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

(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, doubie spaced

with 2,5 cm margins all round. First lines of paragraphs should be indented. Tables and a list of

legends for illustrations should be typed separately, their positions indicated in the text. All

pages should be numbered consecutively.

Major headings of the paper are centred capitals; first subheadings are shouldered small

capitals; second subheadings are shouldered italics; third subheadings are indented, shouldered

italics. Further subdivisions should be avoided, as also enumeration (never roman numerals)

of headings and abbreviations.

Footnotes should be avoided unless they are short and essential.

Only generic and specific names should be underlined to indicate italics; all other marking

up should be left to editor and publisher.

4. ILLUSTRATIONS should be reducible to a size not exceeding 12 « 18 cm (19 cm including

legend); the reduction or enlargement required should be indicated; originals larger than

35 x 47 cm should not be submitted; photographs should be rectangular in shape and final

size. A metric scale should appear with all illustrations, otherwise magnification or reduction

should be given in the legend; if the latter, then the final reduction or enlargement should be

taken into consideration.

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 year of publication given in text, e.g.:

‘Smith (1969) describes .. .’

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

“As described (Smith 1969a, 19695; Jones 1971)’

‘As described (Haughton & Broom 1927)...’

‘As described (Haughton et a/. 1927)...’

Note: no comma separating name and year

Dagination indicated by colon, not p.

names of joint authors connected by ampersand

et al. in text for more than two joint authors, but names of all authors given in list of references.

(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. gén. 74: 627-634.

Konun, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.

Ann. Mag. nat. Hist. (13) 2: 309-320.

Konn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.

Bull. Bingham oceanogr. Coll. 17 (4): 1-51.

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 77 Band

February 1979 Februarie

Part 6 Deel

CRETACEOUS FAUNAS FROM ZULULAND

AND NATAL, SOUTH AFRICA

THE AMMONITE SUPERFAMILY

HAPLOCERATACEAE ZITTEL, 1884

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

Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

word uitgegee in dele op ongereelde tye na beskikbaarheid

van stof

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

OUT OF PRINT/UIT DRUK

1, 2(1-3, 5-8), 3(1-2, 4-5, 8, t.-p.i.), 5(1-3, 5, 7-9),

6(1, t.—p.i.), 71-4), 8, 9(1-2, 7), 10(1-3),

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

Kopieregnavrae na die Suid-Afrikaanse Museum

ISBN 0 908407 63 7

Printed in South Africa by In Suid-Afrika gedruk deur

The Rustica Press, Pty., Ltd., Die Rustica-pers, Edms., Bpk.,

Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

CRETACEOUS FAUNAS FROM ZULULAND AND NATAL,

SOUTH AFRICA

THE AMMONITE SUPERFAMILY HAPLOCERATACEAE ZITTEL, 1884

WILLIAM JAMES KENNEDY

Geological Collections, University Museum, Oxford

HERBERT CHRISTIAN KLINGER

South African Museum, Cape Town

(With 20 figures)

LMS. accepted 18 October 1978]

ABSTRACT

Representatives of the superfamily Haplocerataceae Zittel, 1884, are locally common in

rocks of Barremian age in northern Zululand, where Sanmartinoceras (Sanmartinoceras)

africanum sp. nov. occurs at all stages of development from larval shells to discs which are still

septate at a diameter of 120 mm, and represent the largest known specimens of the genus.

Sanmartinoceras (Theganeceras) nodosum sp. nov., of Lower Aptian age, Sanmartinoceras

(Sinzovia) trautscholdi (Sinzow), and an Aconeceras sp. of Upper Aptian age are other repre-

sentatives of the Aconeceratidae, whilst the Binneyitidae are represented by specimens of

Borissiakoceras of Middle Cenomanian age. In addition to systematic descriptions, the

problems of recognizing dimorphism in these genera are discussed, and certain resultant

taxonomic problems noted, and an annotated list of species referred to the two families is

included.

CONTENTS

PAGE

Introduction . : ee 3 3 o>

Location of specimens : , : . 86

Field localities. : : 2 ; 86

Dimensions of specimens . ; : . 86

Suture terminology. . - F » 86

Systematic palaeontology . : , Si

Annotated list of species referred to the

Aconeceratidae and Binneyitidae . 5 lia

Acknowledgements. ; : , a alls

References. ; ; ; ; ‘ . 120

INTRODUCTION

The Haplocerataceae are the longest ranging of the Ammonitina, first

appearing in the Middle Jurassic (Bajocian) and ranging to the Upper Cretaceous

(Coniacian). As Casey has noted, two broad morphologies recur within the

superfamily: firstly, smooth platycones, typified by the Haploceratidae and

Binneyitidae, and secondly, variously ribbed or strigate oxycones with entire or

Ann. S. Afr. Mus. 77 (6), 1979: 85-121, 20 figs.

86 ANNALS OF THE SOUTH AFRICAN MUSEUM

crenulate keels, typified by the Oppeliinae, Strigoceratidae and Aconeceratidae

(Casey 1961a: 118). In the Cretaceous of Zululand both groups are represented,

in the Upper Barremian and Aptian the Aconeceratidae are not uncommon at

some levels, whilst the Binneyitidae occur as a great rarity in rocks 6f Middle

Cenomanian age. Although only a few species are represented in the authors’

collections, they are of some significance, for their specimens of Aconeceratidae

allow, for the first time, the description of the detailed ontogeny of the widely

occurring genus Sanmartinoceras Bonarelli, 1921, as well as clarifying some

aspects of its stratigraphic distribution, whilst the present records of the

binneyitid Borissiakoceras extend the known distribution of the group.

The following species are described below: Aconeceras sp., compared to

A. walshense (Etheridge); Sanmartinoceras (Sanmartinoceras) africanum sp. nov.;

Sanmartinoceras (Theganeceras) nodosum sp. nov.; Sanmartinoceras (Sinzovia)

trautscholdi (Sinzow); and Borissiakoceras sp.

To this is added an annotated list of genera, subgenera and species referred

to the two families.

LOCATION OF SPECIMENS

The following abbreviations are used to indicate the repositories of the

material studied:

BMNH British Museum (Natural History), London

GSM Institute of Geological Sciences, London

MNHP Muséum d’Histoire Naturelle, Paris

OUM = Oxford University Museum, Oxford

SAM South African Museum, Cape Town

SAS South African Geological Survey, Pretoria

FIELD LOCALITIES

Details of localities mentioned in the text are given by Kennedy & Klinger

(1975); fuller descriptions of sections are deposited in the Palaeontology

Department of the British Museum (Natural History), London; Geological

Survey, Pretoria; and South African Museum, Cape Town.

DIMENSIONS OF SPECIMENS

All dimensions given below are in millimetres:

D = diameter, Wb = whorl breadth, Wh = whorl height, U = umbilical

diameter.

Figures in parentheses are dimensions as a percentage of the total diameter.

SUTURE TERMINOLOGY

The suture terminology of Wedekind (1916), recently reviewed and dis-

cussed by Kullman & Wiedmann (1970) is followed here:

Is = Internal lobe with septal lobe, U = Umbilical lobe, L = Lateral lobe,

E = External lobe.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 87

SYSTEMATIC PALAEONTOLOGY

Phylum MOLLUSCA

Class CEPHALOPODA Cuvier, 1797

Subclass AMMONOIDEA Zittel, 1884

Order AMMONITIDA Zittel, 1884

Suborder AMMONITINA Zittel, 1889

Superfamily HAPLOCERATACEAE Zittel, 1884

Family Aconeceratidae Spath, 1923

Discussion

The Aconeceratidae Spath, 1923, is a group of small to medium-sized,

compressed, involute, commonly oxycone ammonites with flat or convex sides

and a fastigiate to keeled venter. Ornament consists of weak to strong flexuous,

commonly biconvex ribs and growth striae and a keel which may be minutely

crenulate.

