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

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FISCHER, P. H., DUVAL, M. & RAFFY, A. 1933. Etudes sur les échanges respiratoires des littorines.

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ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 105 Band

February 1997 Februarie

Part” 1 Deel

CRETACEOUS FAUNAS FROM ZULULAND AND

NATAL, SOUTH AFRICA. THE AMMONITE

FAMILY BACULITIDAE GILL, 1871

(EXCLUDING THE GENUS EUBACULITES)

HERBERT CHRISTIAN KLINGER

WILLIAM JAMES KENNEDY

Cape Town Kaapstad

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CRETACEOUS FAUNAS FROM ZULULAND AND NATAL,

SOUTH AFRICA.

THE AMMONITE FAMILY BACULITIDAE GILL, 1871

(EXCLUDING THE GENUS EUBACULITES)

HERBERT CHRISTIAN KLINGER

Division of Earth Sciences, South African Museum, Cape Town

WILLIAM JAMES KENNEDY

Geological Collections, University Museum, Oxford

(With 132 figures)

[MS accepted 17 May 1995]

ABSTRACT

Representatives of the ammonite family Baculitidae are common faunal elements in the

mid- and Upper Cretaceous of Zululand. The genus Lechites is rare, being represented by

a single species in the Upper Albian, L. gaudini (Pictet & Campiche); Sciponoceras is

represented by rare Cenomanian S. roto Cieslifiski, S$. baculoides (Mantell) and common

S. cucullatum Collignon. Baculites is very common, and represented by Coniacian B. yoko-

yamai Tokunaga & Shimizu, Coniacian to Santonian B. capensis Woods, Coniacian to

Campanian B. bailyi Woods, and Campanian B. sulcatus Baily, B. increscens Collignon,

B. vanhoepeni Venzo, B. duharti Hunicken, Baculites sp. aff. B. rectus Marshall and

B. nibelae sp. nov.

CONTENTS

PAGE

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INTRODUCTION

Members of the ammonite family Baculitidae are conspicuous faunal

elements in the mid- and Upper Cretaceous of Zululand. Early representatives

of the family belonging to the genus Lechites are rare, and so far only one

species, L. gaudini (Pictet & Campiche), represented by two specimens, is

known from the Upper Albian of Zululand. Sciponoceras is locally common in

the Cenomanian; the majority of specimens belong to the Lower Cenomanian

S. cucullatum Collignon. Sciponoceras roto Cieslinski and S. baculoides

(Mantell) are rare in the Lower and Middle Cenomanian, respectively. Turonian

sediments are absent in outcrops in Zululand and there is thus no baculitid

record of this stage. The earliest Middle Coniacian baculitids include smooth, or

Ann. S. Afr. Mus. 105 (1), 1997: 1-206, 132 figs, appendix.

D ANNALS OF THE SOUTH AFRICAN MUSEUM

only rarely ribbed forms, referable to B. yokoyamai Tokunaga & Shimizu and

B. bailyi Woods. Nodose baculitids of the group of B. capensis Woods first

appear in the Middle Coniacian, and become very abundant in the Upper

Coniacian of Zululand where they, in places, formed the major part of the

biomass. Concretions crowded with B. capensis are common in parts of the

Middle and Upper Coniacian. Baculites capensis and B. bailyi are common up

to the Lower or the Middle Santonian in southern Africa, but both occur as late

as the Early Campanian. Ribbed B. sulcatus Baily occurs in the Lower Cam-

panian of Pondoland. In Zululand and Natal, ribbed to nodose baculitids refer-

able to B. increscens Collignon, B. vanhoepeni Venzo and B. nibelae sp. nov.

occur in the Lower and Middle, and possibly part of the Upper Campanian,

respectively. A smooth species, B. duharti Hunicken is recorded from the

Middle Campanian.

In the Maastrichtian of Zululand, the genus Eubaculites replaces Baculites

completely, and locally forms a major part of the total biomass. Representatives

of the genus Eubaculites from Zululand have been described earlier by Klinger

(1976) and Klinger & Kennedy (1993) and include E. carinatus (Morton),

E. labyrinthicus (Morton), E. latecarinatus (Brunnschweiler) and E. simplex

(Kossmat).

CLASSIFICATION

Rigel

Until recently, authorship of the family Baculitidae was ascribed to Meek

(1876) (see e.g. Wright 1957: L218; Luppov & Drushchits 1958: 64). How-

ever, the name Baculitidae had been introduced five years earlier by Gill, in

1871, and authorship rests with him, as correctly indicated by Wright (1981:

172).

Wiedmann (1962a: 179) suggested that Cretaceous ptychoceratitid, poly-

ptychoceratid, hamitid and baculitid heteromorphs could be united as sub-

families into a single family, Baculitidae, and later (Wiedmann 19625: 93) even

suggested including bochianitids in this ‘super’ family Baculitidae. This radical

reorganization has found few supporters apart from Scholz (1979). According to

a recent review of the higher taxa of Jurassic and Cretaceous Ammonoidea by

Besnosov & Michailova (1991), Wiedmann’s classification would involve

lumping genera of two different orders into a single family! We regard the

family Baculitidae as a rather conservative group that probably had its origins in

the genus Hamites in the Albian, and retained a more or less straight shell

throughout its existence right up to the end of the Cretaceous Period.

The earliest baculitid, Lechites, arose in the Upper Albian from the genus

Hamites. The adult shell of Lechites consists of a single, straight shaft; as yet,

the ammonitella is unknown. Ornament is simple, consisting of circum-

peripheral ribs. Constrictions are rare. One small branch of Lechites, elevated

to subgeneric rank as L. (Tuberolechites) by Cooper & Kennedy (1977),

developed feeble tubercles on the venter. There is considerable variation in the

symmetry of the umbilical lobe in Lechites (see e.g. Wiedmann & Dieni 1968:

63, text-fig. 36; Scholz 1979: 14, text-fig. 5A); it may differ on either side of

the same individual. Recent comprehensive reviews of the genus by Cooper &

CRETACEOUS FAUNAS FROM SOUTH AFRICA 3

Kennedy (1977) and Scholz (1979) differed somewhat in details, but covered

most of the taxonomic problems. The distribution of the genus is more or less

cosmopolitan.

Baculites (?) (Protobaculites) ambiguus Collignon (1964: 9, pl. 319

(fig. 1375)), recorded from a single specimen from the Lower Cenomanian of

Madagascar, was regarded by Collignon as a possible precursor of Baculites, as

indicated by the etymology. Ornament consists of annular ribs and periodic

constrictions between every fifth and sixth rib. The suture line has a trifid

umbilical (U) lobe and a large internal (I) lobe. Kennedy & Wright (1994) have

shown that Baculites (?) (Protobaculites) ambiguus is, in fact, not a baculitid,

but a Hemiptychoceras.

Lechites gave rise to the next-oldest baculitid genus, Sciponoceras, in the

Early Cenomanian. The main feature of Sciponoceras is the regular occurrence

of constrictions. The last representatives of Sciponoceras occur in the Upper

Turonian. As in Lechites, Sciponoceras has a more or less cosmopolitan

distribution.

The first true Baculites occur in the Lower Turonian and differ from

contemporary Sciponoceras mainly by the loss of regular constrictions. The

earliest, Turonian, Baculites are cosmopolitan but later representatives become

restricted to distinct biogeographic regions. The genus Baculites persists up to

the end of the Maastrichtian, and is amongst the last ammonite genera before

the extinction of the order Ammonoidea at the K/T boundary (see e.g.

Birkelund 1979, 1993; Ward & Kennedy 1993).

Hamites

Lechites Criobaculites

Sciponoceras Trachybaculites

L. (Tuberolechites) Fresvillia

Baculites

AVGILITNOWA AO ANHDO'IAHd

Pseudobaculites

Boehmoceras \ Eubaculites

Fig. 1. Suggested phylogeny of the ammonite family Baculitidae Gill, 1871.

Time axis not to scale.

4 ANNALS OF THE SOUTH AFRICAN MUSEUM

During the Late Coniacian, an insignificant baculitid lineage, Pseudo-

baculites, arose from Baculites in the United States Western Interior. It differed

from contemporary Baculites mainly in being larger, having a more rapid rate

of taper, and a far more complex suture line. Pseudobaculites is an endemic

lineage, the last representatives occurring in the Lower Maastrichtian; no

representatives are known outside the U.S. Western Interior.

Boehmoceras, another short-lived but apparently slightly more widely

distributed lineage, evolved from Baculites during the Late Santonian. The shell

is coiled in an open criocone, and it is interpreted as a recoiled baculitid. It is so

far known only from the Upper Santonian of Germany, Sweden, Austria,

France, Mississippi, Alabama and Texas; it is rare in Europe, but common in

the Gulf Coast region of the U.S.A.

During the Late Campanian, the genus Eubaculites arose from Baculites,

differing from the latter mainly by the development of a tabulate or acute venter

and pyriform whorl section. Eubaculites is the dominant baculitid of the Maas-

trichtian in the Southern Hemisphere, excluding Antarctica and New Zealand. It

is rare in Western Europe, and absent or not yet recorded from the Middle East

and North and West Africa. It is also absent from the U.S. Western Interior, but

occurs in the Pacific, Gulf Coast and Atlantic Seaboard regions of the U.S.A.

A number of aberrant baculitids are known from the Maastrichtian but their

origins and relations to Baculites are not clear. They all probably merit generic

separation and include:

(1) Fresvillia Kennedy, 1986, type species Fresvillia constricta Kennedy, 1986a

(p. 62, pl. 14 (figs 39-42), text-fig. 10a) from the Upper Maastrichtian of

France. Also included in the genus is Baculites teres Forbes, 1846 (p. 115,

pl. 10 (fig. 5)), recorded from the Upper Maastrichtian of Southern India and

tentatively from the Maastrichtian of California (Matsumoto 1959: 163, pl. 45

(figs Sa-d, 6a-—c), text-figs 82a—-c, 83) and southern Alaska (Jones 1963: 29,

pl. 16 (figs 10-12, 14), text-fig. 14).

(2) Trachybaculites Cobban & Kennedy, 1995, type species Baculites columna

Morton, 1834 (p. 44, pl. 19 (fig. 8)) (see Cobban & Kennedy 1992c, 1995)

from the Maastrichtian of Alabama, Texas, Mississippi, California and South

Dakota. This is a heterochronous homoeomorph of Albian Lechites with

circumperipheral ribbing, but with apparently simplified sutures.

Baculites lechitides Brunnschweiler, 1966 (p. 23, pl. 1 (figs 1-3), text-

fig. 8) from the Upper Maastrichtian of Western Australia probably also belongs

here and possibly B. kegeli Oliveira (1957: 22, pl. 2 (figs 6-7), text-fig. 1) from

the Maastrichtian of Brazil. According to Cobban & Kennedy (1995: 29),

Baculites vicentei Stinnesbeck, 1986 (p. 203, pl. 9 (fig. 4), pl. 10 (figs 3-6),

text-figs 23a—c) from the Maastrichtian of Chile also belongs to Trachybaculites.

(3) The group of Hamites trabeatus Morton, 1834 (p. 45, pl. 15 (fig. 3)) from

the Upper Maastrichtian of Alabama, appears to be an endogastrically recoiled

baculitid lineage, homoeomorphic with Santonian Boehmoceras but for the

position of the siphuncle. Baculites? sp. of Cobban & Kennedy (1992c: 68,

figs 1.1-1.4, 3.1) from the Maastrichtian Fox Hills Formation of South Dakota

belongs here.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 5

The name Criobaculites gen. nov. is here proposed for this group, with type

species Hamites trabeatus Morton, 1834. Diagnosis: Criocone coiled baculitids

with siphuncle inside the curve (i.e. endogastric).

(4) ?The group of Baculites paradoxus Pervinquiére, 1907 (p. 94, pl. 4

(figs 10-11), text-fig. 24) from the Upper Maastrichtian of Tunisia. We are not

certain if this group belongs to the family Baculitidae at all. These are minute,

straight shells with trigonal saddles and lobes, reminiscent of some Scipono-

ceras. Baculites indét. of Pervinquiére (1907: 95, pl. 4 (fig. 12a—b), text-fig. 25)

may also belong here. It differs from B. paradoxus mainly in possession of

constrictions. Both are of very dubious baculitid affinities and are better

regarded as allied to Phylloptychoceras.

LOCATION OF SPECIMENS

The following abbreviations are used to indicate the repositories of material

referred to in the text.

SAM _ South African Museum, Cape Town

NMB_ National Museum, Bloemfontein—presently housed in the South

African Museum

™ Transvaal Museum, Pretoria

MRC Prof. M. R. Cooper Collection, University of Durban-Westville

SAS Geological Survey, Pretoria—presently housed in the South African

Museum

GD Institut des Sciences de la Terre de 1l’Université Dijon (ex

Collignon Collection)

BMNH The Natural History Museum, London

MNHP Muséum National d’Histoire Naturelle, Paris

CPC Facultad de Ciencias Exactas, Fisicas y Naturales, Universidad

Nacional de Cérdoba, Argentina

GO Department of Geological Sciences, University of Bologna

FIELD LOCALITIES

Details of field localities are given in Kennedy & Klinger (1975); fuller

descriptions of localities are deposited in the Department of Palaeontology,

Natural History Museum, London, Geological Survey of South Africa, Pretoria,

and Division of Earth Sciences, South African Museum, Cape Town. Additional

localities are referred to in the text.

DIMENSIONS

All dimensions are given in millimetres. MxWb—maximum whorl breadth

(in mm); MxWh—maximum whorl height (Gn mm); MnWb—minimum whorl

breadth Gn mm); MnWh—minimum whorl height (in mm); D—distance between

maximum and minimum whorl height and breadth measurements (in mm);

Wb/Wh—ratio of whorl breadth to whorl height; Ri—Rib or tubercle index:

6 ANNALS OF THE SOUTH AFRICAN MUSEUM

number of ribs or tubercles per whorl height; Ti—taper index (after Matsumoto

& Obata 1963: 4) (Ti = MxWh - MnWh/ D x 100).

SUTURE TERMINOLOGY

The suture terminology of Wedekind (1916) reviewed and discussed by

Kullmann & Wiedmann (1970) is followed here: I = internal lobe, U =

umbilical lobe, L = lateral lobe, E = external lobe.

SYSTEMATIC PALAEONTOLOGY

Class CEPHALOPODA Cuvier, 1797

Order AMMONOIDEA Zittel, 1884

Suborder ANCYLOCERATINA Wiedmann, 1960

Family Baculitidae Gill, 1871

Genus Lechites Nowak, 1908

Type species. Baculites gaudini Pictet & Campiche (1861: 112, pl. 55

(figs 5-9)) by the original designation of Nowak (1908: 350).

Diagnosis

Shell straight, with simple ornament consisting of prorsiradiate ribs, some-

times grouped in pairs; shallow constrictions may be present. Ventral tubercles

present in some. Dimorphic; macroconchs with trumpet-shaped aperture, micro-

conch with down-turned aperture and incipient lappets.

Discussion

Lechites is the oldest of the Baculitidae; it first appeared in the Late Albian

and, as Spath (1941: 660) had previously suggested, was presumably derived

from Hamites. Middle Albian records, as Baculites Sanctae-Crucis Pictet &

Campiche (1861: 109, pl. 55 (figs 1-4)) are probably misidentified protaniso-

ceratids and not Lechites (see e.g. Spath 1939: 572-573; 1941: 661 footnote).

Recent work by Cooper & Kennedy (1977) and Scholz (1979) showed that

some specimens of Lechites in the Upper Albian and Lower Cenomanian

acquire incipient to distinct ventral tubercles, and are homoeomorphs of /dio-

hamites and Pseudoxybeloceras fragments to a certain extent. Cooper &

Kennedy (1977) separated the tuberculate forms as a separate subgenus of

Lechites, Tuberolechites (type species Tuberolechites regifex Cooper &

Kennedy, 1977: 654, fig. 8 (1-15)). Initially Scholz also considered separating

them at least at subgeneric level. On closer examination, Scholz (1979: 15)

found that all transitions occur between simply ribbed L. gaudini and tuberculate

forms in the condensed sequences he studied. According to Scholz, the tubercu-

late forms occur in the Upper Albian and again in the ‘Vraconnian’; he thus

regarded the acquisition of tubercles as an iterative, rather than as a unique

monophyletic event. Consequently, Scholz merely separated tuberculate and

normally ribbed at subspecific level, as Lechites gaudini s.s. and L. gaudini

nodosus Scholz (1979: 15, pl. 1 (figs 11-16)).

CRETACEOUS FAUNAS FROM SOUTH AFRICA a

It is important to note that, in spite of their unique morphology, tuberculate

forms of Lechites are rare. Cooper & Kennedy (1977: 645, 654) examined

nearly 200 specimens of L. gaudini but found only four tuberculate examples;

Scholz (1979: 12, 15) had nearly 700 specimens of L. gaudini, of which only 32

from France and nine from Hungary were tuberculate.

From the extensive descriptions of e.g. Spath (1941), Renz (1968), Cooper

& Kennedy (1977) and Scholz (1979), it is obvious that there is considerable

variation in the shape and density of ribbing, and that transitions occur between

most of the different ‘species’ attributed to the genus. A reduction in the number

of names, as suggested by Cooper & Kennedy (1977) and Scholz (1979), is

probably justifiable.

Occurrence

Lechites is known with confidence only from the Upper Albian. It has been

recorded from Western and Central Europe, North Africa, ?Somalia, Madagas-

car, Zululand, India, North, Central and South America, Antarctica (Thomson

1984: 88—as Baculites; Moncrieff & Kelly 1993: 5) and Japan (Hokkaido).

Lechites gaudini (Pictet & Campiche, 1861)

Figs 2A-C, M, 3

1861 Baculites Gaudini Pictet & Campiche, p. 112, pl. 55 (figs 5-9).

1933 Baculites Gaudini Pictet & Campiche; Collignon, p. 73, pl. 5 (fig. 8, 8a).

non 1936 ~~ Baculites Gaudini Pictet & Campiche: Venzo, p. 118 [60], pl. 10 [6] (fig. 3).

[= Sciponoceras ?cucullatum]

1941 ~=+Lechites gaudini (Pictet & Campiche); Spath, p. 662, pl. 72 (figs 4-7, 9-10),

text-fig. 242. (cum. synon.)

1941 Lechites communis Spath, p. 666, text-fig. 244.

1947 = Lechites Gaudini var. raricosta Breistroffer, p. 78.

? 1968 = Lechites italicus Wiedmann & Dieni, p. 64, pl. 6 (fig. 10), text-fig. 37.

1968 = Lechites campichei Renz, p. 82, pl. 17 (figs 8a-c, 9a-c, 10a-b), text-fig. 29m.

