Troisieme note. Mém.
Inst. scient. Madagascar 7A: 53-67.

LINDBERG, K. 1957. Cyclopides (Crustacés copépodes) de la Céte d’Ivoire. Bull. Inst. fr. Afr.
noire (Sér. A) 19: 134-179.

MONCHENKO, V. I. 1974. A new Halicyclops (Crustacea, Copepoda) species from the Dnieper—
Bug estuary. Vestn. zool. 3: 47-52. (In Russian with Russian and English summaries.)

Norman, A. M. 1903. New generic names for some Entomostraca and Cirripedia. Ann. Mag.
nat. Hist. (Series 7) 11: 367-369.

ONABAMIRO, S. D. 1952. Four new species of Cyclops sensu lat. (Crustacea: Copepoda)
from Nigeria. Proc. zool. Soc. Lond. 122: 253-266.

PETROVSKI, T. K. 1955. Beitrag. zur Kenntnis der Copepoden. Acta Mus. maced. Sci. nat. 3:
71-104.

PLesaA, C. 1956. Quelques remarques sur les Halicyclopes (Cruaracés copépodes) des eaux
roumaines. Folia balc. 1: 25-28.

PLesA, C. 1961. New Cyclopoides (Crustacea, Copepoda) of the Interstitial Fauna from the
Beaches of Ghana. J/] W. Afr. Sci. Ass. 7: 1-13.

Witson, M. S. 1958. The Copepod genus Halicyclops in North America with description of a
new species from Lake Pontchartrain, Louisiana, and the Texas coast. Tulane Stud. Zool.
6: 176-189.

Ooi ee eM is AT, Sl LT a) NN SS Se NY RR es A a A Rt ES rt Am AU

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

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

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

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

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

Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.

Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (figs 8a—b).
Nucula largillierti Philippi, 1861: 87.

Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

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

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

In describing new species, one specimen must be designated as the holotype; other speci-
mens mentioned in the original description are to be designated paratypes; additional material
not regarded as paratypes should be listed separately. The complete data (registration number,
depository, description of specimen, locality, collector, date) of the holotype and paratypes
must be recorded, e.g.:

Holotype
SAM-A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach,
Port Elizabeth (33°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

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

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

(c) Scientific names, but not their vernacular derivatives
e.g. Therocephalia, but therocephalian

Punctuation should be loose, omitting all not strictly necessary

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Roman numerals should be converted to arabic, except when forming part of the title of a
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‘Revision of the Crustacea. Part VIII. The Amphipoda.’ Rea

Specific name must not stand alone, but be preceded by the generic name or its abbreviation
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Name of new genus or species is not to be included in the title: it should be included in the
abstract, counter to Recommendation 23 of the Code, to meet the requirements of
Biological Abstracts.

T. WOOLDRIDGE

A NEW SPECIES OF HALICYCLOPS
(COPEPODA, CYCLOPOIDA)
FROM ESTUARIES IN TRANSKEI, SOUTHERN AFRICA

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