The following genera and subgenera have been placed in the family:

Genus Protaconeceras Casey, 1954, Hauterivian

Genus Aconeceras Hyatt, 1903, Barremian to Lower Albian

Genus Sanmartinoceras Bonarelli, 1921 (in Bonarelli & Nagera 1921),

Barremian to Lower Albian

Subgenus Sanmartinoceras s.s., Barremian to Aptian

Subgenus Sinzovia Sazonova, 1958, Aptian

Subgenus Theganeceras Whitehouse, 1926, Lower Aptian

Genus Doridiscus Casey, 1961, Aptian

Genus Nothodiscus Casey in Collignon, 1962, Aptian

Genus Gyaloceras Whitehouse, 1927, Upper Aptian.

Aconeceratids have a wide geographic distribution extending from

Greenland to Antarctica, although their occurrence is sporadic. In Europe at

least, they occur in enormous numbers in some clay facies as in southern

France, where the blue marls of the Fosse Vocontienne yield thousands of

Aconeceras and Casey (1961b: 122) regarded them as an open sea group which

invaded the neritic zone only in periods favourable for their growth. As will be

seen below, however, their local abundance in carbonaceous nearshore sand-

stones of the South African Barremian is scarcely compatible with such a

generalization.

The Haplocerataceae as a whole are characterized by striking dimorphism

(Makowski 1962, Callomon 1963, Kennedy & Cobban 1976 with references),

but within the Aconeceratidae our knowledge is such that it remains most

difficult to recognize dimorphic pairs. In Aconeceras itself, the many hundreds

of specimens from the French Aptian examined are all pyritic phragmocones

and it would appear that juveniles, at least, are most difficult to split into

dimorphs, as Palframan (1969) found in some Jurassic haploceratids. The

authors do know, however, of a few specimens referrable to the family in which

88 ANNALS OF THE SOUTH AFRICAN MUSEUM

apertures are preserved: the original material of S. (Sanmartinoceras)

groenlandicum Rosenkrantz, 1934 (in Bogvad & Rosenkrantz), illustrated here

as Figure 1A-C, is an obvious microconch with prominent lappets and a

rostrum. Equally, Gyaloceras smithi Whitehouse, 1927, appears to be a female

or macroconch. The specimen is reproduced here as Figure 2A. In most cases,

however, the shell ornament of aconeceratids (as opposed to apertural form)

gives little clue to dimorphism and a markedly biconvex growth line, indicating

a long rostrum and blunt lappets, occurs, in the present material at least, in both

small and large (if not mature specimens). The authors would, however, suggest

that the presence of a strong spiral depression may indicate the possible presence

of a long lappet (Figs 1A—B, 5C). The topic is returned to below under the

discussion of Sanmartinoceras.

Genus Aconeceras Hyatt, 1903

Type species

Ammonites nisus d’Orbigny, 1841.

Discussion

Aconeceras is represented by a single, poor specimen from the Aptian of

Zululand ; in consequence the reader is referred to Casey’s (19615: 123) extensive

remarks on the genus.

Occurrence

The genus ranges from Barremian to Lower Albian and is widely distributed

from western Europe and the U.S.S.R. to east Africa, Madagascar, Nepal,

Antarctica and eastern Australia.

Aconeceras sp.

Material

One specimen only, BMNH C80002, from the Makatini Formation,

Aptian III, locality 166, Mfongosi Spruit, northern Zululand.

Description

The specimen is an external mould of an oxycone individual with an

original maximum whorl height of over 20 mm. Coiling is very involute with a

small, pit-like steep-sided umbilicus. The whorl section is high, compressed with

the greatest breadth well below mid-fiank. The flanks are distinctly flattened and

the venter fastigiate.

The specimen is somewhat worn and there is no obvious trace of ornament

preserved.

Discussion

Although poorly preserved, this specimen is clearly referrable to the genus

Aconeceras. Amongst described species it is perhaps to be compared with the

CRETACEOUS FAUNAS FROM SOUTH AFRICA

‘TX = “(HEST) ZlURIUSSOY 32 peaseg Jo AdoDd “pest ‘zIUeIUASOY WINIIPUDIUIOAS (SDABIOUILADULUDG)) SDaIOUIJ ADDS)

[ ‘3hy

90 ANNALS OF THE SOUTH AFRICAN MUSEUM

pbs

mots

Fig. 2. A. Gyaloceras smithi Whitehouse, 1926. B—D. Aconeceras walshense (Etheridge), 1892.

After Whitehouse (1926, 1927a). x1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 91

Fig. 3. A-C. Sanmartinoceras (Sanmartinoceras) fontinale (Hudleston), 1890.

D-G. Sanmartinoceras (Sanmartinoceras) olenae (Tenison-Woods), 1883. After Whitehouse

(1927a). x1.

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 4. Sanmartinoceras (Sanmartinoceras) africanum sp. nov., paratypes.

A. SAS H54/41/e. B-D. BMNH C79988. E-H. BMNH C79982. x1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 93

Australian Aconeceras walshense (Etheridge) (see Whitehouse 1927a: 114, pl. 16

(figs 2-3); text-figs 1, 6-7) (Fig. 2B—D herein).

Occurrence

Upper Aptian of Zululand.

Genus Sanmartinoceras Bonarelli, 1921 (in Bonarelli & Nagera 1921)

Type species

Sanmartinoceras patagonicum Bonarelli, 1921 (in Bonarelli & Nagera 1921).

Diagnosis

Stout oxycones, initially smooth, with falciform striae and ribs, the latter

weak on the inner flank, sometimes bunched, and strengthening into distinctive

concave ribs on the outer flank. Keel finely denticulate.

Discussion

Sanmartinoceras was originally based upon crushed specimens from the

Aptian of Lago San Martin, Argentina. Leanza (1970, text-fig. 14) (see Fig. SA-B

herein) has figured uncrushed topotypes, whilst Howarth (1958) and Thomson

(1974) have figured other material from the sub-Antarctic Islands. Specimens

are illustrated here as Figure SC—D. Specimens of S. patagonicum with apertures

preserved are unknown, but Rosenkrantz (in Bogvad & Rosenkrantz 1934: 20,

pl. 4 (fig. 3), pl. 5 (figs 1-5)) figured a species, S. groenlandicum Rosenkrantz,

with a distinctive rostrum and lappets. These features, indicative of a micro-

conch, have been incorporated into the generic diagnosis. Examination of these

examples shows that the lateral lappets are associated with a distinct spiral

groove (Fig. 1A—B). In the Antarctic specimens (Fig. 5C—D) similarly sized

individuals show this groove developed to varying degrees at the same diameter,

and the authors suspect it may prove a criterion for differentiation of immature

males and females at similar diameters, in the type species at least. In the case

of Australian Sanmartinoceras the problem is even more tantalizing. Whitehouse

(1926; 1927a) described four aconeceratids from the Aptian of Walsh River.

Two, Sanmartinoceras fontinale (Hudleston) (Fig. 3D-G, 7A-H) and S. olenae

(Tenison-Woods) (Fig. 3A—C) have strong falcoid ribs and prominent rostra.

The others are very feebly ornamented. Aconeceras walshense (Etheridge)

(Fig. 2B—D) has falcoid growth striae, whilst Gyaloceras smithi Whitehouse

(Fig. 2A) has an inflated whorl and fastigiate venter. 1he authors strongly

suspect that these species are dimorphs but again cannot resolve the problem

fully.

Within Sanmartinoceras, three subgenera have been recognized. In addition

to Sanmartinoceras sensu stricto, Theganeceras Whitehouse, 1926, was treated

as a subgenus by both Wright (1957) and Casey (19616). Only three species,

S. (T.) falcatum (von Koenen) (Fig. 6I-J) S. (T.) scalatum (von Koenen)

(Fig. 6H) from the Lower Aptian of northern Germany, and S. (T.) grande

94 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 5. Sanmartinoceras (Sanmartinoceras) paragonicum Bonarelli, 1921.

A-B. Topotype, after Leanza (1970). C. BMNH C49055. D. BMNH C49057.

C_D. from Alexander Land. x1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 95

Fig. 6. A. Sanmartinoceras (Sinzovia) aff. trautscholdi (Sinzow), 1870. B-—D. Sanmartinoceras

(Sinzovia) stolleyi Casey, 1961. BMNH C14365 and 14362, from the Upper Aptian of Bekum,

near Hildesheim, Germany. E-F. Sanmartinoceras (Sinzovia) trautscholdi (Sinzow), 1870.