1968 Lechites vraconensis Renz, p. 82, pl. 17 (figs 1la-c, 12a-b, 14a-b, 15a-c),

text-fig. 29b, g-h, k.

? 1971 = Lechites fasciata Scholz, p. 431, figs 1-2.

1977 Lechites gaudini (Pictet & Campiche); Cooper & Kennedy, p. 644, figs 1

(1-38), 2 (1-30), 3, 4 (1-18), 5 (1-15), 6-7, 8 (16-26). (cum. synon.)

1978 — Lechites gaudini (Pictet & Camp.); Scholz, pl. 3 (figs 1, 8).

1979 = Lechites gaudini gaudini (Pictet & Campiche); Scholz, p. 12, pl. 1 (figs 1-9),

text-fig. SA-B.

1982 = Lechites aff. gaudini Pictet & Campiche; Renz, p. 59, pl. 20 (fig. 7a-b).

1983 = Lechites gaudini (Pict. & Camp.); Horvath, pl. 1 (fig. 2).

1984 Lechites gaudini (Pict. & Camp.); Mitiu, p. 83, pl. 2 (figs 6-9).

1984 Lechites gaudini (Pict. & Camp.) transition to L. communis Spath; Mitiu,

p. 85, pl. 2 (figs 10-11).

1984 = Lechites communis Spath, Mitiu, p. 85, pl. 2 (figs 12-13).

‘1985 = Lechites gaudini (Pictet & Campiche); Immel & Seyed-Emami, p. 112, pl. 7

(hig):

1991 Lechites communis (Pictet & Campiche); Ivanov, pl. 4 (fig. 11).

Type

Lectotype, by subsequent designation of Spath (1941, p. 663) the specimen

figured by Pictet & Campiche (1861, pl. 55 (fig. Sa-c)) and refigured by Renz

ANNALS OF THE SOUTH AFRICAN MUSEUM

Figs 2

CRETACEOUS FAUNAS FROM SOUTH AFRICA 2

(1968, pl. 17 (fig. 3)) from the Upper Albian Sroliczkaia dispar zone of Sainte-

Croix, Kanton Waadt, Switzerland.

Material

SAM-PCZ8766 from locality 182, Zululand, Mzinene Formation. Outcrops

here extend from uppermost Albian to Lower Cenomanian; SAS Z19 from

Haughton’s (1936: 299) locality Z19 on the Pongola River, north of the

confluence with the Mfongozi Creek, Zululand, Mzinene Formation, Upper

Albian.

Description

SAM-PCZ8766 is part of a phragmocone with parts of the original shell

preserved; SAS Z19 is an internal mould of part of the body chamber with the

last septum preserved. The whorl section in both specimens is ovoid, higher

Mmaewice with Wb: Wh ratios 7.7: 8.8 = 0.87 and 7.8: 10.2 = 0.76,

respectively. PCZ8766 is finely ribbed, with 5 ribs per whorl height; Z19 is

more coarsely ribbed with 3 ribs per whorl height.

The suture is partially exposed in PCZ8766 (Fig. 3); the umbilical lobe (U)

is distinctly smaller than the lateral lobe (L) and asymmetric.

Discussion

Lechites gaudini is quite variable as far as density and shape of ribbing is

concerned, and several varietal or specific names have been given to extreme

morphologies. PCZ8766 could be referred to Lechites gaudini s.s., whereas

Z19 could be referred to L. raricosta Breistroffer, 1947. We follow Cooper &

Kennedy (1977: 644), Wiedmann & Dieni (1968: 62) and Scholz (1979: 12) in

including these variants within L. gaudini s.s. Lechites antanimangaensis Col-

lignon (1964: 34, pl. 325 (fig. 1451)) is very similar to our coarsely ribbed

specimen (Z19). However, it has been suggested by Cooper & Kennedy (1977:

653) and Wright & Kennedy (1995: 314) that L. antanimangaensis may be a

Sciponoceras body chamber.

Occurrence

The species is common in the Upper Albian, Stoliczkaia dispar zone. It has

been recorded from England, France, Switzerland, Romania, Hungary, Sar-

dinia, Algeria, Madagascar, southern India, Hokkaido, Mexico, Venezuela and

Antarctica.

Fig. 2 (see facing page). A-C, M. Lechites gaudini (Pictet & Campiche, 1861).

A-C. SAS Z19 from Haughton’s (1936: 299) locality Z19, Zululand, Mzinene Formation,

Albian. M. SAM-PCZ8766 from locality 182, Mzinene Formation; the locality ranges from

the Upper Albian to Lower Cenomanian I. D-F. Sciponoceras baculoides (Mantell, 1822).

SAS A705 from an unspecified horizon on the Skoenberg, Mzinene Formation, Cenomanian.

G-L, N-O. Sciponoceras cucullatum Collignon (1964). G-H. NMBD1004a.

I-K. SAS H210/34. L, N. NMBD1004b. O. NMBD1002a. All from an unspecified

horizon on the Skoenberg, Mzinene Formation, Cenomanian. P. Sciponoceras roto

Cieslinski, 1959. SAS EM191 from an unspecified horizon at locality 181, Zululand,

Mzinene Formation, Cenomanian I or II. All x |

10 ANNALS OF THE SOUTH AFRICAN MUSEUM

(0) 5 mm

[ici Ne ee ae eae

Fig. 3. Lechites gaudini (Pictet & Campiche, 1861).

Suture line of SAM-PCZ8766. Scale bar for size.

Genus Sciponoceras Hyatt, 1894

(= Cyrtochilus Meek, 1876, non Jakowlew, 1875; Cyrtochilella Strand, 1929)

Type species: Hamites baculoides Mantell, 1822: 123, pl. 23 (figs 6-7), by

the original designation of Hyatt (1894: 578). |

Diagnosis

According to the most comprehensive, recent review by Wright & Kennedy

(1981: 112), ‘Sciponoceras is distinguished . . . by its marked constrictions.

These are straight and prorsiradiate in early forms but in later ones are rursi-

radiate on the inner quarter or third of the flank and then turn forward. There

are also well-marked, weakly branching ribs on the body chamber or the whole

Shell, following the same course as the constrictions. The apertural features

appear to be of specific as well as sexual significance; some are relatively

simple, others have a long rostrum, broad folds and a high collar, others have

long lateral lappets.’

Discussion

Extensive discussions on Sciponoceras are given by Matsumoto (1959),

Matsumoto & Obata (1963), Kennedy (1971), Cobban & Scott (1972), Wright

(1979), Wright & Kennedy (1981), Kennedy & Juignet (1983) and Wright &

Kennedy (1995), and nothing new can be added to these.

Occurrence

Transitions from Lechites to Sciponoceras first occur in the Upper Albian

Stoliczkaia dispar Zone faunas of the Anglo-Paris Basin. The genus extends to

the Upper Turonian. The genus has a world-wide distribution (see Matsumoto

CRETACEOUS FAUNAS FROM SOUTH AFRICA 11

1973: 422, fig. 1), including Europe as far east as Transcaspia, the Middle East,

North Africa, Madagascar, Zululand, ?Angola, India, northern Australia, New

Zealand, various localities in North America in the Western Interior, Gulf Coast

and California, Mexico, and in Argentina in South America.

Sciponoceras baculoides (Mantell, 1822)

Fig. 2D-F

1822 Hamites baculoides Mantell, p. 123, pl. 23 (figs 6-7).

1959 Sciponoceras baculoide (Mantell); Matsumoto, p. 104, pl. 31 (fig. la-d), text-

fig. 2a—b. (cum. synon.)

non 1973 Sciponoceras baculoide (Mantell); Henderson, p. 81, text-fig. 4a-d, fig. 6 nos

4a-c, Sa-c, 7a-c.

? 1975 Sciponoceras baculoides (Mantell); Forster, p. 166, pl. 4 (fig. 6), text-fig. 36.

1980 Sciponoceras baculoide (Mantell); Marcinowski, p. 252, pl. 3 (figs 17-20).

1983 Sciponoceras baculoides (Mantell); Kennedy & Juignet, p. 19, figs 11(a)-(y),

12(a)-(bb), 13(a)-(w), 14(a)-(n) (cum. synon.)

1983 Sciponoceras baculoide (Mantell); Marcinowski & Walaszczyk, pl. 1 (fig. 3).

1983 Sciponoceras baculoide (Mantell); Kaplan er al. pl. 5 (fig. 2).

1987 Sciponoceras baculoides (Mantell); Wright & Kennedy, p. 177, pl. 37

(ie 13):

1991 Sciponoceras baculoide (Mantell); Delamette & Kennedy, p. 462, figs 17.6,

he AA ATS.

1992 Sciponoceras baculoide (Mantell); Thomel, pl. 6 (fig. 7), pl. 10 (fig. 2),

pl. 11 (figs 1-3), pl. 19 (fig. 4).

1995 Sciponoceras baculoides (Mantell); Wright & Kennedy, p. 317, pl. 95

(figs 1-3, 5-10), pl. 96 (figs 1-7), pl. 97 (figs 1-5), pl. 98 (figs 29-32),

text-figs 129H, 132R-S, 133A-C, M-FF. (cum. synon.)

Type

Lectotype BMNH 8612, by subsequent designation of Kennedy (1971: 9), is

the larger specimen on the block figured by Mantell (1822, pl. 23 (fig. 6)) from

the lower Middle Cenomanian of Hamsey, Sussex, refigured by Kennedy (1971,

pl. 2 (fig. 5a—b)).

Material

SAS A705 from an unspecified horizon and locality on the Skoenberg, Zulu-

land, Mzinene Formation, Cenomanian.

Description

The specimen is a phragmocone fragment, preserved as an internal mould

in glauconitic silt. Part of a constriction is preserved at the smaller end, and

faint prorsiradiate ribs are visible on the flanks and over the venter. The whorl

section is ovoid, Wb: Wh 10.7: 11.7 (= 0.91).

Discussion

Extensive descriptions and discussions can be found in Kennedy (1971),

Juignet & Kennedy (1976), Kennedy & Juignet (1983) and Wright & Kennedy

(1995).

1 ANNALS OF THE SOUTH AFRICAN MUSEUM

Occurrence

Sciponoceras baculoides has been reported from the Lower to Upper Ceno-

manian, but many of these records are difficult to substantiate. According to

Kennedy & Juignet (1983: 22), the species is abundant only in the lower part of

the Middle Cenomanian in southern England and France. It extends to the lower

Upper Cenomanian. Other records are from Germany, Poland, Romania, Cali-

fornia, North Africa, southern India, Madagascar, Mozambique and Zululand.

Sciponoceras cucullatum Collignon, 1964

Figs 2G-L, N-O, 4-5, 6A-D

? 1936 = Baculites Gaudini Pictet & Campiche; Venzo, p. 118 [60], pl. 10 [6] (fig. 3).

1964 Sciponoceras cucullatum Collignon, p. 38, pl. 326 (fig. 1458).

Type

Holotype by monotypy is the specimen figured by Collignon (1964, pl. 326

(fig. 1458)) from the ‘Lower’ Cenomanian, Zone of Mantelliceras mantelli and

Calycoceras newboldi of gisement 505, west of the falls of Mahaboboka

(Manera), Madagascar; herein refigured in Figure 6A-C.

Material

SAM-PCZ7499, PCZ9137, PCZ9139-9140, NMB D1002, D1002a-b,

D1003a—c, D1004, D1004a-1, SAS H210/23, H30-31, H33-35, SAS A106,

1063-1064, 1069, all from an imprecisely located: horizon on the Skoenberg at

locality 61. Judging by the preservation, the specimens seem to have come from

the higher parts of the section. Mzinene Formation, upper Lower or lower

Middle Cenomanian.

Dimensions

Specimen MxWb MxWh Wb/Wh MnWb MnWh Wb/Wh D Ti _ Ri

D1002a oll Pee OA Sul 6.0 0.85 30 4.0 —

D1002b To) 853) OFS See all O30) 37 3.24 —

H210/34 0. WZ O72 US NOES OL TZ 2) (Re —

D10041 Do) Nabe OT CV eo 0,1 34 3.8 5

BANOS) NOG W2e O93 Bo UE Ole 31) dee —

D1004d Wet NOK OTT 102 Tae 30269 38 4.5 =

D1004b kai, NS O70) 1320) 1a TOS 59 3:05 3

D1003b We NSO) OZ 12:0) 14-3) O90 72, QO. 9)

D1003c Ne WeallSi3) Oso _ — — -~

A1063 ICA TZ) 2-25) 16.2 16.2 1.0 40 2.5 —

D1004 IMs ZZ) Wace) — 21.2 — 83° 220 —

H210/31 OFS) 220) 0286 NSe2 232 Oe, 46." 14 —

D1003a 230 2407 O93 18.5) 214 OES 110 3% 6

Fig. 4 (see facing page). Sciponoceras cucullatum Collignon, 1964. A-C. NMBD1003a.

D. NMBD1003b, both from an imprecisely located horizon on the Skoenberg at locality 61,

Zululand, Mzinene Formation, ?Middle Cenomanian. Arrow points to shallow constriction.

Both x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA

14 ANNALS OF THE SOUTH AFRICAN MUSEUM

Description

Preservation of the material from this locality is characteristic: specimens

are either preserved as internal moulds in a combination of glauconitic silt and

sparry calcite, or with part of the original shell preserved in recrystallized

sparry calcite. Unfortunately, recrystallization of the calcite, plus the fact that

most specimens were found eroded on the surface, has obliterated most of the

surface ornament.

The species reaches large size; the largest Zululand specimen, D1003a

(Fig. 4A—C) is still septate at a whorl height of nearly 25 mm. The Madagascan

holotype (Fig. 6A-C) is adult with a fully developed, recurved, hooded aperture

at Wb = 20.7 mm.

There is considerable variation in whorl shape; it varies from circular, to

high oval, with Wb: Wh ratio as little as 0.67. This variation is partially due to

ontogenetic change: smaller specimens are generally more compressed laterally

than large specimens, which tend to be almost circular in cross section.

Where the shell is preserved, ornament is seen to consist of ribbing that is

distinctly prorsiradiate on the flanks, transverse or feebly convex on the venter,

and also straight and transverse, but weakened on the dorsum. Density of rib-

bing is approximately 5 per whorl height. On internal moulds the surface is

completely smooth. Constrictions are very rare, e.g. D1003a (Fig. 4A—C); these

seem to follow the same direction as the ribs and are strongest over the venter.

As a result of the preservation, only parts of the suture are visible.

Discussion

We have not yet been able to examine Venzo’s (1936) described material,

but the figure and the locality data indicate that it is very probable that his Bacu-

lites gaudini (Venzo 1936: 118 [60], pl. 10 [6] (fig. 3)) belongs to this species,

and not to Lechites gaudini. Because of the rare and weak development of con-

strictions it is quite easy to understand Venzo’s error in referring this species to

Lechites rather than to Sciponoceras.

In retrospect, we suspect that our earlier reference to §S. roto from Zululand

(Kennedy & Klinger 1975: 277) may, in part, have been based on this species.

As is shown below, S. roto lacks the distinctive prorsiradiate ribbing of the

present material and has regular constrictions, thus easily distinguishing it from

S. cucullatum.

As far as the large size is concerned, S. cucullatum is close to S. santa-

crucense Leanza (1970: 212, pl. 11 (figs 1-7)) (Fig. 6E herein), a rather poorly

known species from Santa Cruz Province, Argentina. This species was initially

dated as Cenomanian, but this was based on incorrect stratigraphic information.

Sciponoceras santacrucense occurs below a level with abundant Placenticeras at

Puesto el Alamo (Riccardi & Aguirre Urreta 1988; personal observation), which

would probably date it as late Turonian. Sciponoceras santacrucense has dis-

tinct, regular, albeit widely spaced constrictions that easily distinguish it from

S. cucullatum.

Sciponoceras gracile (Shumard, 1860) (see Kennedy 1988: 108, pl. 20

(figs 1-14, 17-20), text-fig. 38 for the most recent review) from the Upper

Cenomanian, grows to similar large size, but the ornament consists of fine ribs

CRETACEOUS FAUNAS FROM SOUTH AFRICA 15

Fig. 5. Sciponoceras cucullatum Collignon, 1964. A-D. NMBD1004. E-F. NMBD1004c.

G-H. NMBD1002. J-L. NMBD1004d. All from an imprecisely located horizon on the

Skoenberg at locality 61, Zululand, Mzinene Formation, 7Middle Cenomanian. All x 1.

16 ANNALS OF THE SOUTH AFRICAN MUSEUM

Cae

Fig. 6. A-D. Sciponoceras cucullatum Collignon, 1964. Plaster cast of the holotype from

the Lower Cenomanian zone of Mantelliceras mantelli and Calycoceras newboldi of gise-

ment 505, west of the falls of Mahaboboka (Manera), Madagascar. D. NMBD1004 from an

imprecisely located horizon on the Skoenberg at locality 61, Zululand, Mzinene Formation,

?Middle Cenomanian. E. Sciponoceras santacrucense Leanza, 1970. Impression of specimen

from the Turonian of Puesta el Alamo, Santa Cruz Province, Argentina. All x I.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 17

that are strongest on the ventral half of the flanks, and constrictions occur fairly

regularly at distances ranging from | to 1.5 major diameters, according to

Wright & Kennedy (1981: 113).

Sciponoceras kossmati (Nowak, 1908) is difficult to interpret (see Wright &

Kennedy 1981: 114), but it appears that most of the material from California

and Hokkaido, described by Matsumoto (1959: 106, pl. 31 (figs 2a—b, 3), text-

figs 4a—b, 5a—b, 6a-b) and Matsumoto & Obata (1963: 13, pl. 3 (fig. 2), pl. 4

(fig. 1), pl. 5 (figs 1-3), pl. 6 (figs 3-5), text-figs 5-25) as S. kossmati, is indis-

tinguishable from S. gracile.

Sciponoceras orientale Matsumoto & Obata (1963: 18, pl. 3 (fig. 1), pl. 6

(figs 1-2), pl. 7 (figs 1-6), pl. 9 (fig. 6), text-figs 33-49) from the Lower and

Middle Turonian of Hokkaido is also quite large, but is distinguished by the

frequent constrictions and coarse ribbing.

The recurved aperture of the holotype of S. cucullatum is totally unlike any

of the known apertures of other Sciponoceras species.

Occurrence

‘Lower’ Cenomanian of Madagascar, Lower and ?Middle Cenomanian,

Zululand.

Sciponoceras roto Cieslinski, 1959

Fig. 2P

1907 Baculites baculoides Mantell; Pervinquiére, p. 92 (pars), pl. 4 (fig. 8 only),

non pl. 4 (fig. 7), text-fig. 22.

1940 = Cyrtochilus pervinquierei Breistroffer, p. 99 (29). (nomen nudum)

1959 = Sciponoceras roto Cieslinski, p. 39, pl. 4 (fig. 10), text-fig. 14 (II).

1971 = Sciponoceras roto Cieslinski; Kennedy, p. 10, pl. 3 (fig. 7).

1972 Sciponoceras roto Cieslinski; Hancock er al. pl. 81 (fig. 8).

1975 Sciponoceras roto Cieslinski; Kennedy & Klinger, p. 277 (pars).

1979 = Sciponoceras cf. roto Cieslinski; Kennedy er al. p. 10, pl. 1 (fig. 4).

1980 Sciponoceras roto Cieslinski; Marcinowski, p. 254, pl. 3 (figs 14-15).

1985 Sciponoceras roto Cieslinski; Marcinowski & Walaszczyk, pl. 1 (fig. 9).

1991 Sciponoceras roto Cieslinski; Delamette & Kennedy, p. 460, figs 17.8-17.13,

17.16-17.23.

1995 Sciponoceras roto Cieslinski; Wright & Kennedy, p. 315, pl. 94 (figs 13-19),

pl. 95 (fig. 4), pl. 98 (fig. 28), text-figs 131J-L, N. (cum. synon.)

Type

Cieslinski based this species on nine syntypes; as yet, no lectotype has been

designated.

Material

SAS EM191 from an unspecified horizon at locality 181, Zululand, Mzinene

Formation, Cenomanian I-II.

Description

The specimen consists of part of the phragmocone and body chamber. The

whorl section is circular; Wb: Wh 9.5: 9.5 (= 1.0) at the greatest diameter.

18 ANNALS OF THE SOUTH AFRICAN MUSEUM

Apart from two prominent constrictions spaced at about two whorl heights

apart, the surface of the internal mould is nearly smooth. Faint ribs occur on the

venter.

Discussion

Sciponoceras roto is easily identified by the circular whorl section and lack

of ornament between the constrictions. The specimen from Zululand differs

from the European material in having the constrictions slightly closer spaced—

twice the whorl height compared to three times. We do not know if this feature

is significant.

Breistroffer (1940: 99 (29)) erected a new species, Cyrtochilus pervin-

quierei, for one of Pervinquiére’s specimens of Baculites baculoides. As Wright

& Kennedy (1995: 315) have noted, this is a nomen nudum. In any case, the

specimen is close to, if not identical with, S. rofo.

As discussed above, earlier records of S. roto from Zululand (Kennedy &

Klinger 1975: 277) are probably partially based on misidentified S. cucullatum.

Occurrence

Sciponoceras roto 1s rare and known from the Lower Cenomanian of

Poland, southern England, France, Germany, Mangyschlak, Iran, Tunisia,

Madagascar and Zululand.

Genus Baculites Lamarck, 1799

(= Homaloceratites Hupsch, 1768 (non binom.); Euhomaloceras Spath, 1926)

Type species. Baculites vertebralis Lamarck, 1799: 80, by subsequent

designation of Meek (1876: 80).

Diagnosis

Planispiral ammonitella followed by straight or curved shaft; body chamber

may curve in some. Size variable, up to 2 m in length; probably dimorphic.

Ornament variable, from nearly smooth with only growth striae around periph-

ery, to crescentic ribs that are strongest on dorsolateral region, but may be

circumperipheral, to distinct dorsolateral nodes, which may be round, crescentic

or, rarely, longitudinally elongated. In some, the venter may be distinctly corru-

gated, associated either with or without ventrolateral ribbing. Aperture simple,

in line with long axis of shell, short dorsal rostrum, prominent lateral sinus and

straight to slightly curved ventral rostrum; may be flared in some species.

Suture variable, ranging from simple, subquadrate lobes and saddles with open

bases with simple incisions or phylloid folioles, to highly dendritic, with saddles

and lobes with slender bases.

Discussion

Of all the heteromorph ammonite genera, the genus Baculites probably

poses most problems in identification and global stratigraphic correlation. The

great number of monotypical and/or endemic Baculites species that exist in the

CRETACEOUS FAUNAS FROM SOUTH AFRICA 19

literature, as well as gross misidentifications (e.g. B. ovatus, typically from the

Upper Campanian of North America, allegedly from the Coniacian of Vene-

zuela (Reyment 1958) or B. inornatus, typically Campanian but reportedly from

the Lower Coniacian of Venezuela (Renz 1982)), all bear testimony to this

problem.

Reasons for the taxonomic difficulties are due to a combination of various

factors. These include:

(1) Extreme degree of intraspecific variation. This is especially obvious in

ornate species; these may vary from completely smooth to heavily ribbed or

tuberculate (see e.g. B. anceps or B. capensis).

(2) Inornate (smooth) species are virtually impossible to identify unless they

have a very characteristic whorl section and/or suture line and the stratigraphic

position is known.

(3) Apparent endemism of some Baculites species and restriction to distinct

bio(?)geographic regions, 1.e. U.S. Western Interior, Europe and Indo-Pacific

regions, as well as possibly the U.S. Gulf Coast-Atlantic region and North

Africa—Middle East regions.

Details of these problems are to be discussed fully in our forthcoming

synthesis of the family Baculitidae (Klinger & Kennedy in press).

Fortunately, the descriptions and/or revision of the majority of the baculitid

faunas of these major regions are the work of a few persons. Consequently, the

taxonomic criteria within a certain geographic region are more or less constant.

The baculitid faunas of the U.S. Western Interior were dealt with primarily

by Reeside (1927a, 1927b) and especially Cobban (1951 onwards), and recently

with the assistance of Kennedy. The northern extension of the Western Interior

region into West Greenland was covered by Birkelund (1965). In the Indo-

Pacific region, the baculitids of California, Hokkaido and Honshu were mono-

graphed by Matsumoto (1959), Matsumoto & Obata (1963) and Obata & Matsu-

moto (1963), respectively; Ward (1978) revised the baculitids of Washington

State and British Columbia. Those of Madagascar were described virtually

single-handedly by Collignon (1931 to 1971). Major contributions from other

parts of the Indo-Pacific region include New Zealand (Marshall 1926; Hender-

son 1970), Antarctica (Olivero 1984, 1992), the Austral Basin of South America

(Hunicken 1965; Hiinicken et al. 1975, 1980) and Argentina (Leanza 1964,

1967).

In contrast, Baculites is poorly known in the European and Asian regions.

Recent revision of the Upper Cretaceous ammonite faunas of Europe, especially

those of France by Kennedy (1984 onwards) and Kennedy & Summesberger

(1986, 1987), have clarified the taxonomy of most of the Coniacian, Santonian

and Maastrichtian Baculites species. The succession and systematics of the

majority of Baculites species of the Campanian Stage still have to be worked

out.

Distribution of Baculites on either side of the Atlantic is erratic. The oldest

Baculites in Angola is an as yet undescribed Late Turonian—Early Coniacian

species, which is transitional between B. yokoyamai and B. codyensis (M. R.

Cooper collections, S.A. Museum; herein Fig. 131N—R, 132) or B. capensis.

Cooper (1988) recorded B. capensis from the Lower Campanian. Haughton

20 ANNALS OF THE SOUTH AFRICAN MUSEUM

(1925, 1926) and Howarth (1965) described B. subanceps (Figs 130, 131A—H)

from the Upper Campanian or Lower Maastrichtian, and Haas (1943: 13,

figs 15-19) described B. anceps (herein Fig. 131I-M) from the Lower Maas-

trichtian. A single baculite species, B. teicherti is known from the Maastrichtian

of Nigeria.

Baculites from the Atlantic seaboard and the Gulf Coast region of the

U.S.A. (see e.g. Adkins 1929; Stephenson 1941; Cobban 1974; Cobban &

Kennedy 199la, 1991b, 1992a, 1992b, 1993; Kennedy & Cobban) 1993a7

1993b, 1993c, 1993d, 1993e) bear little resemblance to those of Angola or of

the Western Interior.

Fig. 7. A-F. Baculites yokoyamai Tokunaga & Shimizu, 1926. A. NMBD1158/2.

B-C. NMBD1158/3, specimen with distinct ribbing on body chamber. D. NMBD1158/4.

E. NMBD1158/1. F. NMBD1158/7. All from a locality on the Msunduzi River, Van

Hoepen collection, presumably the equivalent of locality 145 of Kennedy & Klinger (1975:

299), St Lucia Formation, Coniacian II. Scale x 1.

The baculitid records from Central America and Brazil are scant. Only

B. yokoyamai is known from the Lower Coniacian of Venezuela and B. kegeli

from the Maastrichtian of Brazil. |

The North African baculitids were last studied by Pervinquiére (1907, 1910)

and, from his descriptions, it appears that Baculites is rare. Lefeld & Uberna

(1992) recently mentioned Baculites in association with Nostoceras from Libya.

The baculitids of the Middle East, specifically Israel, appear to contain a

mixture of European, Indo-Pacific and U.S. Gulf Coast and Atlantic seaboard

CRETACEOUS FAUNAS FROM SOUTH AFRICA 7

elements, as well as some endemics. Dr Z. Lewy (Jerusalem) has shown us an

extensive collection of Campanian to Maastrichtian Baculites, which he intends

to describe. This as yet undescribed collection shows that the older records of

Blanckenhorn (1905), Taubenhaus (1920) and Picard (1929) are badly in need of

revision. Baculitids from Egypt were recently described by Luger & Groschke

(1989) and Hamama & Kassab (1990). These include B. subanceps and

B. ovatus—Indo-Pacific and Gulf Coast-Atlantic region species. The record of

B. scotti, a Western Interior species by Hamama & Kassab (1990: 462, pl. 2

(figs 5-9)) is probably incorrect. Alharithi & Ibrahim (1992) also recorded

B. ovatus from the Maastrichtian of Jordan.

The Upper Coniacian to Lower Maastrichtian Baculites fauna of the U.S.

Western Interior is mainly endemic (see review by Cobban 1994), but Kennedy

(1993) has recently recorded three typical Western Interior species from the

Campanian and Maastrichtian of Belgium. Some parallel development of the

sutures and ornament occurs in the baculites of the Western Interior and of the

Indo-Pacific regions (Klinger & Kennedy in press). Differences between bacu-

lites of the European and Indo-Pacific regions appear to be of lesser magnitude,

and there seems to have been a much greater amount of faunal interchange

between these two regions.

Occurrence

The genus first appears in the Lower Turonian, Zone of Pseudaspidoceras

flexuosum, of the Western Interior of the U.S.A., as B. yokoyamai, and also in

the Lower Turonian of Romania as Baculites aff. undulatus and Baculites sp.

(Szasz 1986); it persists to the end of the Maastrichtian (Birkelund 1979, 1993).

Baculites yokoyamai Tokunaga & Shimizu, 1926

Figs 7-11, 12A-I

1926 Baculites (Lechites) yokoyamai Tokunaga & Shimizu, p. 195, pl. 22

(fig. Sa—b), pl. 26 (fig. 11).

1931 Baculites Besairiei Collignon, p. 37, pl. 5 (figs 6, 6a, 7, 7a, 8, 8a, 9), pl. 9

(fig. 16).

1931 Baculites sulcatus Collignon, non Baily, p. 36, pl. 5 (figs 3, 3a, 4, 4a, 5, 5a,

[ei sa) pl. Oi (fie. 15);

1931 Baculites Roedereri Collignon, p. 38, pl. 5 (figs 10, 10a), pl. 9 (fig. 17).

51 Baculites latelobatus Collignon, p. 38, pl. 5 (igs 11, lla, 12, 12a), pl. 9

(fig. 18).

i 1958 Baculites ovatus Say?; Reyment, p. 7, pl. 1 (figs 1-2), text-figs 1-2.

1959 = Baculites aff. B. yokoyamai Tokunaga & Shimizu; Matsumoto, p. 118, text-

fig. 26.

1963 ~=Baculites yokoyamai Tokunaga & Shimizu; Matsumoto & Obata, p. 30, pl. 8

ie.) 5) aple LO (ies: 1-6). pla (iss 1,-4.2 5).¢pl. 12 (ie. 3), pl. 14

(fig. 4), text-figs 72-87.

1965 = Baculites besairiei Coll.; Collignon, p. 18, pl. 420 (figs 1745-1746).

1972 Baculites cf. B. yokoyamai Tokunaga & Shimizu; Cobban & Scott, p. 48,

pl. 20 (figs 15-21).

1975 Baculites cf. B. yokoyamai Tokunaga & Shimizu; Hattin pl. 8 (figs F, H).

1977 —Baculites cf. B. yokoyamai Tokunaga & Shimizu; Kennedy, p. 271, fig. 17

Cie):

1978 Baculites cf. B. yokoyamai Tokunaga & Shimizu; Hattin & Siemers, text-

ji ae ap as

22 ANNALS OF THE SOUTH AFRICAN MUSEUM

1980 Baculites yokoyamai Tokunaga & Shimizu; Cobban & Hook, p. 13, pl. 4

(figs 9-10).

1982 Baculites inornatus non Meek; Renz, p. 105, pl. 34 (figs 3-4, Sa—b, 6), text-

fig. 80.

1983 Baculites yokoyamai Tokunaga & Shimizu; Cobban & Hook, p. 7, pl. 1

(figs 1-7).

1983 Baculites yokoyamai Tokunaga & Shimizu; Cobban, p. 16, pl. 14 (figs 6-8).

1984 Baculites yokoyamai Tokunaga & Shimizu; Cobban, p. 14, pl. 1 (figs 5-6).

1986 Baculites yokoyamai Tokunaga & Shimizu; Cobban, fig. 3H-I.

1988 Baculites yokoyamai Tokunaga & Shimizu; Kennedy, p. 110, pl. 23

(figs 8-10), text-fig. 29c.

1988 Baculites yokoyamai Tokunaga & Shimizu; Kennedy & Cobban, p. 608, fig. 3

Oy Ss ea sh, NON,

1989 _~—-B. yokoyamai Tokunaga & Shimizu; Kennedy er al. p. 101, fig. 3le-h.

1990 —_ Baculites yokoyamai Tokunaga & Shimizu; Cobban, p. B11, pl. 9 (figs 16-22).

1991a_ Baculites yokoyamai Tokunaga & Shimizu; Kennedy & Cobban, p. 69, pl. 13

(figs 4-10, 17-21, 24-28, 34-37, 41-42), text-fig. 22A.

1992 Baculites yokoyamai Tokunaga & Shimizu; Summesberger, p. 124, pl. 8

(figs 10-11).

Type

Holotype, by monotypy, is the specimen figured by Tokunaga & Shimizu

(1926, pl. 22 (fig. 5a—-b), pl. 26 (fig. 11)) from the lower Futaba Beds in the

upper reaches of the Sakurazawa in Orikiri, Hirono-mura, Fukushima prefec-

ture, north-east Honshu. The type was destroyed by fire during World War II.

A specimen from the collections of the Kyushu University, GK 4580 from the

Bannosawa in Hokkaido, of Coniacian age, zone of Inoceramus uwajimensis,

was to be designated neotype by Matsumoto & Obata (see Matsumoto & Obata

963-73 0=3i))

Material

NMB D1158/1-2, ?3, 4-8, all from a locality on the Msundusi River,

collected by E. C. N. van Hoepen in 1923, presumably equivalent to Kennedy

& Klinger’s (1975: 299) locality 145, St Lucia Formation, Coniacian II

(associated with Forresteria (F.) madagascariensis Collignon, 1965).

_Dimensions

Specimen MxWb MxWh Wb/Wh MnWb MnWh Wb/Wh D Ti

D1158/4 6.4 8.6 0.74 Died 8.1 0.70 20) anes

D1158/7 8.4 UL On 79 9) 0.80 28. 7G

D1158/1 933 10 OWT q-9 10.3 OFT 26° 6

D1158/5 9.3 WAI O77 “= — _ — —

D1158/2 oo 13.0 — _- =

D1158/3 oO NET 0.68 Tod) Oo OO 50. <356

Description

This species is rare in Zululand, and known from one locality only. It is

easily identified by the elliptical whorl section and the very slow rate of taper in

adult specimens. Also, the maximum adult size is small compared to later

species of Baculites in Zululand.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 23

Only one specimen, D1158/1 (Fig. 7E) has faint traces of ribbing over the

venter; all the other specimens are perfectly smooth, both on the shell surface

and on internal moulds.

One specimen, D1158/3 (Fig. 7B-—C)—a body chamber with part of the last

two septa preserved, is identical to the above specimens except that it has

distinct, crescentic lateral ribs, which appear to originate in a curved, dorso-

lateral tubercle. Unfortunately we cannot determine on the basis of this speci-

men whether this ribbing is restricted to the body chamber, or is also present on

the phragmocone.

The suture is partially exposed in D1158/4 (Fig. 8A) and D1158/3

(Fig. 8B), showing very simple, open subquadrate saddles and lobes.

a [Pug vst

Fig. 8. Baculites yokoyamai Tokunaga & Shimizu, 1926. Suture lines.

A. NMBD1158/4. B. NMBD1158/3. Scale bar in mm.

Discussion

Baculites yokoyamai is the best documented species in the early history of

the genus Baculites and appears to have both a wide geographic distribution and

long stratigraphic range. It appears to be the root stock from which the majority

of later Baculites species has evolved.

The name B. yokoyamai has been applied to non-tuberculate baculitids with

elliptical whorl section, primitive suture and of Turonian to Coniacian age. The

Turonian occurrences are best documented in the U.S. Western Interior, where

the species ranges throughout the stage, first appearing in the Lower Turonian

Zone of Pseudaspidoceras flexuosum. It persists into the Lower Coniacian,

Zone of Inoceramus erectus of the Western Interior, from where it has recently

been fully illustrated by Kennedy & Cobban (1991a). In Hokkaido it persists

through the greater part of the Coniacian, although possibly not into the upper-

‘most part, in the Zone of Inoceramus uwajimensis.

24 ANNALS OF THE SOUTH AFRICAN MUSEUM

E U

9) 2mm

E (ee

L G

Fig. 9. Baculites yokoyamai Tokunaga & Shimizu, 1926. A-E. Whorl sections.

A. Japanese specimens (copy of Matsumoto & Obata 1963, text-figs 72-86).

B. NMBD1158/8. C. NMBD1158/6. D. NMBD1158/1. E. NMBD1158/5. All x 1.

F-G. Baculites bailyi Woods, 1906. Suture lines. F. SAM-PCZ8355. G. SAM-PCZ8344.

Our smooth specimens are identical to material described from the U.S.

Western Interior and Hokkaido. Kennedy & Cobban (1991a) have recently

shown that, even though the majority of B. yokoyamai in the lower Coniacian of

the Western Interior are smooth, some forms occur with fold-like ribs on the

upper part of the flanks (e.g. Kennedy & Cobban 1991a, pl. 13 (figs 20, 24).