Copy of Trautschold (1865). G. Sanmartinoceras (Theganeceras) grande Thomson, 1974.

After Thomson (1974). H. Sanmartinoceras (Theganeceras) scalatum von Koenen, 1902.

I-J. Sanmartinoceras (Theganeceras) falcatum von Koenen, 1902. H-J after von Koenen

(1902). All x1.

96 ANNALS OF THE SOUTH AFRICAN MUSEUM

Thomson (Fig. 6G) from the Aptian of Antarctica, have thus far been ascribed

to the subgenus, which Casey (19616: 132) diagnosed as lacking the smooth

juvenile stage of Sanmartinoceras sensu stricto and having finer, more numerous

dense ribs. As is described below, the single Zululand specimen referred to the

subgenus suggests that dimorphism may also present taxonomic problems.

The subgenus Sinzovia Sazonova, 1958, differs from Sanmartinoceras sensu

stricto (according to Casey 19616: 133) in having a very low keel and absence or

poor development of a spiral groove. It differs from Theganeceras in having a

longer juvenile smooth stage and greater tendency to smoothness on the lower

flank. The type species is illustrated here as Figure 6E-F, related German species

as Figure 6A—D, and the only South African representative, S. (S.) trautscholdi

(Sinzow), as Figure 7I-K.

Occurrence

Sanmartinoceras and its subgenera range from Upper Barremian to Aptian

and are known from Greenland, western Europe, the U.S.S.R., Zululand,

Madagascar, Nepal, Australia, Antarctica, Argentina and Papua.

Subgenus Sanmartinoceras sensu stricto

Sanmartinoceras (Sanmartinoceras) africanum sp. nov.

Figs 4A-H; 8A-F; 9A-I; 10A-F; 11A-B; 12A-C; 13A-F; 14A-C; 15F-J;

16-17; 19D-E

Sanmartinoceras Kennedy & Klinger 197: 274.

Types

The holotype is SAS H54; paratypes are BMNH C79977—C80001, SAS

H54/4la-d, SAS ZO(i), SAS LJE13la, SAS LJE112, SAS H54/17, SAS

H54/33, all from the Makatini Formation, Barremian I-II, locality 170,

Mlambongwenya Spruit, northern Zululand.

Diagnosis

A large (up to 120 mm diameter phragmocone) species of Sanmartinoceras

in which the inner ‘haft’ of the ribs is narrow, straight and prorsiradiate and the

outer ‘blade’ broad and markedly concave.

Dimensions

D Wb Wh Wb:Wh U

Holotype, SAS H54 —(—) 20,5(—) 42-5(—) 0,48 —(—)

C79985 aye J 27,3(100) 6,6(24) 14,8(54) 0,45 4,1(15)

C79978 ss .. 120,0(100) 28,8(24) 69,8(58) 0,41 730)

C79984 .. —.. —-115,0(100) 24,0(21) 65,5(57) 0.37. ~—«:11,3(9,8)

CRETACEOUS FAUNAS FROM SOUTH AFRICA 97

Fig. 7. Sanmartinoceras (Sanmartinoceras) fontinale (Hudleston), 1890. Holotype, BMNH

C5306, Upper Aptian, Primrose Springs, north of Lake Eyre, south Australia. A-D x1;

E-H x2. I-K. Sanmartinoceras (Sinzovia) trautscholdi (Sinzow, 1870). SAM-—PCZ5919. x1.

98 ANNALS OF THE SOUTH AFRICAN MUSEUM

D E F

Fig. 8. Sanmartinoceras (Sanmartinoceras) africanum sp. nov.

A-C. Holotype SAS 54; D-F paratype BMNH C79989. x1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA

Fig. 9. Sanmartinoceras (Sanmartinoceras) africanum sp. nov., paratypes.

A-D. SAS H54/416. E-H. SAS H54/41a. I. BMNH C79996. All x2.

100 ANNALS OF THE SOUTH AFRICAN MUSEUM

Cc D E F

Fig. 10. Sanmartinoceras (Sanmartinoceras) africanum sp. nov., paratypes.

A-B. BMNH C79984. C-F. SAS H54/41c. A-B x1; C-F x2.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 101

Fig. 11. Sanmartinoceras (Sanmartinoceras) africanum sp. nov., paratypes.

A. BMNH C79984. B. SAS H54/41d. A x1; B x2.

102 ANNALS OF THE SOUTH AFRICAN MUSEUM

Description

The barrel-shaped protoconch is succeeded briefly by rather stout whorls,

beyond which, up to a diameter of 0,5-0,8 mm, the shell is moderately involute,

slightly compressed, with flattened flanks and a rounded venter. All the available

specimens are partially or wholly exfoliated, and any ornament, if present, must

have been weak. A few specimens begin to show traces of a ventral ridge, but

not a keel at this diameter, although these features may be an artefact of

preservation.

From 0,8 mm onwards the shell is an oxycone, the coiling is very involute,

with a tiny, crater-like umbilicus. The umbilical wall is flat and subvertical, with

an abruptly rounded shoulder. The whorl section varies markedly; stout

individuals have a whorl breadth to height ratio of 0,6; in slender individuals the

figure is 0,4. Maximum whorl breadth is low on the flanks; the inner flanks are

flattened, the outer flanks convergent; a distinct ventrolateral shoulder is

developed, and there is a sharp keel which, when well preserved, can be seen to

be minutely crenulated (Fig. 9E-H).

Ornament varies greatly. In some individuals (Figs 11B, 13C—D) there are

only the finest growth striae. These arise at the umbilical seam, but are very

weak. They pass forwards across the inner flank, strengthening as they do so,

and sometimes splitting (Fig. 15G—J). They are at their maximum development

at mid-flank, where they flex gently backwards, giving rise to a clear concavity

which extends across the outer third of the flank (Fig. 13D). Striae decline on

the ventrolateral shoulder and project strongly forwards to meet the keel at an

acute angle. Individual striae correspond to individual crenulations on the keel.

In robust specimens, e.g. BMNH C79983, striae are accompanied by distant,

pronounced ribs, which are strongest at mid-flank; there is every intermediate

between.

Middle growth stages are also characterized by a wide range of variation

(Figs 4E-H, 8A-F). The holotype demonstrates the features of a strongly

ornamented individual. It is wholly septate (Fig. 8B) retaining traces of shell.

Coiling is very involute, with most of the inner whorls being covered. The tiny

umbilicus is deep, with a high, flat, subvertical wall, abruptly rounded shoulder,

and faint umbilical carina. The whorl section is compressed (breadth to height

ratio is 0,48) with faintly concave inner flanks, a weakly inflated mid-flank

region converging to distinct shoulders, clearly demarcated from a high,

septate ventral keel. In none of the medium-sized specimens does the keel bear

serrations. Ornament consists of fine, rectiradiate striae on the inner part of the

flank, effaced at a spiral mid-flank depression. The outer flank bears low, blunt,

concave ribs, declining at the ventrolateral shoulder, but projected forwards to

meet the keel.

The largest available specimens are up to 120 mm in diameter, and are still

wholly septate (Fig. 12A—C). In these, the whorls are quite robust, and ornament

consists of striae on the inner flank with blunt, concave ribs on the outer flank.

None of the present specimens has the aperture preserved. A few fragments show

103

CRETACEOUS FAUNAS FROM SOUTH AFRICA

‘IX ‘8L66L0 HNWa odAjered “aou ‘ds wnuvoisfo (spsav0ulADULUDG) SDADIOUIZADUUDS) “Z| “BIT

104 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 13. Sanmartinoceras (Sanmartinoceras) africanum sp. Nov., paratypes.

A-B, F. SAS Zo(i). C. SAS LJE 131a. D. SAS H54/41a. E. BMNH C79979.

A-D, E-F x1; C-D x2.

105

CRETACEOUS FAUNAS FROM SOUTH AFRICA

‘TX A-d ‘1X O-V ‘oT ‘ou

WOT}OB]JOD Jedoog *yY ‘WP “ds spsavoyviss1iog *J-C ‘TIT ALT SVS edAyered “Aou ‘ds wnupa14fo (spsad0uljADUUDG) SDABIOUIJADUUUDS' *D-Y “pT “BIT

106 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 15. A-E. Borissiakoceras sp., BMNH C80003. F-J. Sanmartinoceras (Sanmartinoceras)

africanum sp. nov., paratypes. F is SAS LJE 112; G—J is BMNH C79985.