D1158/3 (Fig. 7B-C) could possibly be interpreted as a more strongly sculp-

tured variant of B. yokoyamai, and we see no need for separating it from the

latter at present. Nevertheless, the similarity in ornament between D1158/3 and

B. sweetgrassensis Cobban (1951: 820, pl. 118 (figs 6-9), text-figs 1-3) (see

also Kennedy & Cobban 199la: 70, pl. 14 (figs 24-25, 29-34, 38-42)) is

CRETACEOUS FAUNAS FROM SOUTH AFRICA 25

striking. This latter species, however, has a distinctive ovoid whorl section with

a narrow venter.

The relationship between ribbed D1158/3 and smooth forms of B. yoko-

yamai is very similar to that of B. sweetgrassensis (ribbed) and B. mariasensis

(smooth) (Cobban 1951: 818, pl. 118 (figs 10-12), text-figs 4-7)—see also

Kennedy & Cobban 1991a: 69, pl. 13 (figs 11-16, 22-23, 29-33, 38-40), pl. 14

(figs 1-23, 26-28, 43-48), text-fig. 25E). The latter species both occur in the

Lower Coniacian Zone of /noceramus erectus, and, according to the original

description of Cobban (1951: 820), ‘B. sweefgrassensis is an uncommon species

. . . . The types were collected with B. mariasensis in the Colorado shale’.

Their co-occurrence, as well as the ratio of smooth to ribbed forms suggests

that B. sweetgrassensis may be no more than a ribbed form of B. mariasensis,

in the same relation as the ribbed Zululand specimen of B. yokoyamai to the

majority of smooth specimens.

Cobban (1951: 817) and Cobban & Reeside (1952) initially identified what is

now known as B. yokoyamai in North America, as Baculites cf. B. besairiei.

Matsumoto (1959: 117) subsequently suggested that B. besairiei Collignon

(1931: 37, pl. 5 (figs 6-9), pl. 9 (fig. 16)) might be a synonym of B. yokoyamai.

Later, Matsumoto & Obata (1963: 34) concluded confidently that ‘all the

described characters of Baculites besairiei Collignon are the same as those of

B. yokoyamai’.

Collignon had 721 baculitid specimens from the apparently condensed or

secondarily concentrated ferruginous conglomerate of Mahagaga, initially dated

as Santonian (Collignon 1931), but later as Late Coniacian (Besairie & Collig-

non 1959). Of these, 531 specimens were discarded as being too fragmentary,

leaving 190 specimens to be studied. The majority, 150 specimens, were

Seremred to 6. Desairiei Collignon (1931: 37, pl. 5 (figs 6, 6a, 7, 7a,.8, 8a, 9),

pl. 9 (fig. 16)); lectotype herein designated, is the largest figured specimen pl. 5

(fig. 6)) (herein Fig. 1OA-C). Collignon (1931: 37) described the variation in

strength of ribbing over the venter in B. besairiei in detail. In typical forms, the

ribs are oblique over the flanks and cross the venter in a sharp chevron. In

some, ribbing is thin, in others thick, and in some alternating thick and thin.

In addition to B. besairiei, Collignon described three other non-tuberculate

baculitids from this assemblage. These include B. sulcatus Collignon, non Baily

(Collignon 1931: 36, pl. 5 (figs 3-5, 13), pl. 9 (fig. 15)) (24 specimens) (herein

Fig. 11D-L); B. roedereri (Collignon 1931: 38, pl. 5 (fig. 10), pl. 9 (fig. 17))

(herein Fig. 12G-I); holotype by monotypy is the figured specimen; B. late-

lobatus (Collignon 1931: 38, pl. 5 (figs 11-12), pl. 9 (fig. 18)) (8 specimens)

(herein Fig. 12A-F); lectotype, herein designated, is the largest of the two

figured specimens (Collignon 1931, pl. 5 (fig. 11)) (herein Fig. 12A-C).

We have been able to examine the figured and some of the unfigured

material and agree with Matsumoto & Obata that B. besairiei is probably a

synonym of B. yokoyamai. Certainly as far as the elliptical whorl section and

variation in ornament are concerned, there is no valid reason for maintaining

them as separate species.

Furthermore, Collignon’s B. sulcatus definitely does not belong to this Cam-

panian Pondoland species; it lacks the typical strong lateral ornament. Instead,

we regard the 24 specimens described by Collignon under that name as slightly

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 10

CRETACEOUS FAUNAS FROM SOUTH AFRICA fa)

more strongly ribbed variants of B. yokoyamai. Externally, B. roedereri is

indistinguishable from B. besairiei, i.e. B. yokoyamai. According to Collignon

(1931: 38) it differs in the suture line: the terminations of the lobes are rounded;

the first (E/L) saddle is very large and divided by a prominent lobule; the

second lateral saddle (L/U) is narrow and much higher than the first; and the

antisiphona! saddle (I/U) is very large and very low, divided by a large, pointed

lobule. The first lateral lobe (L) is narrow and prominent, the second lateral

lobe (U) is smaller than the first and asymmetrical. These differences are trivial

and well within the range of variation of B. yokoyamai (= B. besairiei).

Similarly, B. latelobatus was alleged to differ on sutural details. The anti-

siphonal saddle (U/I) is elongated above the first lateral saddle (E/L), the first

lateral lobe (L) is wide, whereas the second lateral lobe (U) is nearly rudi-

mentary. Again, these differences are well within the limits of variation of

B. yokoyamai (= B. besairiei).

The Mahagaga baculitid assemblage is of extreme interest in that it contains

the earliest representatives of tuberculate Baculites in Madagascar, 1.e.

mevenier Collicnon (1931: 35, pl. S fig. 2, 2a), pl. 9 (fig. 14)) (herein

Fig. 12J-—L) and Baculites cf. B. brevicosta Collignon non Schliter (Collignon

1931: 34, pl. 5 (fig. 1, la), pl. 9 (fig. 13)), but these are numerically insignifi-

cant compared to unornamented B. yokoyamai.

It is difficult to separate B. yokoyamai from other contemporary Turonian

Baculites species. In the Western Interior, the name B. calamus Morrow, 1935

(p. 473, pl. 49 (fig. 8a—b)) is used for Turonian Baculites with weak lateral ribs

(fide Cobban & Scott 1972: 49). However, as shown by e.g. Cobban & Hook

(1983, pl. 1 (figs 3-6)) and recently Kennedy & Cobban (1991a), some speci-

mens of B. yokoyamai also have weak to prominent lateral ornament, and it is

difficult to maintain them as separate species. Compared to B. yokoyamai,

B. calamus is extremely rare. We are not quite sure on how many specimens

Morrow based his species; he only mentioned the holotype but, according to

Cobban & Scott (1972: 49), ‘The species is uncommon; we found it in only one

bed at one locality’ (in Colorado). According to Cobban & Scott (1972: 32,

table 3), B. yokoyamai occurs throughout virtually the whole of the Turonian,

whereas B. calamus occurs only in the Middle Turonian Collignoniceras

woollgari Zone. Rarity alone can not be a criterion for invalidating the species

but, combined with the overlapping morphologies, we doubt that B. calamus

can be separated satisfactorily from B. yokoyamai.

The only other Turonian Baculites species is B. undulatus d’Orbigny, 1850.

Matsumoto & Obata (1963: 28) ascribed authorship of this species to Roman &

Mazeran, who indeed first provided a description of this species and designated

the lectotype (Roman & Mazeran 1913: 11, pl. 4 (figs 6-8)). Although poorly

defined, d’Orbigny’s Prodrome species are valid, and authorship of

Fig. 10 (see facing page). Baculites yokoyamai Tokunaga & Shimizu, 1926. The figured

specimens of B. besairiei Collignon, 1931. A-C. Lectotype, the original of Collignon, pl. 5

(fig. 6). D-F. The original of Collignon, pl. 5 (fig. 9). G-I. The original of Collignon,

pl. 5 (fig. 8). J-L. The original of Collignon, pl. 5 (fig. 7). All from the ferruginous

conglomerate at Mahagaga, Madagascar. All unregistered and housed in the collections in

Dijon. All x 2.

28 ANNALS OF THE SOUTH AFRICAN MUSEUM

ae Gy

Fig. 11. Baculites yokoyamai Tokunaga & Shimizu, 1926. A-C. The original of Collignon

(1931, pl. 5 (fig. 13)) Baculites aff. sulcatus. D-F. The original of Collignon (1931, pl. 5

(fig. 4). G-I. The original of Collignon (1931, pl. 5 (fig. 3)). J-L. The original of Collig-

non (1931, pl. 5 (fig. 5)), all originally referred to Baculites sulcatus. All from the

ferruginous conglomerate at Mahagaga, Madagascar. All unregistered and housed in the

collections in Dijon. All x 2.

B. undulatus rests with d’Orbigny. The lectotype was refigured and described

by Sornay (1955) and Breton & Bavent (1985: 101-102, figs 1-3).

The name B. undulatus has generally been applied to Upper Turonian Bacu-

lites in Western Europe, especially France (d’Orbigny 1850: 19, 21, no. 21;

Roman & Mazeran (1913: 11, pl. 4 (figs 6-8)), and the Chalk Rock of England

Fig. 12 (see facing page). A-I. Baculites yokoyamai Tokunaga & Shimizu, 1926.

A- ‘C. The lectotype of B. latelobatus, the original of Collignon (1931, pl. 5 (fig. 11)).

D-F. A paralectotype of B. latelobatus, the original of Collignon (1931, pl. 5 (fig. 12)).

G-I. The holotype of B. roedereri, the original of Collignon (1931, pl. 5 (fig. 10)).

J-O. Baculites capensis Woods, 1906. J-L. The lectotype of Baculites boulei Collignon, the

original of Collignon (1931, pl. 5 (fig. 2)). M-O. The original of Baculites cf. B. aspero-

anceps of Collignon (1931, pl. 3 (fig. 7)). A-L from the ferruginous conglomerate at

Mahagaga; M-O from Mokotibe-Tsianaloky. All unregistered and housed in the Soc:

in Dijon. A-L x 2; M-O x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA

Fig. 12

30 ANNALS OF THE SOUTH AFRICAN MUSEUM

(Wright 1979: 287, pl. 1 (figs 6-8), pl. 7 (fig. 11)), where it is only known

from a few localities; Fritsch & Schloenbach (1872: 49) recorded numerous

baculitids from several localities in Bohemia that they considered to probably

belong to B. undulatus. Unfortunately, the specimens were nearly all

secondarily compressed, and none were figured; this makes it difficult to

confirm their Bohemian records. That of Jahn (1895: 136, pl. 8 (fig. 8a—c)),

as Baculites n. sp., however, confirms the occurrence of B. undulatus in

Bohemia. Matsumoto & Obata (1963: 28, pl. 8 (fig. 4), pl. 9 (figs 1-5), pl. 11

(figs 2-3), text-figs 62-71) recorded B. undulatus from the Upper Turonian of

Hokkaido.

Kennedy ef al. (1989: 101, fig. 311) recently recorded B. undulatus from the

Upper Turonian Prionocyclus macombi and P. wyomingensis zones of Trans-

Pecos Texas and New Mexico. This figured specimen is very similar to the

Middle Turonian B. calamus, and we suspect that they may be allied, possibly

as a local subspecies or variant of B. undulatus.

Szasz (1986: 120, pl. 1 (figs 1-2)) recorded Baculites aff. B. undulatus from

the Lower Turonian of Romania, accompanied by a questionable baculitid with

ornament very similar to that of B. calamus. If the Romanian material is cor-

rectly identified as B. undulatus, this suggests that B. yokoyamai and

B. undulatus appeared more or less at the same time in Europe and in North

America.

Baculites undulatus differs from B. yokoyamai mainly in having coarser,

less oblique ribbing. Dorsolateral bullae develop in some adult specimens of

both B. undulatus (see e.g. Wright 1979, pl. 7 (fig. 11)) and B. yokoyamai (see

above and NMB D1158/3—Fig. 7B-C), and seem to be of little systematic

value.

It thus appears that differences between B. yokoyamai and B. undulatus are

found mainly in the ribbing—that of the former being more delicate than that of

the latter. In addition, B. undulatus has a narrower stratigraphic distribution,

being restricted mainly to the Upper Turonian, although it might already occur

in the Lower Turonian, whereas B. yokoyamai occurs throughout the Turonian,

all of the Lower Coniacian and probably persists into the Upper Coniacian.

Differences between B. yokoyamai and B. bailyi are discussed below.

Feebly ornamented baculitids recorded from the Coniacian of Venezuela,

and misidentified as B. inornatus Meek (a Campanian species) by Renz (1982:

105, pl. 34 (figs 3-6), text-fig. 80) and as B. ovatus Say? (of Late Campanian

age) by Reyment (1958: 7, pl. 1 (figs 1-2), text-figs 1-2), are good examples of

B. yokoyamai, thus extending the geographic range of the species to South

America.

Occurrence

Baculites yokoyamai is known from the Turonian and Lower Coniacian

of the U.S. Western Interior, Lower Coniacian of Venezuela, and Coniacian of

Hokkaido (though apparently not extending to the uppermost part of the stage),

Middle Coniacian of Zululand, and Upper Coniacian of Madagascar. Dubious

specimens have recently been recorded from the Middle Turonian of Austria

(Summesberger 1992: 124).

CRETACEOUS FAUNAS FROM SOUTH AFRICA 31

a ed

Fig. 13. Baculites bailyi Woods, 1906. A. Copy of the suture line figured by Woods (1906,

pl. 44 (fig. 5)) based on a paratype in the now missing Griesbach collection. B. Copy of

Baily (1855, pl. 11 (fig. 5a-b)) of the holotype of B. bailyi (as B. sulcatus). C-E. The

holotype, BMNH C11372. F-H. SAM-PCO5994a. I-K. SAM-PCO5994b. Both from

dredge samples off the Natal South Coast. All except A x 1. Photographs in C-E by

courtesy of Natural History Museum, London.

32 ANNALS OF THE SOUTH AFRICAN MUSEUM

Baculites bailyi Woods, 1906

Figs 13-23, ?34M-R, 67K-Q, 78A-B

1855 = Baculites sulcatus Baily, p. 457 (pars), pl. 11 (fig. 5a-b only, non Sc).

i 1904 Baculites sp. Etheridge, p. 90, pl. 3 (fig. 24).

1906 Baculites Bailyi Woods, p. 341, pl. 44 (fig. 5).

? 1921 Baculites Bailyi Woods; van Hoepen, p. 18, pl. 3 (figs 9-10).

1921 Baculites bailyi H. Woods; Spath, p. 261.

1922 Baculites bailyi Woods; Spath, p. 146.

y 1930 Baculites Bailyi Woods; Besairie, p. 223, pl. 21 (fig. 7 only, non 6).

1963 ~=Baculites bailyi Woods; Matsumoto & Obata, p. 35, pl. 20 (figs 1-2), pl. 21

(fig. 5), text-figs 88-89, 116-120, 140-142.

1969 ~— Baculites bailyi Woods; Collignon, p. 21, pl. 520 (fig. 2051).

1977 — Baculites bailyi Woods; Klinger & Kennedy, p. 75, fig. 5D.

1978 Baculites bailyi Woods; Ward, p. 1148, pl. 1 (figs 5-7), text-fig. SA-D.

1984 ~=Baculites bailyi Woods; Olivero, p. 57, pl. 1 (figs 1-5), text-fig. la—b.

1985 Baculites bailyi Woods; Klinger, p. 5, fig. 4E-H.

? 1988 Baculites cf. kirki Matsumoto; Riccardi & Aguirre-Urreta, p. C389, pl. 3

(figs 4-8).

Types

Holotype, by original designation of Woods (1906: 342) is the specimen

figured by Baily (1855, pl. 11 (fig. 5a—b)), BMNH C11372 (herein refigured as

Fig. 13C-E), from an unspecified horizon at the type section of the Mzamba

Formation, Mzamba Estuary, Pondoland, Transkei. Probable paratypes are

SAM- 4822 and SAM-13103 (Fig. 67M), both also from an unspecified horizon

at the Mzamba Estuary.

Material

We have more than one hundred specimens from the following localities:

(1) Locality 1, Mzamba Formation, Pondoland, Transkei, Santonian II.

(2) Locality 10, Zululand (locality d of Spath 1921¢ 221-222) 35S aeons

Formation; the locality extends from Coniacian II at the base to probably as

high as Campanian.

(3) Locality 13, Zululand, St Lucia Formation, Coniacian II or HI.

(4) Locality 72, Zululand, St Lucia Formation, Coniacian III.

(5S) Locality 73, Zululand, St Lucia Formation, Coniacian IV (associated with

Peroniceras (Zuluiceras) modestum).

(6) Locality 74, Zululand, St Lucia Formation, the section extends from

Santonian I to Campanian I.

(7) Locality 77, Zululand, St Lucia Formation, Coniacian V.

(8) Locality 78, Zululand, St Lucia Formation, Santonian I-II.

(9) Locality 80, Zululand, St Lucia Formation, Coniacian V.

(10) The top of the section at locality 92, Zululand, St Lucia Formation,

Coniacian III.

Fig. 14 (see facing page). Baculites bailyi Woods, 1906. A-I. SAM-PCZ8391.

J-Q. SAM-PCZ8357. Both from locality 99, Zululand, St Lucia Formation, Santonian I]

or II. A-D, J-M x 1, E-H, N-Q x 2,1 x 4.

CRETACEOUS FAUNAS FROM SOUTH AFRICA

‘Sly

34 ANNALS OF THE SOUTH AFRICAN MUSEUM

(11) Locality 96, Zululand, St Lucia Formation, Coniacian V or Santonian I.

(12) Locality 98, Zululand, St Lucia Formation, Santonian 71.

(13) Locality 99, Zululand, St Lucia Formation, Santonian I or II.

(14) Locality 100, Zululand, St Lucia Formation, Santonian I.

(15) Locality 101, Zululand, St Lucia Formation, Santonian II or III.

(16) Field locality H40, small exposure at pump house on the banks of the

Nyalazi River, on Nyalazi Sugar Estate, 2 km east of Nyalazi River Settlement,

Zululand, St Lucia Formation, Campanian 71.

(17) Field locality H135, loose concretions next to railway line, near Lake View

siding, Zululand, St Lucia Formation, Coniacian ?V.

(18) Dredge samples off the Natal South Coast.

Dimensions

A list of dimensions is given in the appendix.