A-E, G-J x2; F x1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 107

weak to strong strigations (Figs 4B—D, 13E). Where well preserved, this appears

to be a feature of the dorsal shell layer, and not of the original external shell

surface. That this occurs in some very large specimens suggests the species may

have reached diameters close to 200 mm.

The available specimens show the details of suture development to

advantage, as shown in Figure 16.

Mature sutures are deeply and intricately subdivided, as shown in Figure 17.

Discussion

The specimens show the ontogeny and range of intraspecific variation far

better than in any previously described Sanmartinoceras whilst the specimens

are the largest known for the genus. It is most unfortunate that none of the

specimens retains apertures, but the rather distinct spiral flank depression

present in the holotype suggests that it may be a microconch; the largest

individuals (Figs 1OA—B, 11A, 12A—D) are probably macroconchs.

Thomson (1974: 23) reviewed criteria used to differentiate species of

Sanmartinoceras as follows:

(i) strength and form of ribbing,

(ii) the way in which ribs appear (i.e. abruptly or increasing gradually in

strength),

(iii) the size of the individual at which ribs first appear.

In his discussion, however, he concluded that the form of the falcate ribbing

alone was sufficient to separate species. Figure 18 compares the line of ribs and

striae in the five described species; on this criterion alone it can be seen that the

straight ‘haft’ and form of ‘sickle’ of the present species are quite different from

the markedly biconcave rib of S. olenae, S. fontinale and S. patagonicum.

A straight ‘haft’ characterizes S. groenlandicum (Fig. 18), but here the ‘blade’

is much more deeply concave with the inner half almost straight. As noted

elsewhere, the authors do not regard the presence or absence of a lateral spiral

groove as of specific significance, whilst the present specimens vary greatly in

strength of ornament and growth stage at which it appears. The other feature

which separates S. africanum from other species is its great size, but this may

be no more than an artefact of preservation.

Occurrence

Upper Barremian of Zululand only.

Subgenus Theganeceras Whitehouse, 1927

Theganoceras nodosum sp. nov.

Figs 19A-C, 20

Holotype

SAM-PCZ5708 from the Makatini Formation, Aptian I, locality 170,

Mlambongwenya Spruit, northern Zululand.

108 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 16. Suture development in Sanmartinoceras (Sanmartinoceras) africanum sp. nov.

A-C. BMNH C79979. D-E. SAS H54/41A. All x 12,5.

109

CRETACEOUS FAUNAS FROM SOUTH AFRICA

“STIX “OTb/PSH SVS ‘Aou ‘ds wnupoisfo (SDAZIOUIJADUUDG) SDABIOUIJADUIUDG JO SOINNS Ine “LT “Si

110 ANNALS OF THE SOUTH AFRICAN MUSEUM

geen

A B

Vig

Fig. 18. Rib patterns in Sanmartinoceras (Sanmartinoceras) compared.

A. S. (S.) olenae. B-C. S. (S.) fontinale. D. S. (S.) africanum. E. S. (S.) groenlandicum.

F-G. S. (S.) patagonicum.

Diagnosis

A large species of Theganeceras with broad, flat, falcoid ribs on the inner

whorls which bear umbilical bullae and incipient ventral clavi. Outer whorls

ornamented by bullae, flexuous growth lines, folds and constrictions.

Description

The unique holotype of this species comprises a crushed body chamber and

a nucleus. Growth of calcite between shell and sedimentary infilling has

produced an unduly thick ‘shell’ of partly diagenetic origin.

The inner whorl, at a diameter of approximately 38 mm, bears fourteen to

sixteen closely spaced, low, broad, sickle-shaped ribs which are straight and

prorsiradiate on the inner flank, flexing backwards at mid-flank and becoming

markedly concave on the outer flank. The ribs are strong and well developed

down to the umbilicus where a distinct bulla is present, whilst on the ventro-

lateral shoulder they strengthen into an incipient clavus. The ribs themselves

and interspaces between bear dense striae, sometimes strengthened into riblets,

giving some ribs a bunched, fasciculate appearance. The venter is fastigiate, and

bears strongly projected striae.

CRETACEOUS FAUNAS FROM SOUTH AFRICA ay

Fig. 19. A-C. Sanmartinoceras (Theganeceras) nodosum sp. nov., holotype, SAM-PCZ5708. Sle

D-E. S. (Sanmartinoceras) africanum sp. nov., patatype, BMNH specimen. x1.

112 ANNALS OF THE SOUTH AFRICAN MUSEUM

The outer body whorl shows coiling to have been involute with a moderately

large, deep, crater-like umbilicus with a flat, outwards-sloping wall. The whorl

section is compressed, lanceolate, so that the overall shell form was an oxycone.

The whorl breadth to height ratio is estimated as approximately 0,38, the

greatest breadth being close to the angular umbilical shoulder. The preserved

fragment bears seven small, sharp, comma-shaped bullae at the shoulder.

These give rise to bundles of fine, dense, flexuous striae which, being crowded

close to the umbilicus, appear as ribs, which fade out at a short distance from

the bulla. The striae flex forwards to mid-flank, then backwards, to form the

very shallow blade of a sickle. They are strongly projected across the ventro-

lateral shoulder. Paralleling these striae are irregular, low folds and constrictions.

The imperfectly exposed suture is illustrated in Figure 20.

Discussion

The presence of well-developed ribs which extend to the umbilical shoulder

indicates this specimen to be a Theganeceras. It differs from all previously

described species in having broader, less markedly flexed ribs when young, as

well as possessing umbilical bullae, which feature is unique for the subgenus.

Only S. (Theganeceras) grande Thomson (1974, pl. 4 (fig. 9)) approaches this.

specimen in size; the unique holotype of that species is quite distinct, however,

lacking bullae and having strongly flexed ribs and striae (Fig. 6G).

Equally, no S. (Sinzovia) bears bullae, and the closest species, S. (S.) stolleyi

Casey (Fig. 6A—D), has ribs which efface on the inner flank. The presence of

bullae also distinguishes our specimen from described S. (Sanmartinoceras) at

comparable dimensions.

Occurrence

Lower Aptian I of northern Zululand only.

Subgenus Sinzovia Sazonova, 1958

Sanmartinoceras (Sinzovia) trautscholdi (Sinzow)

Figs 4I-K, 6E-F

Ammonites bicurvatus Trautschold (non Michelin), 1865: 22, pl. 3 (figs 17a-c).

Oppellia Trautscholdi Sinzow, 1870: 118, pl. A (figs 1, la—b only).

Sinzovia trautscholdi (Sinzow): Sazonova, 1958: 128, pl. 6 (fig. 2), pl. 8 (figs 1, 1a, 3-7),

pl. 10 (figs 4-5) (with synonymy). Druschchitz & Kudryavtseva, 1960: pl. 42 (figs 7a—b).

Collignon 1962: 31, pl. 229 (fig. 974).

? Sanmartinoceras (Sinzovia) sp. cf. trautscholdi (Sinzow): Casey 1961b: 136, pl. 26 (fig. 7).

Holotype

Trautschold’s original of his plate 3 (fig. 17a—c), from the Lower Aptian of

Simbirsk (now Polivna) in the Stalingrad area of Russia, on which Sinzow

(1870: 118) based his species. The original figures are reproduced here as

Figure 6E-F.

113

CRETACEOUS FAUNAS FROM SOUTH AFRICA

“STIX “SOLSZOd-INNVS ‘odAjofoy 9y3 Jo sinjns Jenszed “Aou ‘ds wnsopou (spsadauvsay [,) SDAQIOUIJADUUDS “QT ‘BIA

114 ANNALS OF THE SOUTH AFRICAN MUSEUM

Material

SAM-PCZ5919 from the Makatini Formation, Aptian IV at locality 152,

Mkuze Game Reserve, northern Zululand.