Max. Wb. (mm) MxWb/MxWh MnWb/MnWh Ti

0-4.9 OF5 0.71 5.3

5.0-9.9 OMS 0.76 5.4

10.0-14.9 0.76 0.75 4.15

15.0-19.9 0.77 0.76 Zot ‘

Description

Woods’s (1906: 341) original description is as follows: ‘Shell increasing in

diameter very slowly. Section oval, compressed, rather narrower on the

siphonal than on the antisiphonal side. A small carina is seen in specimens

which have the shell well-preserved. Lobes and saddles narrow and rather deep

. Ornamentation consists of small, inconspicuous ribs, which bend rapidly

backwards from the siphonal margin; below the middle of ‘the Shell they curve

round and pass forward over the anti-siphonal margin.’

Our material shows that this is essentially an inornate species with ovoid

whorl section, which, in typical specimens, has a venter that is narrower than

the dorsum and may even be acute; but in some, the whorl section may be near-

circular. Lateral ornament consists of fine striae only, which may strengthen on

the ventral half of the flanks and cross the venter with a distinct chevron, and,

in the process, form distinct corrugations, which may be present even on

internal moulds. Irregularly developed, weak dorsolateral nodes occur in a few

specimens.

One of the notable features of B. bailyi (and other baculitids as well) is that

it often occurs in great concentrations in single concretions, sometimes in more

or less parallel arrangement. In some of these concretions, e.g. SAM-PCZ8390

from locality 13, the preservation is exquisite, with original shell material and

protoconchs preserved (Figs 17A-B, 18A-B).

Early whorls with protoconchs are preserved in PCZ8390 (Fig. 17A),

PCZ8390z (Fig. 18A), PCZ8390m (Fig. 18B), PCZ83900, PCZ8390e and

PCZ7213-4. The small diameter of the early coiled section is less than 0.5 mm,

and explains why exposed protoconchs on fracture surfaces of concretions are

CRETACEOUS FAUNAS FROM SOUTH AFRICA 35

Fig. 15. Baculites bailyi Woods, 1906. A-C. SAM-PCZ8359, exact locality unknown,

probably Mkweyane (Umkwelane Hill), Zululand. Note distinct ventral corrugation.

D-F. SAM-PCZ8346. G-I. SAM-PCZ8345. J-L. SAM-PCZ8344 (microconch), all from

locality 99, Zululand, St Lucia Formation, Santonian I or II. All x 1.

ANNALS OF THE SOUTH AFRICAN MUSEUM

Lees,

CRETACEOUS FAUNAS FROM SOUTH AFRICA a7

extremely rare. A slight nick-point in the early shaft is also conspicuous. The

presence and preservation of baculitids with protoconchs probably indicate very

low energy levels in this depositional environment.

The whorl section is elliptical at very early diameters, c. 2 mm, and sub-

sequently becomes ovoid, with a narrow to acute vénter, and maximum whorl

breadth just dorsal of mid-flank (Figs 19B-C, F, 20A-I, 21A-I, 22B-E). The

whorl section is quite variable. In some, the section is distinctly tear-shaped,

with a nearly acute venter, often accentuated by chevron ribbing over, and dis-

tinct corrugation of the venter. In others the venter may remain broadly rounded

to large diameters, resulting in a near-circular whorl section.

Details of the ornament vary considerably, partly due to preservation.

Ornament consists of fine striae which cross the whole periphery of the shell

(Fig. 14J—Q), but are generally strongest on the ventral half of the flanks, where

they curve and are strongly prorsiradiate. In some, the striae may be raised in

sheaves on the dorsal part of the flanks, forming slight, crescentic, incipient

tubercles (Fig. 14A—-H). Over the venter some striae may thicken periodically,

separated by weaker ones, or all may thicken. Both types of ornament produce

distinct crenulation and chevron ribbing over the venter (e.g. Figs 1SA-C,

16I-L, 23D-G, P-R).

In the majority of specimens, ornament on the body chamber and phragmo-

cone is the same. In some, however, irregularly spaced dorsolateral nodes may

appear on the body chamber. In PCZ8350 (Fig. 16A-D), albeit slightly patho-

logic, crescentic ribs appear on the body chamber. On PZC8353 (Fig. 34H-I) a

single pair of tubercles occurs; on PCZ8387 (Fig. 34M-O) two pairs of

distantly-spaced tubercles occur. We are not quite sure if these are, indeed,

atypical forms of B. bailyi or weakly ornamented variants of B. capensis—we

suspect the former.

Unfortunately preserved apertures are rare, but the disparate sizes of some

apparently adult specimens (compare e.g. Figs 15G-I and I15J-L) suggest that

the species may be dimorphic.

The suture lines (Figs 19A, D, 22A) of our specimens vary in details, but

are relatively simple with open saddles and lobes.

Discussion

For several reasons, B. bailyi is difficult to interpret on the basis of the

Pondoland material alone. Of the original type series, only the holotype and two

very poorly preserved paratypes remain. The exact stratigraphic location of the

holotype at the type section of the Mzamba Formation is unknown and, despite

being amongst the first ammonites recorded from Mzamba, B. bailyi is

extremely rare there. Most probably the biggest obstacle in referring smooth

Coniacian and Santonian Baculites from Zululand and Pondoland to B. bailyi, as

we had indeed done earlier (Kennedy & Klinger 1975: 278-279), is the

seemingly atypical suture line of B. bailyi as figured by Woods (1906, pl. 44

Fig. 16 (see facing page). Baculites bailyi Woods, 1906. A-D. SAM-PCZ8350 (note

irregular ornament on body chamber). E-H. SAM-PCZ8349 (arrow points to lateral

tubercle). Both from locality 99, Zululand, St Lucia Formation, Santonian I or II.

I-L. SAM-PCZ8356 from locality 100, Zululand, St Lucia Formation, Santonian I.

All <r

ANNALS OF THE SOUTH AFRICAN MUSEUM

einnteon nan

Fig. 17

CRETACEOUS FAUNAS FROM SOUTH AFRICA 39

(fig. 5)) (herein Fig. 13A) and taken from a specimen in the Griesbach

collection. None of our smooth Baculites from Zululand have as narrow saddles

as the specimen in Woods’s figure.

The Griesbach collection was originally housed in the Hamburg Museum

(Woods 1906: 275). This collection could not be traced and, according to Prof.

C. Spaeth (letter 2.2.1992) was probably destroyed during aerial bombardment

of the Museum in 1943. There is thus no way of ascertaining whether the suture

line figured by Woods was indeed accurate or not. Unfortunately, the holotype,

through being septate throughout, still has the shell preserved. Without

damaging the shell, it is impossible to determine whether it has a suture like that

of the material described here, or of the lost Griesbach specimen.

Woods (1906: 432) also mentioned that ‘Several other specimens were col-

lected by the (Cape of Good Hope Geological) Survey.’ Only two of the Survey

specimens (SAM-13103, 4822) could be traced in the collections of the S.A.

Museum. These are poorly preserved internal moulds and do not show the

sutures.

As mentioned above, B. bailyi is a rarity at Mzamba. All our specimens of

B. bailyi from Mzamba were collected in the basal beds, where it occurs with

B. capensis. Most of these specimens are small but we are reasonably confident

that these are indeed consistently smooth Baculites and not merely smooth forms

of B. capensis. Those specimens from which sutures could be traced, e.g.

PCP6765 (Fig. 19A) and PCP8729 (Fig. 19E), show considerable variation in

sutural details, but none are quite like Woods’s figure.

Apart from our own material, the largest fossil collection from Mzamba to

date was made by Van Hoepen in 1919 and is housed in the Transvaal Museum

(Van Hoepen 1921). Unfortunately, no precise stratigraphic data are available

for this collection; the locality is merely given as Mzamba Estuary. The Van

Hoepen collection is of extreme interest in that it contains a baculitid assem-

blage that we had not encountered at the type section. Six of these baculites

were tentatively referred to B. bailyi by Van Hoepen, “There is, however, still

an element of doubt’ (Van Hoepen 1921: 18). These are all juvenile specimens

with whorl sections varying from circular to elliptical. It is impossible to ident-

ify them any more precisely as Baculites. All the other baculitids were referred

to B. sulcatus by Van Hoepen. Details of these are given below (p. 114). What

is important, however, is that these differ from typical B. sulcatus from the

uppermost beds at Mzamba, and can possibly be regarded as early forms

connecting with B. capensis from the basal beds. Ornament in these specimens

of B. sulcatus varies from individuals with extremely strong lateral nodes

projected forwards over the flanks and over the venter as strong ribs and with

Fig. 17 (see facing page). Baculites bailyi Woods, 1906. A. SAM-PCZ8390.

B. SAM-PCZ8390b. Parts of same concretion showing protoconchs. Both from locality 13,

Zululand, St Lucia Formation, Coniacian II or III. C. SAM-PCZ8039. D-F. SAM-

PCZ8367 from locality 99, Zululand, St Lucia Formation, Santonian I or II. G-H.

SAM-PCZ8390j from locality 13, Zululand. I. SAM-PCZ8020 from locality 101, Zululand,

St Lucia Formation, Santonian II or IJ. (Note weak tubercle.) J-K. SAM-PCZ8364b from

locality 80, Zululand, St Lucia Formation, Coniacian V. Scale bar for size for A and B;

D-F. IEK x I: G-H «2;

40 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 18. Baculites bailyi Woods, 1906. A. SAM-PCZ8390z. Concretion crowded with

juvenile specimens. B. SAM-PCZ8390m. SEM photograph of barrel-shaped protoconch and

early straight shaft. From locality 13, Zululand, St Lucia Formation, Coniacian II or III.

Axes Bel:

CRETACEOUS FAUNAS FROM SOUTH AFRICA 4]

Fig. 19. Baculites bailyi Woods, 1906. Whorl section and suture lines.

A-B. SAM-PCP6765. C-D. SAM-PCP6766. E. SAM-PCP8729.

F. NMBD108. Venter in whorl section pointing downward. Scale bar for size.

numerous ventral intercalatories, to specimens with smooth flanks and only

Suggestions of ventral corrugations. The suture line of one of the latter (smooth

specimens), TM 540m (Fig. 70C) is nearly identical to that figured by Woods of

the specimen of B. bailyi in the (now lost) Griesbach collection. This suggests—

but it is impossible to prove—that the suture line figured by Woods may

possibly have been taken from a smooth form of B. sulcatus.

Three baculitids (Fig. 13F-K) dredged off the Natal South Coast (Klinger

1985) are derived from what are probably offshore equivalents of the Mzamba

Formation. They occur in a faunal assemblage similar to that described from

42 ANNALS OF THE SOUTH AFRICAN MUSEUM

O 10 mm

Fig. 20. Baculites bailyi Woods, 1906. Whorl sections.

A-D. SAM-PCZ8710. E. SAM-PCZ9845. F. SAM-PCZ8356.

G. SAM-PCZ8374. H. SAS H28/3. I. SAM-PCZ9943.

Venter pointing downward. Scale bar for size.

Mzamba by Van Hoepen (1920, 1921). As far as whorl section and lack of

ornament is concerned, they are unmistakably B. bailyi when compared to the

holotype. Unfortunately only parts of the suture are visible in two of the

specimens but the tops of the saddles appear wider than in Woods’s figure.

These rather inconclusive data from the type locality of B. bailyi indicate

that the species is coeval with B. capensis in the Middle Santonian, and may

occur as late as the early Campanian, where it is partly coeval with B. sulcatus.

The suture line figured by Woods does not seem to be characteristic of typical

B. bailyi. In all likelihood, it was either drawn incorrectly, or is from a.smooth

form of B. sulcatus.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 43

This interpretation of B. bailyi seems to verify Matsumoto & Obata’s (1963:

35), Ward’s (1978: 1148) and Olivero’s (1984: 57) records of this species from

the Upper Santonian of Hokkaido, Upper Santonian to Lower Campanian of

British Columbia and Lower Campanian of James.Ross Island, respectively.

Furthermore, none of the figured suture lines of their specimens has narrow

saddles and lobes, as in Woods’s figure. Matsumoto & Obata (1963: 37) also

commented that the sutures of their specimens of B. bailyi differed from the

Pondoland material. The two sutures illustrated by Matsumoto & Obata (1963,

text-figs 88-89) also show minor differences in the width of the individual

elements.

A B

Cc D E

G H :

O 10 mm

Fig. 21. Baculites bailyi Woods, 1906. A. Whorl sections.

A. SAM-PCZ8389. B. SAM-PCZ8346. C. SAM-PCZ8345.

D. SAM-PCZ8373. E. SAM-PCZ8347. F. SAM-PCZ8375.

G. SAM-PCZ8384. H. SAM-PCZ8384. I. SASH135.

Venter pointing downward. Scale bar for size.

mi ANNALS OF THE SOUTH AFRICAN MUSEUM

L |

5 U

€.

O 5 mm

ie ee ee

0 5 mm

EARDAEe Coen aire ees |

Fig. 22. Baculites bailyi Woods, 1906. Suture line and whorl sections.

A-C. SAM-PCZ83901. D-E. SAM-PCZ8390a. Venter in whorl

section pointing downward. Scale bar for size.

Collignon (1969: 21, pl. 520 (fig. 2051)) confirmed the rarity of the species

in Madagascar, where only two specimens are known, from the Lower

Campanian. Earlier records of B. bailyi from the Santonian of the Montagne de

Francais (Madagascar) by Besairie (1930: 223, pl. 21 (figs 6-7)) are probably

partially incorrect. One of these specimens (Besairie’s fig. 6) has distinct

dorsolateral ribbing and definitely does not belong here; the other is probably

B. bailyi.

Fig. 23 (see facing page). Baculites bailyi Woods, 1906. A-C. SAM-PCZ8379.

D-G. SAM-PCZ8354. H-I. SAM-PCZ8353. J-M. SAM-PCZ8355. Note lateral tubercle

in H. N-O. SAM-PCZ12782, all from locality 98, Zululand, St Lucia Formation, Coniacian

2V. P-R. SAM-PCZ8016 from locality 96, Zululand, St Lucia Formation, Coniacian V or

Santonian I. S-U. SAM-PCZ8390 from locality 13, Zululand, St Lucia Formation,

Coniacian II-III. All x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA

Fig. 23

46 ANNALS OF THE SOUTH AFRICAN MUSEUM

In contrast to Pondoland, B. bailyi is locally very common in Zululand,

especially on the north-eastern part of the floodplain of the Hluhluwe River at

Nkundusi, where it occurs in the uppermost Coniacian and/or Lower Santonian.

Elsewhere in Zululand, B. bailyi first occurs in the second or third divisions of

the Coniacian. Initially we thought that these smooth Coniacian specimens of

Baculites were the same as B. besairiei from Madagascar. However, as dis-

cussed above, B. besairiei (including B. roedereri and B. latelobatus) is a

synonym of B. yokoyamai. We admit that it is difficult to separate isolated

specimens of B. yokoyamai, B. bailyi and smooth or juvenile B. capensis;

identification is mainly by association. These three species have overlapping

Stratigraphic ranges and morphologies in the Coniacian, but they generally do

not occur together. The whorl section of B. yokoyamai is predominantly

elliptical, whereas that of B. bailyi is generally ovoid to circular. Completely

smooth specimens of B. capensis are rare, but can generally be distinguished

from B. bailyi by the more elliptical whorl section. Juvenile specimens of some

early forms of B. capensis, where tuberculation only develops in late stages

of growth—e.g. Figure 49J, are difficult to impossible to separate from

B. bailyi.

It is equally difficult or impossible, on external morphology alone, to dis-

tinguish between B. bailyi and Baculites sp. aff. B. rectus (see p. 47). Both are

completely smooth and have similar whorl sections. Suturally, however, they

are totally different. Even small phragmocone fragments of Baculites sp. aff.

B. rectus show the highly complex suture that immediately distinguishes it from

B. bailyi as figured by Woods (1906), Matsumoto & Obata (1963), Ward (1978)

or Olivero (1984).

Baculites uedae Matsumoto & Obata (1963: 40, pl. 20 (figs 5-7), pl. 21

(figs 1, 3, 6), text-figs 91-92, 121-129), from the Santonian of Hokkaido,

appears indistinguishable from our Zululand B. bailyi. It is virtually identical to

some Santonian microconchs of B. bailyi (see e.g. Fig. 1SJ-L) from Zululand

and we suspect that they may be synonyms. According to Matsumoto & Obata

(1963: 43), B. bailyi and B. uedae are nearly contemporary in Hokkaido but, in

all cases except one, occur at different localities. This seems to fit the general

pattern of Baculites distribution in Zululand, where individual variants of the

Same species are generally clustered at different localities but at the same

stratigraphic level.

Baculites kirki Matsumoto (1959: 143, pl. 43 (figs 1-3), text-figs 53a—b,

54-57, 58a—-b) from the Santonian of California and possibly Hokkaido (Matsu-

moto & Obata 1963: 65, pl. 18 (fig. 2), text-fig. 114) resembles some of our

specimens of B. bailyi with an acute venter, but in that species a distinct

rounded ventral keel develops. However, the specimens figured by Riccardi &

Aguirre Urreta (1988, pl. 3 (figs 4-8)) as Baculites cf. B. kirki from the San-

tonian of Cerro Indice in Patagonia are indistinguishable from some of our

B. bailyi with very narrow venters, e.g. NMBD1075/1, and are probably con-

specific, as are probably the Santonian specimens from James Ross Island,

figured by Olivero (1992, pl. 1 (figs 1-3)).

Baculites fuchsi Redtenbacher (1873: 134, pl. 30 (fig. 15); see also Summes-

berger 1979: 113, pl. 1 (figs 2-4), text-figs 2-3; Immel er al. 1982: 28, pl. 11

(fig. 8)), from the Santonian of the Gosau Beds of Austria, is indistinguishable

CRETACEOUS FAUNAS FROM SOUTH AFRICA 47

from B. bailyi. Unfortunately, the Austrian species is too poorly known for

definite comments. However, the association of smooth B. fuchsi with nodose

B. incurvatus in the Gosau Beds is remarkably similar to the Pondoland associ-

ation of B. bailyi with B. capensis. Given more material, it is possible that these

European species may prove to be senior synonyms of B. bailyi and B. capensis

(see also p. 92 onwards).

Baculites nugssuaquensis Birkelund (1965: 48, pl. 4 (fig. 1), pl. 5

(figs 1-4), pl. 6 (figs 1-2), text-figs 35-41), from the Santonian of West Green-

land, is a homoeomorph of B. bailyi. Some specimens of B. nugssuaquensis

with smooth flanks and ventral corrugations (e.g. Birkelund 1965, pl. 6 (fig. 2))

are indistinguishable from similarly ornamented B. bailyi, e.g. PCZ8359

(Fig. 1SA-C). This species, however, belongs to the northern extension of the

U.S. Western Interior baculitid lineage and any resemblance is due only to

convergence.