Description

The specimen is a beautifully preserved disc of which the last half of a

whorl is body chamber. It retains extensive areas of replaced shell, in consequence

of which the sutures are not fully decipherable. The dimensions are as follows:

D Wb Wh Wb: Wh U

43,0 10,0(23,3) 22,0(51,2) 0,45 —_-9,0(20,9)

Coiling is very involute, oxycone, with two-thirds of the previous whorl

being covered. The umbilicus is small and shallow (20% of diameter), with a

flat, subvertical wall. The umbilical shoulder is sharp and angular. The whorl

section is very compressed, with a breadth to height ratio of 0,45. The inner

flank is flattened to concave, with gently rounded, convergent outer flanks,

converging to an acute, delicately keeled venter.

The inner, concave part of the flank is virtually smooth where shell is

present, due to weathering. On the mould, however, there are delicate, concave,

prorsiradiate, bunched striae which sweep forwards to a distinct spiral ridge at

mid-flank and a parallel groove external to this. On the ridge and groove, the

striae flex backwards in a marked convex projection. On the outer flank, they

flex backwards to form a ‘blade’ corresponding to the bunches of ‘handles’ to

what is an overall sickle-like ornament. The blades are strong, concave, blunt

ribs, strong over most of the last whorl, although effacing on the venter, which

is smooth on either side of a narrow, sharp ventral keel.

The sutures cannot be seen.

Discussion

On the basis of weakness of inner flank ornament, and strength and style

of ribs on the outer flank, the authors would refer their single specimen to

Sinzow’s species. It is particularly close to the specimen from Ulyanovsk,

Russia, illustrated by Casey (1961), text-fig. 143 (1a—b)).

Sinzovia stolleyi Casey (19616, text-fig. 143(g)) is more distantly and

narrowly ribbed, as can be seen from Figure 6A—D herein.

Sinzovia aptianum (Sarasin) (1893: 155, pls 4-6 (fig. 12a—c); Casey 19615:

134, pl. 26 (fig. 6a—b)) has blunter, fewer ribs on the outer flank.

Sanmartinoceras ? (Sinzovia ?) sp. nov. of Casey (19615: 136, text-fig. 43(f))

is a Lower Albian species of uncertain affinity.

Occurrence

Lower Aptian of the U.S.S.R. and southern England; Upper Aptian of

Madagascar and Zululand.

CRETACEOUS FAUNAS FROM SOUTH AFRICA iS

Family Binneyitidae Reeside, 1927

The Binneyitidae are a family of micromorphs, characteristically platycone,

with much reduced sutures. Two genera, Borissiakoceras Arkhangelsky, 1916,

and Binneyites Reeside, 1927, were referred to the family by Wright (1957) who

suggested derivation from the Acanthocerataceae, but subsequent work has

revealed the presence of passage forms linking the family to the Aconeceratidae

via Falciferella Casey, 1954. This Middle to Upper Albian genus has a vestigal

keel when young and can be linked to the platycone but still feebly carinate

Aptian aconeceratids Doridiscus Casey, 1961, and Nothodiscus Casey, 1962

(in Collignon 1962). The evolutionary origins of the family thus involved a

change from oxycone to platycone with corresponding loss of keel, and sutural

simplification.

Genus Borissiakoceras Arkhangelsky, 1916

Type species

Borissiakoceras mirabile Arkhangelsky, 1916.

Diagnosis

Small, compressed, moderately involute to moderately evolute platycones.

Flanks typically smooth, sometimes bearing narrow, falcoid ribs. A few species

bear ventrolateral tubercles. Suture simple with narrow, bifid lobes and broader,

bifid or trifid saddles.

Discussion

Borissiakoceras is a distinctive genus, differing obviously from the later

(Coniacian) Binneyites Reeside, 1927, which has stronger lateral and ventro-

lateral ornament, sharp ventrolateral shoulders and more auxiliary elements in

the suture. Johnsonites Cobban, 1961, has a simpler suture and a flat or concave

venter.

Borissiakoceras is clearly descended from Falciferella, species of which

extend to the Upper Albian. It is the only binneyitid in which dimorphism has

been demonstrated (Kennedy & Cobban 1976: pl. 1 (figs 3-4)); males have

stronger, more markedly biconcave growth striae and ribs, reflecting a similar

aperture, with a short rostrum.

Occurrence

Borissiakoceras is best known from North America, ranging from Texas

(Stephenson 1952, 1955) to Kansas and Colorado (Morrow 1935). Other United

States occurrences are summarized by Cobban (1961). It also occurs in British

Columbia (Warren & Stelck 1958) and Alaska (Cobban & Gryc 1961). In these

regions it ranges from Middle Cenomanian to Lower Turonian. The type

species comes from Turkestan and is probably of late Cenomanian age;

116 ANNALS OF THE SOUTH AFRICAN MUSEUM

Bodylevsky & Shulinga (1958) record the genus from the Turonian—Coniacian

of the northern U.S.S.R., and Kennedy & Juignet (1973) record it from the

Middle Cenomanian of Normandy, whilst the present authors have seen a

fragment from the Lower Cenomanian of Sarthe (Sorbonne collections).

Wright (1963) records a doubtful species from the Middle Cenomanian of

northern Australia, and the Engonoceratidae gen. et sp. nov. of Henderson

(1973: 106, pl. 14 (fig. 8)) belongs here, as do the Middle Cenomanian specimens

from Zululand described below.

Borissiakoceras sp.

Figs 14D-F, 15A-E

Material

Two specimens, BMNH C80003, and no. 16 in M. R. Cooper’s collection,

University of Natal, Durban, both from the Middle Cenomanian (Cenomanian

III) Locality 62, the Skoenberg, Zululand.

Description

The specimens comprise a small phragmocone and a body chamber with a

maximum whorl height of 7,4 mm. Coiling is involute with a small, shallow

umbilicus comprising approximately 20 per cent of the diameter. The overall

form is platycone, the whorl breadth to height ratio being approximately 0,5,

with a low umbilical wall, flattened inner, and slightly convergent outer flanks,

abruptly and narrowly rounded ventrolateral shoulder, and somewhat flattened

venter. Both specimens are corroded, so that no trace of any original ornament

remains. The suture is poorly exposed, but much simplified, with such elements

as are visible resembling those of B. mirabile.

Discussion

Overall shell form and suture indicate these specimens to be Borissiakoceras,

the first representatives of the genus, and indeed the Binneyitidae, to be described

from Africa. Because of poor preservation it is not possible to identify them

fully. They most closely resemble feebly ornamented species such as B. mirabile

(see Kennedy & Juignet 1973) and B. orbiculatum (see Cobban, 1961: 750,

pl. 88 (figs 15-41), text-figs 5a-f).

Occurrence

Middle Cenomanian of Zululand.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 117

ANNOTATED LIST OF SPECIES REFERRED TO THE

ACONECERATIDAE AND BINNEYITIDAE

Family Aconeceratidae Spath, 1923

Genus Protaconeceras Casey, 1954

Type species: Oppelia patagoniensis Favre, 1908, by original designation.

Protaconeceras patagoniense (Favre), 1908: 634, pl. 34 (fig. 7), pl. 37 (figs 3-5), text-fig. 6.

Lower Hauterivian of Patagonia.

Protaconeceras spp. nov. Casey, 1954: 270, pl. 7 (fig. 7), text-fig. 2. Upper Hauterivian,

England.

Genus Aconeceras Hyatt, 1903

(= Adolphia Stolley, 1907; Adolphites Hennig, 1932)

Type species: Ammonites nisus d’Orbigny, 1841, by monotypy.

Aconeceras nisus (d’Orbigny), 1841: 184, pl. 55 (figs 7-9). Casey 19616: 128. Upper Aptian of

western Europe, notably Gargasian clays of the Vocontian Trough. Also recorded from

Madagascar and elsewhere, although many records are dubious.

Aconeceras haugi (Sarasin), 1893: 156, pls 4-6, text-fig. lla—c. Lower Aptian of western

Europe, Nepal (Bordet et a/. 1971), and Madagascar (Collignon 1962) where it is said to

be of Upper Aptian age.

Aconeceras neonisoides Casey, 19616: 129, pl. 26 (figs 1, 9-10), text-fig. 41d—e. Lower Albian

of southern England and north Africa (Sornay 1955; Dubourdieu 1956).