Occurrence

Middle Santonian to Lower Campanian of Pondoland, offshore Natal Coast,

Coniacian IJ to Campanian I of Zululand, Lower Campanian of Madagascar,

James Ross Island, Antarctica, Upper Santonian of Hokkaido and Upper

Santonian to Lower Campanian of British Columbia.

Baculites sp. aff. B. rectus Marshall, 1926

Figs 24-26

Compare

1926 Baculites rectus Marshall, p. 154, pl. 19 (fig. 1), pl. 32 (figs 9-10).

1953 Baculites aff. rectus Marshall; Spath, p. 19, pl. 7 (fig. 2a—c).

1970 Baculites rectus Marshall; Henderson, p. 23, pl. 3 (figs 2-3), text-

fig. 6.

1984 Baculites rectus Marshall; Olivero, p. 64, pl. 1 (figs 6-9), text-

nes lc, 2.

Material

SAM-PC5068, glauconitic concretion with numerous baculitid fragments,

labelled ‘Sugar Terminus Foundations, Durban—Donated’. This locality is

probably the same as ‘Maydon Wharf Sugar Terminal Site’ referred to by

Kennedy ef al. (1973) and later (Kennedy & Klinger 1975: 282) as locality 5 of

the Mzamba Formation, and dated as Santonian III and Campanian 7]; several

horizons are clearly represented (Kennedy & Klinger 1975: 282).

Description

The concretion contains numerous baculitids with whorl heights ranging

from c. 2 mm to more than 20 mm. The whorl section (Fig. 25E-F) is typically

ovoid, in some with a very narrow venter. Different sizes of body chambers

Suggest dimorphism. The aperture is slightly flared. The body chambers are all

smooth.

48 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 24. Baculites sp. aff. B. rectus Marshall, 1926. A-B. Parts of same concretion,

SAM-PC5068 from Maydon Wharf, Sugar Terminal site, locality 5, Mzamba Formation,

imprecisely dated as Santonian I]J-Campanian I. Both x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 49

O mm 5

Fig. 25. Baculites sp. aff. B. rectus Marshall, 1926. A-B. SAM-PC5068b.

C-D. SAM-PC5068a. A-B x 2; C-D x 1. E-F. Whorl sections. Venter pointing upward.

Scale bar for size.

The majority of specimens are smooth, but several phragmocone fragments

have low, rib-like swellings on the dorsal half of the flanks, and distinct,

prorsiradiate sulci on the ventral half.

The suture is very complex, dendritic, with saddles and lobes with narrow

stems and some phylloid folioles (Fig. 26).

Discussion

We initially thought that this was a concretion with B. bailyi, but the

complex suture line immediately rules out this identification. Even small

specimens can easily be distinguished as being totally different from B. bailyi on

the basis of the sutural complexity. Unfortunately, we do not have a definite age

50 ANNALS OF THE SOUTH AFRICAN MUSEUM

E L U i

O 5

eee

Fig. 26. Baculites sp. aff. B. rectus Marshall, 1926.

Suture line of SAM-PC5068. Scale bar for size.

for these specimens but we assume them to be younger than the highest beds

exposed at the Mzamba Cliff, above the level of B. sulcatus. The association of

an inflated pachydiscid nucleus possibly confirms this. At any rate, the degree of

complexity of the suture line is greater than that of any of the other smooth

baculitids yet known from southern Africa. The origins of this species are —

obscure. The similar whorl sections suggest derivation from B. bailyi,

Fig. 27 (see facing page). Baculites capensis Woods, 1906. A-C. SAM-4825b,

paralectotype. D-H. SAM-4823, lectotype. I-K. SAM-PCP8753. L. SAM-PCP8050.

All from the basal beds at locality 1, Pondoland, Transkei, Mzamba Formation, Santonian II.

All’ > +1’.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 51

are case ee

Sy ANNALS OF THE SOUTH AFRICAN MUSEUM

associated with a sudden increase in sutural complexity. Alternatively, it could

be regarded as an immigrant.

There are several smooth baculitid species with complex sutures with which

our specimens can be compared.

Closest affinity seems to be with B. rectus Marshall (1926: 154, pl. 19

(fig. 1), pl. 32 (figs 9-10)), best known, but imprecisely dated, as Campanian-

Maastrichtian from New Zealand (see also Henderson 1970: 23, pl. 3 (figs 2-

3), text-fig. 6). Spath (1953: 19, pl. 7 (fig. 2a-e)) also tentatively recorded the

species from Graham Land, Antarctica. Olivero (1984: 64, pl. 1 (figs 6-9), text-

figs lc, 2) also reported the species from the Campanian of James Ross Island,

Antarctica. Collignon’s (1977: 10) Baculites sp. indet. from the Campanian of

New Caledonia may also belong here. None of these specimens has quite as

complex a suture line as the Durban specimens—that figured by Olivero (1984,

text-fig. 2) is the closest match. The suture figured by Spath (1953, pl. 7

(fig. 3)) has very narrow saddles, and resembles that of B. bailyi as figured by

Woods (1906, pl. 44 (fig. 5)), but is more finely incised.

Baculites chicoensis Trask (1856: 85, pl. 2 (fig. 2)) from the Upper Cam-

panian of California and British Columbia, and possibly southern Saghalin (Gra-

bovskaya 1984) has a comparable complex suture line, but the whorl section is

different—an incipient ventral keel appears in later growth stages. Baculites

chicoensis yezoensis Matsumoto & Miyauchi (1984: 70, pl. 25 (figs 1-5), text-

figs 11B-C) from the Lower Campanian of Hokkaido is closer to the Durban

specimens in retaining a narrow but not perceptibly keeled venter, as in the

nominate subspecies. : :

Baculites rex Anderson (1958: 191, pl. 49 (fig. 2)) from the Upper Cam-

panian and Maastrichtian of California, British Columbia and Hokkaido has a

more ‘lytoceratine’ or jagged suture and apparently grows to enormous size, as

the name implies.

Baculites hochstetteri Liebus (1902: 119, pl. 6 (figs 4-6)), imprecisely dated

as Upper Senonian of the Silesian Carpathian region, has sutures indistinguish-

able from our specimens. Unfortunately, this species is known only by the

figured syntypes and further comparisons are impossible. The types, originally

housed in Munich are lost, presumably destroyed during World War II.

Baculites regina Obata & Matsumoto (1963: 85, pl. 22 (figs 3-6), pl. 23

(figs 1-2), pl. 24 Cfigs 1-5), pl. 25 (figs 3-5), pl. 27 (figs 1, 6-7, 9), text-

figs 191-196, 200-214) from the Campanian of Honshu, has a similarly com-

plex suture line, but the adult, subpentagonal whorl section is immediately

distinctive.

Baculites duharti Hiinicken (in Hiinicken et al. 1975: 116, pl. 1 (figs 1-4),

pl. 2 (figs 1-2), pl. 3 (figs 5-8), text-figs 2a-d, 3a-c, 4-5)), from the Middle

and/or Upper Campanian of Tierra del Fuego, and also from the Middle

Campanian of Zululand (p. 178), is superficially similar to Baculites sp. aff.

B. rectus, but has a much simpler suture line, and the elliptical whorl section

typical of the B. capensis group.

Occurrence

Campanian s.1. of Durban.

1906

1907

1921

1971

1921

1922

1923

? 1925

? 1930

1931

1932

1936

1958

1959

1963

1966

non 1973

1977

?non 1988

1991b

Types

CRETACEOUS FAUNAS FROM SOUTH AFRICA 39

Baculites capensis Woods, 1906

Figs 27-33, 34A-L, ?34M-R, 35-54

Baculites capensis Woods, p. 342, pl. 44 (figs 6-7).

Baculites vagina Forbes; Boule, Lemoine & Thevenin, p. 65(45), pl. 15

(fig. 3-3a).

Baculites sp. Crick, p. 240.

Baculites capensis H. Woods; Spath, p. 257, pl. 24 (figs 6-7).

Baculites sp. aff. capensis H. Woods; Spath, p. 258.

Baculites cf. aspero-anceps Lasswitz; Spath, p. 259, pl. 24 (figs 4, 4a).

Baculites cf. brevicosta Schluter; Spath, p. 260, pl. 24 (figs 5, 5a).

Baculites sp. cf. sulcatus Baily; Spath, p. 260.

Baculites capensis Woods; Spath, p. 146.

Baculites capensis Woods; Spath, p. 13, text-fig. 3d.

Baculites sp. ind. Spath, p. 31, pl. 1 (fig. 1).

Baculites Bailyi Woods; Besairie, p. 223 (pars), pl. 21 (fig. 6) only.

Baculites cf. aspero-anceps Lasswitz; Collignon, p. 22, pl. 3 (figs 7, 7a),

pl. 9 (fig. 12).

Baculites aff. capensis Woods; Collignon, p. 22, pl. 3 (fig. 6).

Baculites cf. brevicosta Schluter; Collignon, p. 34, pl. 5 (fig. 1, la), pl. 9

(fig. 13).

Baculites Boulei Collignon, p. 35, pl. 5 (fig. 2, 2a), pl. 9 (fig. 14).

Baculites capensis Woods; Besairie, p. 50.

Baculites capensis Woods; Venzo, p. 116 [58].

Baculites capensis Woods var. umsinenensis Venzo, p. 116 [58], pl. 10 [6]

(figs 11-12).

Baculites buttensis Anderson, p. 191, pl. 49 (fig. 6, 6a, 6b).

Baculites aff. B. capensis Woods; Anderson, p. 192, pl. 48 (fig. 8, 8a).

Baculites schencki Matsumoto, p. 113, pl. 32 (figs la-c, 2a-c, 3a-b, 4a-b,

S5a-c, 6a-c), text-figs 12a—-b, 13a—-c, 14a-b, 15-21, 22a-b, 23a-c, 24-25.

Baculites boulei Collignon; Matsumoto, p. 118, pl. 32 (fig. 7a-c), pl. 33

(figs 4a-c, 5a—b, 6a-d, 7a—-b), text-figs 27a—b, 28-32.

Baculites capensis Woods; Matsumoto, p. 121, pl. 33 (figs la-d, 2a—-c, 3a-b),

pl. 45 (figs la-d, 2a-d, 3a-d, 4a-d), text-figs 33a—b, 34a-b.

Baculites capensis Woods; Matsumoto & Obata, p. 47, pl. 14 (fig. 2), pl. 15

(figs 3-5), pl. 19 (fig. 2), text-figs 95-96, 147-151.

Baculites capensis Woods; Collignon, p. 6, pl. 457 (fig. 1862).

Baculites capensis Woods var. tenuetuberculata Collignon, p. 6, pl. 457

(figs 1863).

Baculites malagasyensis Collignon, p. 7, pl. 457 (fig. 1865).

Baculites sp. group of B. capensis Woods; Kennedy & Klinger, p. 101, pl. 4

(figs 1-5), pl. 5 (fig. la-d), pl. 6 (figs 4-5). (= B. vanhoepeni)

Baculites capensis Woods; Klinger & Kennedy, p. 71, figs 2A-F, 3G.

Baculites capensis Woods; Cooper, p. 210, fig. 1G-I.

Baculites capensis Woods; Kennedy & Cobban, p. 182, figs 6: 4,; 8: 1-8; 10:

7-10. 1D=14=512: 25.

Woods based this species on at least four syntypes, which survive in the col-

lections of the South African Museum. Lectotype, by subsequent designation of

Matsumoto & Obata (1963: 48), is the smaller of the two specimens figured by

Woods (1906, pl. 44 (fig. 6a—b)), from an unknown horizon at the type section

of the Mzamba Formation at the Mzamba Estuary, Transkei, South Africa,

SAM-4823—herein refigured as Figure 27D-H. Paralectotypes are SAM-4824,

4825 and 4825b, all from the same locality.

54 ANNALS OF THE SOUTH AFRICAN MUSEUM

Material

We have several hundred catalogued and uncatalogued specimens from the

following localities:

(1) Basal beds and foreshore exposures at locality 1 at the Mzamba River

Estuary, Pondoland, Mzamba Formation, Santonian II.

(2) Locality 6, Zululand, St Lucia Formation, Santonian II-III to Campanian I.

(3) Locality 15, Zululand, St Lucia Formation, Coniacian IV.

(4) Locality 16, Zululand, Coniacian ?III.

(5) Bed B at locality 22, Zululand, St Lucia Formation, Coniacian IV.

(6) Locality 24, Zululand,St Lucia Formation, Coniacian II-V.

(7) Locality 26, Zululand, St Lucia Formation, ?Santonian.

(8) Locality 72, Zululand, St Lucia Formation, Coniacian III.

(9) Locality 73, Zululand, St Lucia Formation, Coniacian IV-V, ?Santonian I.

(10) Locality 80, Zululand, St Lucia Formation, Coniacian V.

(11) Locality 83, Zululand, St Lucia Formation, Coniacian IV.

(12) Locality 85, Zululand, St Lucia Formation, Santonian I.

(13) Locality 88, Zululand, St Lucia Formation, Coniacian IV-V, ?Santonian I.

(14) Localities 89 and 90, Zululand, St Lucia Formation, Coniacian IV.

(15) Locality 91, Zululand, St Lucia Formation, Coniacian IV or V.

(16) Locality 92, Zululand, St Lucia Formation, Coniacian II and III.

(17) Locality 93, Zululand, St Lucia Formation, Coniacian II.

(18) Locality 94, Zululand, St Lucia Formation, Coniacian V—Santonian I.

(19) Locality 98, Zululand, St Lucia Formation, Coniacian ?V.

(20) Field locality H51, Mpisene Creek, east of Nyalazi River Trading Store,

Zululand, St Lucia Formation, ?Santonian.

(21) Field locality H40, exposure at pump house on Nyalazi River, west of

Nyalazi River Trading Store, Zululand, St Lucia Formation, Campanian 71.

Dimensions

A full list of dimensions is given in the appendix.

Max Wb (mm) MxWb/MxWh MnWb/MnWh Ti Tubs/Wh

0-4.9 0.63 0.6 ae) 2

5-9.9 O83 0.74 6.48 Zon

10-14.9 0.73 Ol 3292 pS

IS='9)9) 0.75 0.75 5.64 2.18

20-21 OMT O:73 — 2

Fig. 28 (see facing page). Baculites capensis Woods, 1906. A. Plaster cast of exposed

surface of basal beds of Mzamba Formation on south side of the Mzamba River estuary.

B-E. SAM-4824, paralectotype figured by Woods (1906, pl. 44 (fig. 7a-c)).

F. SAM-4825, unfigured paralectotype. All from the basal beds of the Mzamba Formation

at locality 1, Pondoland, Transkei, Santonian II. All x 1.

Fig. 29 (see overleaf). Baculites capensis Woods, 1906. All specimens from the basal beds

of the Mzamba Formation at locality 1, Pondoland, Transkei, to illustrate the variation

in ornament. A-C. SAM-PCP8664. D. SAM-PCP8665. E. SAM-PCP8755.

F-H. SAM-PCP8663. I. SAM-PCP8360. J. SAM-PCP8052. K. SAM-PCP8053.

L. SAM-PCP8051. M. SAM-PCP8054. N-P. SAM-PCP8241. All x 1:

CRETACEOUS FAUNAS FROM SOUTH AFRICA

Fig. 28

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 29

CRETACEOUS FAUNAS FROM SOUTH AFRICA 57

DU.

Clete

OOO

Fig. 30. Baculites capensis Woods, 1906. Whorl sections of type

series and topotype material. A-B. SAM-4824. C-D. SAM-4823

Gectotype). EF. SAM -—PCP8754. F. SAM-PCP8756.

G. SAM-PCP8662. H. SAM-PCP8665. Venter pointing downward.

Scale bar for size.

Description

This is the most common baculitid in Zululand, Natal and Pondoland, and is

very variable, as here interpreted. In order to illustrate the variation of the

Species, we first describe the types and topotype material collected at the type

locality, the Mzamba River estuary in Pondoland.

58 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 31. Baculites capensis Woods, 1906. Whorl sections of specimens

from the type locality. A. SAM-PCP8360/10. B. SAM-PCP8360/13.

C. SAM-PCP8052. D. SAM-PCP8241. E. SAM-PCP8051.

F. SAM-PCP8644. G. SAM-PCP8663. H. SAM-PCP8663.

I. SAM-PCP6767. Venter pointing downward: Scale bar for size.

Description of type specimens. The lectotype, SAM-4823 (Fig. 27D-—H) is

part of a phragmocone, partially filled with sparry calcite and glauconitic silt.

SAM-4824 (Fig. 28B-E) is non-septate, and, at the larger end, has part of the

aperture preserved. SAM-4825 (Fig. 28F) is a fragment of which the one flank

is corroded. SAM-4825b (Fig. 27A-C) is a large body chamber and, to date,

the largest specimen of B. capensis recorded from the Mzamba Formation.

Woods’s figures of the lectotype and paralectotype are quite accurate and

show the characteristic features of the species—an elliptical whorl section with

CRETACEOUS FAUNAS FROM SOUTH AFRICA 59

the venter as wide as, or only slightly narrower than the dorsum, and parallel

flanks with a shallow, longitudinal depression situated immediately ventral of

the tubercles. This depression is very shallow, as different angles of illumination

of the lectotype illustrate (Fig. 27D-E). In the lectotype (SAM-4823) and para-

lectotypes (SAM-4824, 4825) the tubercles are characteristically pinched and

clavate.

Paralectotype SAM-4825b (Fig. 27A-C) is part of an internal mould of a

body chamber. As mentioned above, it is the largest specimen of B. capensis

known to us from the Mzamba Formation and is obviously a macroconch. Here

the tubercles are much weaker than on the lectotype and conical to bullate,

rather than clavate. The longitudinal depression at midflank is not as prominent

as in the other specimens. Distinct, forwardly projected ribs are visible on the

venter (on the internal mould).

Decriptions of topotype Pondoland material. We have several specimens of

B. capensis, all from the basal beds of the Mzamba Formation, at Mzamba

Chiff.

Juveniles. The juveniles are fortuitously preserved in a small concretion

(SAM-PCP8360) containing about 20 individuals (Fig. 291) and the impressions

of others. The smallest diameter preserved is 1.7 mm. The whorl section in the

early stages is oval (Fig. 31A-B), with the venter as wide, or only slightly

narrower than the dorsum. The appearance of tubercles is variable. On

PCP8360/10 (Fig. 291), an internal mould, the surface of the flanks is perfectly

smooth up to a whorl height of about 5 mm, when weak, crescentic dorsolateral

tubercles start appearing. Faint ribbing over the venter starts appearing at a

slightly earlier stage. In other, larger specimens, the flanks are still smooth,

ornamented by faint striae only. In PCP8360/3 the venter is distinctly crenulated

on the internal mould and the flanks bear low, rounded tubercles.