Aconeceras australonisoides Brunnschweiler, 1959: 11, pl. 1 (fig. la—b). Aptian of western

Australia.

Aconeceras whitehousei Brunnschweiler, 1959: 12, pl. 1 (fig. 2a—b). Aptian of western Australia.

Aconeceras walshense (Etheridge), 1892: 493, pl. 42 (figs 10-11). Whitehouse 1926: 203, pl. 34

(fig. 1), pl. 37 (fig. 3); 1927a: 114, pl. 16 (figs 2-3), text-figs 1, 6-7. Aptian of Queensland.

Aconeceras nisoides (Sarasin), 1893: 155, pls 4-6 (fig. 10a—c), text-figs 3, 5. Lower Aptian of

western Europe.

Aconeceras luppovie (Sazonova), 1958: 130, pl. 8 (fig. 2). Aptian of the U.S.S.R.

Aconeceras saratoviensis (Sazonova), 1958: 130 (= Oppelia trautscholdi Sinzow, 1898, pl. A

(figs 4-5 only)). Lower Aptian of the U.S.S.R.

Genus Sanmartinoceras Bonarelli, 1921

Type species: Sanmartinoceras patagonicum Bonarelli, 1921, by monotypy.

Sanmartinoceras (Sanmartinoceras) patagonicum Bonarelli, 1921, in Bonarelli & Nagera 1921:

27, pl. 5 (figs 3-6). See also Howarth 1958: 5, pl. 1 (figs 6-10); Leanza 1970: 215, fig. 14;

Thomson 1974: 24, pl. 4b-f, text-fig. 7a. Aptian of Argentina and Antarctica. (See

Fig. 6A—D herein.)

Sanmartinoceras (Sanmartinoceras) groenlandium Rosenkrantz, 1934, in Bogvad & Rosen-

krantz 1934: 20, pl. 4 (fig. 3), pl. 5 (figs 1-5). Aptian of east Greenland. (See Fig. 1A—-C

herein.)

Sanmartinoceras (Sanmartinoceras) olenae (Tenison-Woods), 1883: 150, pl. 7 (fig. 8), pl. 8

(fig. 1). Whitehouse 1926: 205, pl. 41 (fig. 3); 1927a: 117, pl. 17 (fig. 6), text-figs 3, 4, 9.

Aptian of Australia. (See Fig. 3A—C herein.)

Sanmartinoceras (Sanmartinoceras) fontinale (Hudleston), 1890: 241, pl. 9 (fig. 1). Whitehouse,

1927a: 116, pl. 17 (figs 2-5). Aptian of Australia. (See Figs 3D-G, 7A-H herein.)

Sanmartinoceras (Sanmartinoceras) africanum Kennedy & Klinger, 1978 sp. nov. (See p. 96.)

Upper Barremian of Zululand.

Subgenus Sinzovia Sazonova, 1958

Type species: Ammonites trautscholdi Sinzow, 1870 (= Ammonites bicurvatus Trautschold,

1865 non Michelin, 1838) by original designation.

Sanmartinoceras (Sinzovia) trautscholdi (Sinzow), 1870: 118-119. See also Ammonites bicurvatus

Trautschold, 1865: 22, pl. 3 (fig. 17a—c) (non Michelin); Casey, 1961b: text-fig. 135a-c.

Lower Aptian of the U.S.S.R. and southern England; Upper? Aptian of Madagascar,

Upper Aptian of Zululand.

118 ANNALS OF THE SOUTH AFRICAN MUSEUM

Samartinoceras (Sinzovia) aptianum (Sarasin), 1893: 155, pls 4-6, fig. 12a—c. Casey 1961b:

134, pl. 26 (fig. 6a—b), text-fig. 43d—e. Aptian of France, southern England and elsewhere

in Europe; Nepal (Bordet et al. 1971). (See Fig. 6E-F herein.)

Sanmartinoceras (Sinzovia) stolleyi Casey, 1961b: 133, 136; text-fig. 135g—h. Upper Aptian of

Germany. (See Fig. 6A—D herein.)

Sanmartinoceras ? (Sinzovia ?) sp. nov. Casey, 1961b: 136, text-fig. 43f. Aptian of England.

vnon Sinzovia luppovie Sazonova, 1958: 130, pl. 8 (fig. 2) = S. luppovi Casey, 19616: 130.

Lower Aptian of the U.S.S.R. An Aconeceras according to Casey (19616: 133).

?non Sinzovia saratoviensis Sazonova, 1958 (= Oppelia trautscholdi Sinzow, 1898, pl. A

(figs 4-Sa only). Lower Aptian of the U.S.S.R. Also an Aconeceras according to Casey

(19615: 133).

Subgenus Theganeceras Whitehouse, 1926

Type species: Oppelia scalata von Koenen, 1902, by original designation.

Sanmartinoceras (Theganeceras) grande Thomson, 1974: 25, pl. 4g. Lower Aptian of Alexander

Island. (See Fig. 6G herein.)

Sanmartinoceras (Theganeceras) scalatum (von Koenen), 1902: 54, pl. 45 (fig. 6). Lower Aptian

of north Germany. (See Fig. 6H herein.)

Sanmartinoceras (Theganeceras) falcatum (yon Koenen), 1902: 48, pl. 45 (figs 7-8). See also

Casey 19616: 132, pl. 26 (fig. 2). Lower Aptian of north Germany and England. (See

fig. 6I-J.)

Sanmartinoceras (Theganeceras) (?) sp. Thomson, 1974: 26, pl. 4k. Lower Aptian of Alexander

Island.

Sanmartinoceras (Theganeceras) nodosum Kennedy & Klinger sp. nov. (See p. 107.) Lower

Aptian of Zululand.

Genus Gyaloceras Whitehouse, 1927

Type species: Gyaloceras smithi Whitehouse, 1927 by original designation.

Gyaloceras smithi Whitehouse, 1927a: 115, pl. 17 (fig. 1), text-fig. 8. Aptian of Australia.

(See Fig. 2A.)

Gyaloceras ibo Reyment, 1955: 15, pl. 2 (figs 1-3), text-figs 3-4. Upper Albian of Nigeria.

Casey (19616: 139) has suggested that this species is ‘a completely new development whose

relationship to the Aconeceratidae is doubtful. Possibly it is congeneric with the

Aconeceras ? described by Haas (1942: 165) from the Upper Albian of Angola’.

? Genus Eofalciferella Brunnschweiler, 1959

Type species: Eofalciferella condoni Brunnschweiler, 1959, by original designation.

Eofalciferella condoni Brunnschweiler, 1959: 13, pl. 1 (figs. 3-4). Aptian of Australia. The only

illustrations of this species are pencil sketches of the unique holotype, a crushed specimen

from the Windalia Radiolarite. The species and genus are best treated as nomen dubia;

Casey (19615: 131) implies in his discussion of the genus that its affinities may be with

Sanmartinoceras.

Family Binneyitidae Reeside, 1927

Genus Falciferella Casey, 1954

Type species: Falciferella milbournei Casey, 1954, by original designation.

Falciferella milbourni Casey, 1954: 274, pl. 7 (figs 1-5), text-fig. 3. Middle Albian of southern

England.

Falciferella malandiandrensis Collignon, 1962: 32, pl. 229 (fig. 975). Upper Aptian of

Madagascar. This species is keeled to a diameter of approximately 45 mm. It may be an

Aconeceras, although Collignon (1962) indicates that it has a Falciferella-like suture.

Genus Doridiscus Casey, 1961

Type species: Doridiscus rotulus Casey, 1961, by original designation.

Doridiscus rotulus Casey, 1961b: 139, pl. 26 (fig. 8a—b), text-fig. 44c-e. Low Upper Aptian of

southern England.

oe sp. noy. indet? Casey, 19615: 140, text-fig. 44f. Upper Lower Aptian of southern

ngland.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 119

Genus Nothodiscus Casey in Collignon, 1962

Type species: Nothodiscus planus Casey, in Collignon, 1962, by original designation.

Nothodiscus planus Casey in Collignon, 1962: 32, pl. 229 (fig. 976). Upper Aptian of Madagascar.