Adult stage. The whorl section shows some variation, depending on whether

seen in nodal or internodal view, and also whether taken on an internal mould

or on the external surface of the shell (Figs 30-31). They are generally near-

elliptical, with parallel flanks and the venter only slightly narrower than the dor-

sum. In some, however, e.g. PC P8644 (Fig. 31F), the whorl section is trigonal,

with a narrowly rounded venter. In others, the venter may be slightly fastigiate,

e.g. SAM-4823 (Fig. 27H).

The size, shape and spacing of the tubercles varies considerably; again this

is partially determined by whether the shell is preserved, or whether seen on the

internal mould. Typical ornament associated with the name B. capensis consists

of longitudinally elongated, either oblongly rounded or dorsoventrally pinched

tubercles, situated near the dorsal quarter of the flanks, e.g. lectotype

SAM-4823 (Fig. 27D-E), paralectotype SAM-4824 (Fig. 28B-C)—this shows

the tuberculation on the internal mould and in shelly preservation, and

Figure 28A—a plaster cast of a large phragmocone. These longitudinally

elongated tubercles are generally bordered at mid-flank by a very shallow,

longitudinal depression. This depression is hardly visible in transverse (whorl

section) view, but is quite clear under oblique, low lighting. Compare, for

example, Figure 27D-E of the same (lectotype) specimen under different

60 ANNALS OF THE SOUTH AFRICAN MUSEUM

illumination. In typical B. capensis, the spacing of tubercles numbers about two

per whorl height.

In some specimens, the tubercles are distinctly conical to rounded, e.g.

SAM-8753 (Fig. 27I-K) or PCP8050 (Fig. 27L), and in others low and

rounded, e.g. PCP8052 (Fig. 29J) and PCP8665 (Fig. 29D). In a few, e.g.

PCP8662 or PCP8241 (Fig. 29N-P), the tubercles are extremely feebly

developed, so as to be near-absent. In PCP8054 (Fig. 29M) the tubercles are

closer and irregularly spaced.

The surface of the shell bears fine striae, e.g. PCP8050 (Fig. 27L), and

PCP8052 (Fig. 29J). In some, e.g. PCP8664 (Fig. 29A-C), these are quite

strongly developed, and even visible on internal moulds, e.g. SAM-4825b

(Fig. 27A-C). Ventral corrugations are variably developed, ranging from absent

to extremely prominent, even on internal moulds.

Parts of the aperture are preserved in paralectotype SAM-4824 (Fig. 28C);

paralectotype SAM-4825b (Fig. 27A-C)) is a larger body chamber; both are

perfectly straight. PCP8050 (Fig. 27L) is a smaller, slightly curved body

chamber. The former two are probably macroconchs; the latter a microconch.

Description of Zululand material. In Zululand, B. capensis may be defined

aS a predominantly nodose baculitid with elliptical to ovoid whorl section; size

and shape of nodes varies, from absent, through weakly conical to strongly

conical, weakly to strongly crescentic, longitudinally elongated, obliquely

elongated, close, widely or irregularly spaced; some with shallow longitudinal

depression at midflank. Suture simple.

As can be deduced from this definition, the material is extremely variable

(see e.g. Fig. 33) and several morphotypes, based primarily on the presence and

shape of the tubercles, can be recognized. Some of these morphotypes appear to

be related to stratigraphic occurrence—others are conspicuous at particular

Fig. 32 (see facing page). Baculites capensis Woods, 1906. Two specimens to illustrate

extremes of variation in ornament. A-C. NMBD1052/1 from locality 72, Zululand, St

Lucia Formation, Coniacian III. D-F. SAS A2101 from locality 73, Zululand, St Lucia

Formation, Coniacian IV-V. Both x 1.

Fig. 33 (see overleaf). Baculites capensis Woods, 1906. Diverse specimens to illustrate

variation in strength and shape of tuberculation. A-C. SAM-PCZ11999 from locality 91,

Zululand, St Lucia Formation, Coniacian IV or V. D. SAM-PCZ12000. E. SAM-

PCZ8676, both from locality 83, Zululand, St Lucia Formation, Coniacian TV. F. SAM-—

PCZ12001 from locality 85, Zululand, St Lucia Formation, Santonian I. G. SAM-4832

from locality 1, Pondoland, Mzamba Formation, Santonian II. H. NMBD1028/3.

I. SAS A361. J. SAS A333. L. SAM-PCZ8013, all from locality 72, Zululand, St Lucia

Formation, Coniacian II]. K. SAS Z1795f. M. SAS Z1795g. Both from locality 85.

All <i

Fig. 34 (see overleaf). A-L. Baculites capensis Woods, 1906. A-C. SAM-5454.

D-F. SAM-5480. G. SAM-5458. H. SAM-5486. I. SAM-5461. J. SAM-5472.

K. SAM-PCZ8674b. L. SAS A1610a. All from Mkweyane (‘Umkwelane Hill’), probably

locality 10, Zululand, St Lucia Formation. All specimens with ‘brevicosta’ or ‘schencki’

type of ornament. M-R. Two specimens with atypical, widely spaced tubercles. It 1S

uncertain if these are atypical B. capensis (form 12) or nodose variants of B. bailyi.

M-O. SAM-PCZ8387. P-R. SAS PCZ8349. Both from locality 98, Zululand, St Lucia

Formation, Coniacian V, associated with typical B. bailyi fauna. All x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA

%E Sly

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 33

CRETACEOUS FAUNAS FROM SOUTH AFRICA

Fig. 34

64 ANNALS OF THE SOUTH AFRICAN MUSEUM

localities. These morphological types are merely convenient reference points on

which to base the descriptions of this material and have no taxonomic status

whatsoever. Some have, in the past, been given varietal or specific names, and

we try as far as possible to identify our morphotypes with these names. Their

stratigraphic and phylogenetic relationships are discussed below.

Form | (= Baculites sp. aff. B. capensis Spath, 1921: 258). Predominantly

smooth to feebly ornamented forms. These have the whorl section of typical

B. capensis, but lack lateral ornament, e.g. NMBD1052/1 (Fig. 32A-C). In

some, e.g. SAM-5467 (Fig. 48J), ornament consists of ventral corrugations

only (= Baculites sp. cf. sulcatus of Spath 1921: 260). In others, e.g.

SAM-PCZ8760 (Fig. 421), NMB D1124 (Fig. 42J) and SAM-PCZ8027

(Fig. 42K-M), feeble lateral ornament may occur, as far as can be seen, on the

body chamber only. Specimens reach maturity at very different sizes, e.g.

SAM-5484c at Wh = 10 mm and PCZ7210 at Wh = 20 mm.

Form 2 (= Baculites cf. brevicosta Spath, 1921: 260, pl. 24 (fig. 5, 5a));

(= B. schencki Matsumoto, 1959: 113, pl. 32 (figs la-c, 2a—c, 3a—b, 4a—b,

Sa-c, 6a-c), text-figs 12a—b, 13a—-c, 14a—b, 15-21, 22a—b, 23a—c, 24-25). Small

forms with closely spaced, crescentic tubercles situated near the dorsolateral

edge of the flanks and ovoid whorl section (Fig. 34I-L, 36M-N, 42E). The

tubercles may be narrow and pinched, or broad and low. Dorsally and ventrally

the tubercles fade into fine striae. Whorl section generally ovoid.

Form 3 (= Baculites cf. asperoanceps Spath, 1921: 259, pl. 24 (fig. 4, 4a)).

Small forms with closely spaced, generally conical to low, rounded tubercles

situated near the dorsolateral edge of the flanks, e.g. SAM-PCZ8691 (Fig. 35A)

and SAM-PCZ8699 (Fig. 35E). Forms with more widely spaced tubercles cor-

respond to B. boulei Collignon (1931: 35, pl. 5 (fig. 2), pl. 9 (fig. 14)). A weak

longitudinal groove may be visible at mid-flank. e.g. SAM-5480 (Fig. 34D-F),

SAM-PCZ8675 (Fig. 35F) and PCZ8674b.

Form 4 (= ‘incurvatus’). Generally large forms with prominent conical to

rounded tubercles, situated near the dorsal third of the flanks, e.g. SAM-

PCZ9974 (Fig. 42A-D), SAS Z1795a-f (Fig. 44A-F) and SAS Z632a-c

(Fig. 45A-E, I-J).

Fig. 35 (see facing page). Baculites capensis Woods, 1906. A. SAM-PCZ8691.

B. SAM-PCZ8692. C. SAM-PCZ8749. D. SAM-PCZ8682. E. SAM-PCZ8699.

F. SAM-PCZ8675. All from locality 83, Zululand, St Lucia Formation, Coniacian IV.

G. SAM-PCZ8725. H. SAM-PCZ8720. J-K. SAM—PCZ8715. L. SAM-PCZ8711.

M. SAM-PCZ8713. N-P. SAM-PCZ8717. Q. SAM-PCZ8762. R. SAM-PCZ8747.

All from locality 85, Zululand, St Lucia Formation, Santonian I. All x 1.

Fig. 36 (see overleaf). Baculites capensis Woods, 1906. A-C. SAM-PCZ12002.

D-F. SAM-PCZ12003. G-I. SAM-PCZ12004. J-L. SAM-PCZ12005. O. SAS A1493b.

All from locality 22, Zululand, St Lucia Formation, Coniacian IV. M. SAM-PCZ12006.

N. SAM- 16026, both from Mkweyane (‘Umkwelane Hill’), probably locality 10, Zululand.

All x 1

Fig. 37 (see overleaf). Baculites capensis Woods, 1906. A-B. SAM-PCZ8674, concretion

to show co-occurrence of smooth and weakly nodose (boulei) forms of B. capensis. From

locality 83, Zululand, St Lucia Formation, Coniacian IV. Both x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA

Fig. 35

ANNALS OF THE SOUTH AFRICAN MUSEUM

CRETACEOUS FAUNAS FROM SOUTH AFRICA 67

68 ANNALS OF THE SOUTH AFRICAN MUSEUM

E |

O 10

A a dite Es)

L U

B ) 5

eee |

Fig. 38. Baculites capensis Woods, 1906. Suture lines and whorl

section. A. SASH13/6. B. SAS A335. C. SAM-1670b. Venter in

whorl section pointing downward. Scale bar for size.

Form 5 (= var. tenuetuberculata Collignon, 1966: 6, pl. 457 (figs 1863-4).

Generally large forms with weak conical to rounded tubercles, e.g. SAM-

PCZ12002 (Fig. 36A-C). These are probably only large forms (macroconchs)

of form 3.

Form 6. Generally large forms with prominent conical to crescentic, or

transversely elongated tubercles, situated near dorsal third of flanks, e.g.

NMB D1052a (Fig. 43A-C), NMB D1052c (Fig. 43F-H), SAS Z1795a

(Fig. 46A-C) and SAS A353 (Fig. 47A-B).

Form 7 (= capensis typical form). Generally large forms with rounded to

longitudinally elongated tubercles near the dorsal third of the flanks. Longi-

tudinal depression often present near midflank; whorl section varies from sub-

trigonal to distinctly elliptical, e.g. SAM-4823 (Fig. 27D-H) and SAM-4824

(Fig. 28B-E). In some, e.g. SAS Z632a (Fig. 45A-C), the transition from

CRETACEOUS FAUNAS FROM SOUTH AFRICA 69

conical (‘incurvatus’) to longitudinally elongated (capensis) tubercles can be

observed on the same specimen.

Form 8 (= B. capensis in Collignon 1966: 6, pl. 457 (fig. 1862)). Tubercles

extremely elongated longitudinally, assuming crescentic outline in dorsal view,

e.g. NMBD1028/3 (Fig. 50A-C).

Form 9 (= var. umsinenensis Venzo, 1936: 116 [58], pl. 10 [6] (figs 11-

12)). Tubercles elongated obliquely, some with shallow, longitudinal depression

at mid-flank. Tubercles closely or widely spaced, generally situated high on the

flank or near the dorsolateral edge, e.g. SAM-PCZ12016 (Fig. SOG-H), SAM-

PCZ12017 (Fig. SOM), NMB D1028/12 (Fig. SON) and SAS A361 (Fig. 33],

49C-E). Some specimens with very large tubercles, e.g. NMB D1028/3

(Fig. SOA-C), are transitional between forms 8 and 9.

Form 10 (= B. malagasyensis Collignon, 1966: 7, pl. 457 (fig. 1865)).

Tubercles are of the capensis or incurvatus type, but irregularly spaced or

doubled.

EC:

Fig. 39. Baculites capensis Woods, 1906. Suture lines and whorl

sections. A. SAM-PCZ12007. B. SAM-PCZ12006. C. SAM-

PCZ12008. Venter in whor! sections pointing downward.

Scale bar for size.

70 ANNALS OF THE SOUTH AFRICAN MUSEUM

Ze Wi

Fig. 40. Baculites capensis Woods, 1906. Suture lines and whorl

sections. A. SAS A2101. B. SAM-PCZ8039. C. SAM-PCZ12009.

Venter in whorl sections pointing downward. Scale bar for size.

Form 11. (= ?B. sparsinodosus Collignon, 1969: 23, pl. 521 (figs 2052-

2054)). Rare, ?7Lower Campanian forms with large, rounded, distantly spaced

tubercles, e.g. SAM-PCZ8022 (Fig. 544A), SAM-PCZ9971 (Fig. 54B) and

SAM-PCZ9972 (Fig. 54C-E). |

?Form 12. Rare Santonian specimens with very widely spaced tubercles,

e.g. SAM-PCZ8387 (Fig. 34M-O) and SAM-PCZ8349 (Fig. 34P-R). We are

not quite sure if these are true B. capensis or merely rare, tuberculate B. bailyi;

faunal association suggests the latter.

Apertures and dimorphism

The aperture, or part thereof, is preserved in numerous (about 35) speci-

mens (Figs 51A-N, 52A-D). It is simple, consisting of a short, rounded dorsal

CRETACEOUS FAUNAS FROM SOUTH AFRICA WA

Fig. 41. Baculites capensis Woods, 1906. Suture lines and whorl

section. A. SAM-PCZ9974. B-C. NMBD1028g. Venter in whorl

section pointing downward. Scale bar for size.

rostrum, a prominent lateral sinus and a long, ventral rostrum. The dorsal

rostrum may curve slightly outwards, i.e. away from the longitudinal axis of the

shell, whereas the ventral rostrum may curve inwards.

The diameter at which apertures are formed and maturity presumably sets

in, is quite variable (Fig. 53). The smallest whorl height at which indications of

an aperture are present, is 9 mm, whereas some shells reach maximum whorl

heights of 31 mm on the body chamber without having formed an aperture.

Even though specimens at the smaller and larger ends of the distribution range

may be distinguished as micro- and macroconchs respectively, there is no

distinct break in size between them. An alternative interpretation is that this

indicates a variable range of maturation size.

Most body chambers are perfectly straight, but some (e.g. NMB D1052c)

(Fig. 43F-H) are slightly curved.

Tz ANNALS OF THE SOUTH AFRICAN MUSEUM

Suture lines

Details of the sutures are shown in Figures 38A-C, 39A-B, 40A, C, and

41A-B. The suture is very simple with open saddles and lobes.

Discussion

According to our interpretation, B. capensis is a predominantly nodose

baculitid, which first appears in the second or third division of the Coniacian of

Zululand, and persists to the second division of the Santonian in Pondoland and

possibly to the first division of the Campanian in Zululand.

From the descriptions above, and the figures, it can be seen that the pres-

ence and shape of the lateral tubercles is extremely variable. This variation was

already noticed by Spath (1921) in material from Mkweyane (Umkwelane Hill)

collected by A. L. du Toit. Even though he referred several of the morphotypes

present to extant species in open nomenclature, e.g. Baculites sp. aff. capensis,

Baculites cf. aspero-anceps, Baculites cf. brevicosta and Baculites sp. cf.

sulcatus, he admitted that all of these forms were probably only varieties of

B. capensis. Our material not only confirms, but further illustrates the wide

variety of ornament in B. capensis.

The ornament of the different morphotypes suggests that B. capensis could

be derived from smooth B. yokoyamai through strengthening of groups or

sheaves of striae near the dorsolateral parts of the flanks, thus forming weak,

crescentic (‘brevicosta-schencki’) tubercles, as in form 2. Strengthening of these

crescentic nodes could lead to form 3 type of ornament, and further strength-

ening and wider spacing to type 4. Longitudinal elongation of these tubercles

could lead to typical B. capensis-type of ornament. Oblique arrangement of

typical B. capensis tubercles could lead to form 9 “‘umsinenensis’-type of

ornament and ultimately to B. menabensis—B. tanakae to be discussed below.

Alternatively, ‘umsinenensis’-type of ornament could be transitional between

form 2 brevicosta and form 7 (typical capensis). The suggested evolution of

capensis-type ornament from smooth B. yokoyamai is outlined in Figure 55.

Unfortunately, the answer is not that simple. This transition does not involve

a simple change from a population of smooth baculitids through crescentic to

conically nodose to longitudinally elongated tuberculate baculitids. Instead, the

transition seems to involve subtle shifts in proportions of the population. The

appearance and disappearance of distinct morphological features are not strictly

synchronous. Thus, the first appearance of nodose baculitids does not mean that

all smooth forms disappear (see e.g. Figs 29, 37)—instead, smooth forms con-

tinue to exist at least until the first appearance of typical capensis forms. Typical

capensis forms first appear together with weakly nodose forms, long before

Fig. 42 (see facing page). Baculites capensis Woods, 1906. A-D. SAM-PCZ9972.

E. NMBD1124/4. F-G. NMBD1028. H. SAM-PCZ12010. I. SAM-PCZ8760.

J. NMBD1124. K-M. SAM-PCZ8027, all from locality 72, Zululand, St Lucia Formation,

Coniacian III. All x 1.

Fig. 43 (see overleaf). Baculites capensis Woods, 1906. Body chambers.

A-C. NMBD1052a. D-E. NMBD1052d. F-H. NMBD1052c (note slight curvature).

All from locality 73, Zululand, St Lucia Formation, Coniacian IV-V. All x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA We

Fig. 42

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ep “3Id

CRETACEOUS FAUNAS FROM SOUTH AFRICA 75

Fig. 44. Baculites capensis Woods, 1906. A. SAS Z1795a. B. SAS Z1795c.

Cy SAS Z1795f. D. SAS Z1795. E. SAS Z1795d. F. SAS Z17951. All from locality 85,

Zululand, St Lucia Formation, Santonian I. All x 1.

Fig. 45.

ANNALS OF THE SOUTH AFRICAN MUSEUM

Baculites capensis Woods, 1906. A-C. SAS Z632a. D-E. SAS Z632b.