Genus Borissiakoceras Arkhangelsky, 1916

Type species: Borissiakoceras mirabile Arkangelsky, 1916: 55, pl. 8 (figs 2-3), Lower Turonian

of Turkestan. Kennedy & Juignet, 1973: 900, text-figs 1-2, Middle Cenomanian of France.

Borissiakoceras compressum Cobban, 1961: 747, pl. 87 (figs 19-33); pl. 89 (figs 1-9), text-

fig. 4a-k. Middle Cenomanian of the United States Western Interior.

Borissiakoceras reesidei Morrow, 1935: 463, pl. 49 (fig. 7a—b), pl. 50 (fig. 5), text-fig. 8.

Cobban, 1961: 749, pl. 88 (figs 1-14), text-fig. 3h-k. Middle Cenomanian of the United

States Western Interior.

Borissiakoceras orbiculatum Stephenson, 1955: 64, pl. 6 (figs 1-4). Cobban, 1961: 750, pl. 88

(figs 15-41), text-fig. Sa-f. Middle/Upper Cenomanian of Texas and the United States

Western Interior.

Borissiakoceras cf. B. orbiculatum Stephenson, 1955; Cobban 1961: 753, pl. 89 (figs 10-14),

text-fig 5g, i. Uppermost Cenomanian of the Black Hills, United States Western Interior.

Borissiakoceras inconstans Cobban & Gryc, 1961: 187, pl. 38 (figs 30-37), text-fig. 21-1.

Latest ? Cenomanian of Alaska.

Borissiakoceras ashurkoffae Cobban & Gryc, 1961: 188, pl. 38 (figs 38-43), text-fig. 2j-k, m.

Lower Turonian of Alaska.

Borissiakoceras (?) sp. Wright, 1963: 602, pl. 89 (fig. 5). Middle Cenomanian of Bathurst

Island, Australia.

Borissiakoceras ? sp. Lower Cenomanian of Sarthe, France (Sorbonne collections).

Borissiakoceras sp. Kennedy & Klinger, 1978. (See p. 116.) Middle Cenomanian of Zululand.

Genus Johnsonites Cobban, 1961

Type species: Johnsonites sulcatus Cobban, 1961, by original designation.

Johnsonites sulcatus Cobban, 1961: 743, pl. 87 (figs 1-18), text-fig. 3a-g. Middle Cenomanian

of Wyoming and Colorado in the United States Western Interior.

? Johnsonites sp., the original of Stephenson’s (1952: 198, pl. 45 (figs 5-6)) Euhoplites ? sp.

from the Middle Cenomanian of Texas may also belong to this genus.

Genus Binneyites Reeside, 1927

Type species: Binneyites parkensis Reeside, 1927, by original designation.

Binneyites parkensis Reeside, 1927: 5, pl. 3 (figs 1-10). Cobban, 1961: 754, pl. 89 (figs 32-37),

text-fig. 5s, t. Coniacian of Wyoming and Utah in the United States Western Interior.

Binneyites carlilensis Cobban, 1961: 755, pl. 89 (figs 15-22), text-fig. 5h, j-m. Mid-Turonian

of South Dakota and Wyoming in the United States Western Interior.

Binneyites aplatus (Morrow), 1935: 465, pl. 49 (fig. 5), pl. 50 (fig. 6), text-fig. 7. Mid-Turonian

of Kansas in the United States Western Interior.

Binneyites rugosus Cobban, 1961: 756, pl. 89 (figs 26-31), text-fig. 5n—p.

The Engonoceratidae gen. et sp. nov. of Henderson, 1973: 106, fig. 14 (no. 8), text-fig. 15,

is either a Borissiakoceras or Binneyites of Turonian age.

ACKNOWLEDGEMENTS

We thank Dr H. W. Ball, Dr M. K. Howarth and Mr D. Phillips of the

British Museum (Natural History), Dr R. Casey and Mr C. J. Wood (Institute

of Geological Sciences), Mr C. W. Wright and Dr M. R. Cooper (Oxford), for

their advice and useful discussions. The assistance of the staff of the Geological

Collections at Oxford University Museum, and South African Museum,

Cape Town, is gratefully acknowledged as is the financial support of the Trustees

of the Sir Henry Strakosh Bequest, the Royal Society, and Natural Environment

Research Council.

120 ANNALS OF THE SOUTH AFRICAN MUSEUM

REFERENCES

ARKHANGELSKY, A. D. 1916. Les mollusques du Crétacé Supérieur du Turkestan. Trudy

geol. Kom. n.s. 152: 1-57. ;

BopDyLevsky, VY. I. & SHULINGA, N. I. 1958. Jurassic and Cretaceous faunas of the lower

Yenisei River region, U.S.S.R. Trudy nauchno-issled. Inst. Geol. Arkt. 93: 1-196

(In Russian.)

BoGvaD, R. & ROSENKRANTZ, A. 1934. Beitrage zur Kenntnis der unteren Kreide Ostgr6nlands.

Medar. Gronland 93: 15-28.

BONARELLI, G. & NAGERA, J. J. 1921. Observaciones geoldgicas en las immendiaciones del

Lago San Martin (Territorio de Santa Cruz). Boln Dir. gen. Minas Geol. Hidrol., B. Aires

ser. B. (Geoldgia) 27: 1-39.

BORDET, P., COLCHEN, L., KRUMMENACHER, D., LE Fort, P., MOUTERDE, R., & REMy, M.

1971. Recherches Géologiques dans Himalaya du Népal, Région de la Thakkhola.

Paris: Centre National de la Recherche Scientifique.

BRUNNSCHWEILER, R. O. 1959. New Aconeceratinae (Ammonoidea) from the Albian and

Aptian of Australia. Bull. Bur. Miner. Resour. Geol. Geophys. Aust. 54: 1-19.

CALLOMON, J. H. 1963. Sexual dimorphism in Jurassic ammonites. Trans. Leicester lit. phil. Soc.

57: 21-56.

Casey, R. 1954. Falciferella, a new genus of Gault ammonites, with a review of the family

Aconeceratidae in the British Cretaceous. Proc. Geol. Ass. 65: 262-277.

Casey, R. 1961a. A monograph of the Ammonoidea of the Lower Greensand. Palaeontogr.

Soc. (Monogr.) part 2: 45-118.

Casey, R. 19615. A monograph of the Ammonoidea of the Lower Greensand. Palaeontogr.

Soc. (Monogr.) part 3: 119-216.

CosBan, W. A. 1961. The ammonite family Binneyitidae Reeside in the Western Interior of the

United States. J. Paleont. 35: 176-190.

CosBAN, W. A. & Gryc, G. 1961. Ammonites from the Seabee Formation (Cretaceous) of

northern Alaska. J. Paleont. 35: 176-190.

COLLIGNON, M. 1962. Atlas des fossiles carctéristiques de Madagascar (Ammonites) IX (Aptien).

Tananarive: Service Géologique.

DrusHcuitTz, V. V. & KupRYAVTSEVA, M. P. 1960. Atlas of Lower Cretaceous fauna of the

northern Caucasus and Crimea. Moscow: Gostoptekhizdat. (In Russian.)

DusourDIEv, G. 1956. Etude géologique de la région de l’Ouenza (Confins Algéro-Tunisiens).

Publs Serv. Carte géol. Algér. 10: 1-659.

ETHERIDGE, R. 1892. In: Jack, R. L. & ErHertpGe, R. The geology and palaeontology of

Queensland and New Guinea. Brisbane & London.

Favre, F. 1908. Die Ammoniten der unteren Kreide Patagoniens. Neues Jb. Min. Geol.

Beil. Bd. 35: 601-647.

Haas, O. 1942. The Vernay collection of Cretaceous (Albian) ammonites from Angola.

Bull, Amer. Mus. nat. Hist. 81: 1-224.

HENDERSON, R. A. 1973. Clarence and Raukumara Series (Albian-?Santonian) Ammonoidea

from New Zealand. J/ R. Soc. N.Z. 3: 71-123.

HENNIG, E. 1932. Wesen und Wege der Palédontologie. Berlin.

Howarth, M. K. 1958. Upper Jurassic and Cretaceous ammonite faunas of Alexander and

Graham Land. Scient. Rep. Falkld Isl. Depend. Sury. 21: 1-16.

HUuDLEsToN, W. H. 1890. Further notes on some Mollusca from South Australia. Geol.

Mag. (3) 7: 241-246.

Hyatt, A. 1903. Pseudoceratites of the Cretaceous. STANTON, T. W. ed. Monogr. U.S. geol.

Sury. 44: 1-351.

KENNEDY, W. J. & CosBAn, W. A. 1976. Aspects of ammonite biology, biogeography &

biostratigraphy. Spec. Pap. Palaeont. 8: 1-133.

KENNEDY, W. J. & JuIGNET, P. 1973. First record of the ammonite family Binneyitidae

Reeside, 1927 in western Europe. J. Paleont. 47: 900-902.

KENNEDY, W. J. & KLINGER, H. C. 1975. Cretaceous faunas from Zululand and Natal, South

Africa. Introduction, stratigraphy. Bull. Br. Mus. nat. Hist. (Geol.) 25: 263-315.

KOENEN, A. VON. 1902. Die Ammonitiden des Norddeutschen Neocom. (Valanginien,

Hauterivien, Barrémien und Aptien). Abh. preuss. geol. Landesanst. n.s. 24: 1-451.

ANNALS OF THE SOUTH AFRICAN MUSEUM 121

KULLMAN, J. & WIEDMANN, J. 1970. Significance of sutures in phylogeny of Ammonoidea

Paleont. Contr. Univ. Kans. 47: 1-32.

LEANZA, A. F. 1970. Ammonites nuevos 0 pocos conocidos del Aptiano, Albiano y Cenomanino

de los Andes australes con notas acera de su posicion estratigrafica. Revta Soc. geol.

argent. 25: 197-261.

MakowskI, H. 1962. Problems of sexual dimorphism in ammonites. Palaeont. pol. 12: 1-92.

Morrow, A. L. 1935. Cephalopods from the Upper Cretaceous of Kansas. J. Paleont. 9:

463-473.

OrpiGNy, A. D’. 1840-1842. Paléontologie frangaise: Terrains crétacés. 1. Céphalopodes.

Paris: Masson.

PALFRAMAN, F. F. B. 1969. Taxonomy and sexual dimorphism in ammonites. Morphogenetic

evidence in Hecticoceras brightii (Pratt). In: WESTERMANN, G. E. G. ed. Sexual dimorphism

in fossil Metazoa and taxonomic implications: 126-154. Stuttgart: Schweizerbart’sche

Verlagsbuchhandlung.

ReesIDE, J. B. 1927. Cephalopods from the lower part of the Cody Shale of Oregon Basin,

Wyoming. Prof. pap. U.S. geol. Surv. 150-A: 1-19.

REYMENT, R. A. 1955. The Cretaceous Ammonoidea of southern Nigeria and the Southern

Cameroons. Bull. geol. Surv. Nigeria 25: 1-112.

SARASIN, C. 1893. Etude sur les Oppelia du groupe de nisus et les Sonneratia du groupe du

bivurvatus et du raresulcatus. Bull. Soc. géol. Fr. (4) 21: 149-164.

SAZONOVA, I. G. 1958. Lower Cretaceous deposits of the Russian Platform. /n: FLERovoI, O. V.

Mesozoic and Tertiary deposits of the central regions of the Russian Platform. Vses.

Nauchno-Issled. Geol. Razved. Neft. Inst. Moscow: 31-136. (In Russian.)

Sinzow, I. 1870. A geological sketch of the Saratow region. Bull. Soc. Mem. St. Petersb. (2)

5: 103-161. (In Russian.)

SINZow, I. 1898. Bemerkungen iiber einige Ammoniten des Aptien. Odessa.

Sornay, J. 1955. Ammonites nouvelles du Crétacé de la région des Monts du Meéllegue

(Constantine). Bull. Serv. Carte géol. Alger (1) Paléontologie 18: 1-40.

SpaTH, L. F. 1923. A Monograph of the ammonoidea of the Gault. Part 1. Palaeontogr. Soc.

Monogr. 76: 1-72.

STEPHENSON, L. W. 1952. Larger invertebrate fossils of the Woodbine Formation (Cenomanian)

of Texas. Prof. pap. U.S. geol Surv. 242: 1-226.

STEPHENSON, L. W. 1955. Basal Eagle Ford fauna (Cenomanian) in Johnson and Tarrant

counties, Texas. Prof. pap. U.S. geol. Surv, 274—C: 53-67.

STOLLEY, E. 1907. Ueber ein norddeutsches aequivalent der Clansayes Fauna Siidfrankreichs

und der Schweiz. Centrlbl. Min. Geol. u Pal. 1907: 266-270.

TENISON-Woops, J. E. 1883. On some Mesozoic fossils from the Palmer River, Queensland.

J. Proc. R. Soc. N.S.W.16: 147-154.

THOMSON, M. R. A. 1974. Ammonite faunas of the Lower Cretaceous of south-eastern

Alexander Island. Scient. Rep. Falkld Isl. Depend. Surv. 80: 1-44.

TRAUTSCHOLD, H. 1865. Der Inoceramen-Thon von Simbirsk. Bull. Soc. Imp. Nat. Moscou.

34: 432-457.

WarREN, P. S. & STELCK, C. R. 1958. Lower Cenomanian Ammonoidea and Pelecypoda from

Peace River area, western Canada. Bull. geol. Div. Res. Coun. Alberta 2: 36-51.

WEDEKIND, R. 1916. Uber Lobus, Suturallobus und Inzision. Zentbl. Miner. Geol. Paldont.

1916: 185-195.

WHITEHOUSE, F. W. 1926. The Cretaceous Ammonoidea of eastern Australia. Mem. Qd Mus.

8: 195-242.

WHITEHOUSE, F. W. 1927a. Additions to the Cretaceous ammonite fauna of eastern Australia.

Part 1. (Simberskitidae, Aconeceratidae and Parahoplitidae). Mem. Qd Mus. 9: 109-120.

WHITEHOUSE, F. W. 1927b. Additions to the Cretaceous ammonite fauna of eastern Australia.

Part 2. Mem. Od Mus. 9: 200-206.

WRIGHT, C. W. 1957. Mesazoic Ammonoidea. In: Moorg, R. C. ed. Treatise on Invertebrate

Palaeontology, Part L, Mollusca 4: Kansas; New York: University of Kansas Press and

Geological Society of America.

Wricut, C. W. 1963. Cretaceous ammonites from Bathurst Island, northern Australia.

Palaeontology 6: 597-614.

6. SYSTEMATIC papers must conform to the /nternational code of zoological nomenclature

(particularly Articles 22 and 51).

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be

followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.

nov., syn. nov., etc.

An author’s name when cited must follow the name of the taxon without intervening

punctuation and not be abbreviated; if the year is added, a comma must separate author’s

name and year. The author’s name (and date, if cited) must be placed in parentheses if a

species or subspecies is transferred from its original genus. The name of a subsequent user of

a scientific name must be separated from the scientific name by a colon.

Synonymy arrangement should be according to chronology of names, i.e. all published

scientific names by which the species previously has been designated are listed in chronological

order, with all references to that name following in chronological order, e.g.:

Family Nuculanidae

Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-15A

Nucula (Leda) bicuspidata Gould, 1845: 37.

Leda plicifera A. Adams, 1856: 50.

Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).

Nucula largillierti Philippi, 1861: 87.

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

Note punctuation in the above example:

comma separates author’s name and year

semicolon separates more than one reference by the same author

full stop separates references by different authors

figures of plates are enclosed in parentheses to distinguish them from text-figures

dash, not comma, separates consecutive numbers

Synonymy arrangement according to chronology of bibliographic references, whereby

the year is placed in front of each entry, and the synonym repeated in full for each entry, is

not acceptable.

In describing new species, one specimen must be designated as the holotype; other speci-

mens mentioned in the original description are to be designated paratypes; additional material

not regarded as paratypes 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. Aduit 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, it not preceded

by initials or full names

e.g. DuToit but A.L.du Toit; Von Huene but F. von Huene

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.’ et

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. i

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.