F-H. SAS Z632e. I-J. SAS Z632c. All presumably from locality 91, Zululand, St Lucia

Formation, Coniacian IV or V. All x 1. :

CRETACEOUS FAUNAS FROM SOUTH AFRICA Fl

Fig. 46. Baculites capensis Woods, 1906. A-C. SAS Z1795a. Macroconch with part of the

aperture preserved. From locality 85, Zululand, St Lucia Formation, Santonian I. All x 1.

78 ANNALS OF THE SOUTH AFRICAN MUSEUM

capensis-type of ornament reaches its peak. Also, many of these different mor-

phological types, even though stratigraphically contemporaneous, seem to be

concentrated at different localities (see e.g. Fig. 35 (Coniacian IV), Fig. 43

(Coniacian IV-V), Fig. 44 (Coniacian IV-V), Fig. 45A-E, I-J (Coniacian IV or

V), Figs 49C-E, 50A-C, N (Coniacian IV-V), etc.). Thus it is very common

for baculitids from a single locality all to be the same, but different from those

from another locality at more or less the same stratigraphic level.

Forms 1-3, i.e. smooth or with small crescentic or weakly conical tubercles

situated near the dorsolateral edge of the flanks, are most common amongst the

first representatives of the species in the second and third divisions of the Coni-

acian, but typical capensis forms with elongated tubercles also occur. This

includes most of the baculitids described from Umkwelane Hill (Mkweyane)

by Spath (1921). A similar early capensis baculitid assemblage occurs at

locality 15. In the fourth division of the Coniacian tuberculate forms of group 3

(boulei) dominate, but smooth forms still persist, often in the same nodule

(Fig. 37); in addition, typical forms of capensis, including forms 4—9 occur.

In the later part of the Coniacian and in the first two divisions of the San-

tonian, forms 4-9 dominate, but weakly ornamented forms still occur. Normally

we find that, at a given locality, the baculitids will not occur as a random mix-

ture of the different morphological types, but that one form will be dominant

(see e.g. specimens Z1795a, c-f (Figs 44A-F), Z632a-b, e (Fig. 45A—J) and

D1052a, c-d (Fig. 43A-H), all from more or less the same stratigraphic level

but different localities). Based purely on morphological criteria, each of these

morphotypes could be given a different name, but-from a stratigraphic point of

view, this would be illogical.

One morphotype that seems to be most common at, but not exclusively

restricted to, the outcrops along the Mzinene River at localities 71 and 73, is

form 9 (umsinenensis). Typical forms with distinct obliquely elongated tubercles

are best known, and first recorded by Crick (1907: 240) and later by Venzo

(1936: 116 [58], pl. 10 [6] (figs 11-12)) from this locality, but similar

specimens are also known from localities 22, 91, and 83.

Specimens with extremely elongated tubercles (form 8) are rare (Figs 33H,

50A). Forms with extremely distant tuberculation (form 11) are also rare and

have thus far only been found in rubble from excavations at locality 6, which

yielded a mixed Santonian II-III and/or Campanian I fauna. We think that these

baculitids are from the Campanian section of the excavations, but are not sure.

The ornament is comparable to material from the Lower Campanian of Mada-

gascar described by Collignon (1969) as B. sparsinodosus.

Form 12 is extremely rare and is mostly found in association with abundant

Santonian specimens of B. bailyi. Because of this, we suspect that these may

rather be atypical, nodose B. bailyi than B. capensis, but again we cannot be

completely sure.

Form 11 (malagasyensis) is merely a capensis with irregular spacing of the

tubercles.

Fig. 47. Baculites capensis Woods, 1906. Body chamber specimens. A-B. SAS A353.

C. SAM-PCZ8011. D-F. SAM-PCZ12011. All from locality 73, Zululand, St Lucia

Formation, Coniacian IV or V. All x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA

80 ANNALS OF THE SOUTH AFRICAN MUSEUM

Affinities and comparisons

Our interpretation of B. capensis is very broad, and consequently it is very

difficult or in cases impossible to separate it satisfactorily from similar, contem-

porary species known only from a few individuals and where the range of varia-

tion is unknown.

The first problem is separating B. capensis from B. bailyi. It is easy to name

the first nodose specimens B. capensis, but what should we call the smooth

variants? We consider it expedient to regard these smooth specimens as variants

of early B. capensis, albeit mainly by their co-occurrence with these nodose

forms. Fortunately, these smooth forms of B. capensis are generally larger than

B. bailyi, and have the whorl section of the former species, but separation of

isolated specimens remains difficult. Also, in some of the early forms of B. cap-

ensis, e.g. SAM-PCZ8030 (Fig. 49J), tubercles only appear at a relatively late

stage. The early, non-tuberculate parts of these shells are indistinguishable from

B. yokoyamai or B. bailyi.

Contemporary, nodose baculitids appear to be separated into three distinct

bio(?)geographic regions:

A. Indo-Pacific Region

1. Baculites boulei Collignon (1931: 35, pl. 5 (fig. 2, 2a), pl. 9 (fig. 14)).

il. Baculites schencki Matsumoto (1959: 113, pl. 32 (figs 1-6), text-figs 12a—-b,

13a—-c, 14a—b, 15-21, 22a—b, 23a—c, 24-25).

B. Western Interior of North America

i. Baculites codyensis Reeside (1927: 4, pl. 2 (figs 6-9)).

Fig. 48 (see facing page). Baculites capensis Woods, 1906. A. SAM-PCZ7198a from

locality 73, Zululand, St Lucia Formation, Coniacian IV or V. B. SAM-PCZ12012

(H200/84), body chamber with part of aperture preserved. From locality 83, Zululand, St

Lucia Formation, Coniacian IV. C. SAM-PCZ12013, body chamber showing transition

from conical to elongate, capensis tuberculation. From locality 16, Zululand, St Lucia

Formation, Coniacian ?III. D. SAM-PCZ12014 from locality 22, Zululand, St Lucia

Formation, Coniacian IV. E. SAM-PCZ7210, body chamber with hardly perceptible

tubercles from locality 72, Zululand, St Lucia Formation, Coniacian HI. F-H. SAM-5443

from Mkweyane (‘Umkwelane Hill’), Zululand. I. SAM-PCZ12015 from locality 89,

Zululand, St Lucia Formation, Coniacian IV. J. SAM-5467, body chamber with no

tuberculation; the original of Spath’s (1921: 260) Baculites sp. cf. sulcatus from Mkweyane

(‘Umkwelane Hill’), Zululand. All x 1.

Fig. 49 (see overleaf). Baculites. capensis Woods, 1906. A-B. SAS A334.

C-E. SAS A361—specimen with typical ‘umsinenensis’ type of ornament.

F-I. SAS A137—specimen showing shallow longitudinal groove under oblique lighting. All

from locality 73, Zululand, St Lucia Formation, Coniacian IV-V. J. SAM-PCZ8030,

microconch, showing transition from smooth to weakly tuberculate ‘brevicosta’ type of

ornament. From locality 15, Zululand, St Lucia Formation, Coniacian IV. All x 1.

Fig. 50 (see overleaf). Baculites capensis Woods, 1906. A-C. NMBD1028/3.

D-F. SAS A606. G-I. SAM-PCZ12016. J-L. SAS A335. M. SAM-PCZ12017.

N. NMBD1028/12. A-C, G-I, M-N—all specimens with obliquely elongated tubercles of

the ‘umsinenensis’ form 9 type, but note the variation in strength; A-C is closest to form 8;

D-F, with irregularly spaced, incipiently doubled tubercles, as in B. malagasyensis.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 81

Fig. 48

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 49

CRETACEOUS FAUNAS FROM SOUTH AFRICA

Fig. 50

84 ANNALS OF THE SOUTH AFRICAN MUSEUM

C. European Tethyan and Boreal Region

1. Baculites incurvatus Dujardin (1837: 232, pl. 17 (fig. 13)).

il. Baculites brevicosta Schluter (1876: 141, pl. 39 (figs 9-10)).

A. Indo-Pacific. Baculites boulei—Collignon established this species based on

nine syntypes. The lectotype, by subsequent designation of Matsumoto (1959:

118), is the specimen figured by Collignon (1931, pl. 5 (fig. 2, 2a)) (herein

Fig. 12J—L) from Mahagaga, Madagascar.

Baculites boulei has been interpreted in various, contradictory ways,

including by Collignon himself. As discussed above (p. 27), B. boulei was

described from a ferruginous conglomerate north-north-east of Mahagaga. Five

other ‘species’ of Baculites were found in association with B. boulei, one with

arcuate lateral tubercles—, Baculites cf. B. brevicosta Schluter (Collignon 1931:

34, pl. 5 (fig. 1, la), pl. 9 (fig. 13) (3 specimens), and four non-tuberculate

‘species’: B. sulcatus Baily (Collignon 1931: 36, pl. 5 (figs 3-5), pl. 9 (fig. 15))

(24 specimens), B. besairiei Collignon (1931: 37, pl. 5 (figs 6-9), pl. 9

(fig. 16)) (150 specimens), B. roedereri Collignon (1931: 38, pl. 5 (fig. 10,

10a), pl. 9 (fig. 17)) (1 specimen), and B. Jatelobatus Collignon (1931: 38, pl. 5

(figs 11-12), pl. 9 (fig. 18)) (3 specimens). The latter four smooth ‘species’ are

all considered synonyms of B. yokoyamai, as stated above.

. According to Collignon’s (1931: 35) original diagnosis, B. boulei is orna-

mented every 7-8 mm by a large low tubercle, which is arched and concave,

and elongated forward into a thin rib; between two successive tubercles, there

are 4-5 intercalatory ribs, which are visible only in the siphonal region and in

the immediate vicinity of the flanks. The species is similar to B. capensis, but,

according to Collignon (1931: 36) differs by its whorl section being distinctly

oval.

According to Collignon’s description and, judging by the faunal association

and age, it is clear that B. boulei is an early form of B. capensis as here inter-

preted, thus corresponding to form 3.

Collignon later recorded B. boulei (1938: 88, pl. 6 (figs 6-6b)), together

with Baculites incurvatus (Collignon 1938: 88, pl. 6 (figs 4, 4a-5, 5a)) and

Baculites cf. B. aspero-anceps (Collignon 1938: 89, pl. 6 (fig. 7-7b)) from

unequivocal Campanian strata at Andimaka. This is clearly incorrect, as is

Forster’s (1975: 168, pl. 4 (figs 3, 9), text-fig. 37) record of B. boulei from the

Lower Campanian of Mozambique. The suture figured by Forster as that of

B. boulei is more complex than that of Collignon’s original figure. Baculites cf.

B. asperoanceps has a trigonal whorl section, and seems close to B. nibelae sp.

nov. (see p. 162) from the Upper Campanian of Zululand or perhaps B. incre-

scens. (True B. asperoanceps Lasswitz (1904: 16 (236), pl. 3 (15) (fig. la—b))

(lectotype figured herein Fig. 129) is a Campanian species and possibly a

synonym of B. obtusus.)

Fig. 51 (see facing page). Baculites capensis Woods, 1906. Various specimens with parts

of the aperture preserved. A-C. NMBD1028/4. F. NMBD1124. K-N. NMBD1024/10,

all from locality 72, Zululand, St Lucia Formation, Coniacian II or II. D. SAS Z632e from

locality 91, Zululand, St Lucia Formation, Coniacian IV or V. G. SAM-1621b from an

unrecorded locality. H-J. SAM-PCZ12018 from locality 83, Zululand, St Lucia Formation,

Coniacian [V. All x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA

Fig. 51

86 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 52. Baculites capensis Woods, 1906. Two specimens with preserved apertures at

disparate sizes to illustrate dimorphism. A-C. Microconch. NMBD1075 from locality 73,

Zululand, St Lucia Formation, Coniacian IV or V. D. Macroconch. SAS Z1795a from

locality 85, Zululand, St Lucia Formation, Santonian J. Both x 1.

Matsumoto (1959: 118, pl. 32 (fig. 7a-c), pl. 33 (figs 4a—c, 5a-b, 6a-—d,

7a—-b), text-figs 27a—b, 28-32) recorded B. boulei from California, the first

record of the species outside Madagascar. Some of the Californian specimens

were reported together with B. schencki, from ‘Member V of the Redding

area’, which was dated as Coniacian. According to Haggart (1984) and Haggart

& Ward (1989: 226), however, all the Californian occurrences of B. boulei are

Santonian.

Haggart & Ward (1989: 226, fig. 3.7-3.10)) recorded a single specimen of

Baculites cf. B. boulei from the Santonian—Campanian of Vancouver Island.

Ornament is of the brevicosta form 2 type of B. capensis, but this is mainly an

early Coniacian form. As Haggart & Ward (1989: 226) suggested, the

Vancouver specimen resembles B. tanakae and is probably a late form of

B. capensis transitional to B. tanakae.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 87

op)

2; 10

WwW

op)

eo)

= 5

; E < eT

5 10 15 20 25 30

MAXIMUM WHORL HEIGHT IN MM

Fig. 53. Baculites capensis Woods, 1906. Histogram illustrating distribution of maximum

whorl height. Solid circles represent specimens with the aperture preserved. Note the absence

of a sharp break between apparent macro- and microconchs suggesting, perhaps, a uniform

variable range of maturation sizes.

Matsumoto & Obata (1963: 43, pl. 13 (figs 3, 5), pl. 15 (fig. 6), text-

figs 93, 152-155) subsequently described B. boulei from Hokkaido, where it is

partly coeval with both B. capensis and B. schencki, i.e. Coniacian and San-

tonian (Matsumoto & Obata, text-fig. 216).

Kennedy (1986b: 112) regarded B. boulei as having a stouter whorl section

than B. capensis and more closely spaced tubercles, which lie nearer to the

dorsum than in the latter species.

Baculites schencki Matsumoto (1959: 113, pl. 32 (figs la-c, 2a-c, 3a-b,

4a-b, 5a-c, 6a-c), text-figs 12a—b, 13a—c, 14a—b, 15-21, 22a—b, 24-25) typi-

cally has an ovoid whorl section, and closely spaced, crescentic lateral nodes. It

differs in typical forms from B. boulei by the weaker lateral ornament, and the

narrower venter, but intermediate forms occur. This is very similar to, or ident-

ical with what we regard as form 2 of early B. capensis.

According to Matsumoto (1959: 120), the stratigraphic ranges of the two

species overlap in California: B. schencki is more common in the lower part

(of Member IV and V of the Redding area), whereas B. boulei is relatively

common in the upper part.

In comparing B. boulei and B. schencki with B. capensis, Matsumoto (1959:

125) remarked that in California ‘the three species, B. schencki, B. boulei and

B. capensis are nearly contemporary, but their stratigraphic positions of the

ANNALS OF THE SOUTH AFRICAN MUSEUM

‘ty tig “iy ty iy

Fig. 54

CRETACEOUS FAUNAS FROM SOUTH AFRICA 89

maximum abundance are arranged in ascending order’. Our Zululand material

partially confirms this, although the B. schencki-type of ornament is not as

common here as in California. (According to Haggart (1984: 232) no further

Seecmmens Of B. boulei or of B. schencki were collected in the Chico

Formation.) Our material confirms that the stratigraphic ranges of these forms

are largely overlapping.

Thus on stratigraphic grounds alone, there is no justification for formally

separating B. schencki, B. boulei and B. capensis. We admit that, in some

areas, one of the morphologies may be more common, but if different

morphologies are to be the criterion for separating B. schencki and B. boulei

from B. capensis, then we may as well regard all the varieties of B. capensis

described above as different species. We thus regard B. schencki and B. boulei

as synonyms of B. capensis. A case could be argued to refer to B. schencki,

B. boulei and B. capensis as early, intermediate and typical forms but, as noted

above, these morphologies are partly co-eval, and this terminology would be

confusing.

B. Western Interior of North America. In his original description of B. cap-

ensis, Woods (1906: 343) remarked on the similarities between the former and

B. asper Morton (1834: 43, pl. 1 (figs 12-13), pl. 13 (fig. 2)), stating that the

latter differed ‘in having larger and transversely elongated tubercles’. Spath

(1921: 257) also referred to the resemblance between B. capensis and B. asper.

According to Spath, the specimen figured by Meek (1876, pl. 39 (fig. 10a only))

and a specimen from Mississippi in the British Museum ‘are close to the South

African species in all characters but the suture line’. Matsumoto (1959: 125)

also commented on the similarities between B. capensis and B. asper, conclud-

ing that it was ‘probably a parallelism between the entirely separated biogeo-

graphic provinces’. Similarities between the two species were again referred to

by Matsumoto & Obata (1963: 50).

Unfortunately, B. asper appears to be uninterpretable at present. According

to Morton (1834: 44), B. asper was initially discovered by Mr Nuttal at

Cahawba, Alabama, and later found by Mr Conrad at Prairie Bluff, also in

Alabama. These localities are both appreciably younger than the Coniacian-

Santonian range of B. capensis. Cahawba is 1n the Selma Chalk, which is Late

Campanian, and the Prairie Bluff Chalk is Maastrichtian.

Reeside (1962: 116) mentioned that only one of Morton’s specimens sur-

vived. This had a label in Morton’s handwriting that states the locality as being

Prairie Bluff—i.e. Maastrichtian. Reeside concluded that Conrad’s locality data,

accepted by Morton, was erroneous, and interpreted B. asper as a Coniacian,

Santonian and early Campanian species.

Kennedy & Cobban (1991a: 72), on the other hand, regarded all the Ameri-

can Coniacian-Santonian records of ‘B. asper’ non Morton as B. codyensis

Fig. 54 (see facing page). Baculites capensis, form 11. Compare with B. sparsinodosus

Collignon, 1969. A. SAM-PCZ8022. B. SAM-PCZ9971. C-E. SAM-PCZ9972. All

from rubble from excavations at locality 6, Zululand, St Lucia Formation, Santonian II-III to

Campanian I; presumably from the Campanian. Note the absence of tubercles on the body

chambers in A and B. All x 1.

90 ANNALS OF THE SOUTH AFRICAN MUSEUM

e F

Fig. 55. Baculites capensis Woods, 1906. Suggested evolution of ornament, from

B. yokoyamai (A), through B. brevicosta (B), schencki (C), boulei (D), to typical capensis

(E) and umsinenensis (F), and ultimately to B. menabensis type (Fig. 57) (see p. 93).

Reeside (1927a: 4, pl. 2 (figs 6-19)). Typical forms of B. codyensis are easily

distinguished from all varieties of B. capensis. These have crescentic, rib-like

lateral tubercles, which may be projected prominently over the venter, super-

ficially resembling B. sulcatus. Atypical forms of B. codyensis, previously mis-

identified as B. asper, have strong, distant bullae and a stouter whorl section

(see e.g. Kennedy & Cobban 1991a, pl. 16). These are identical to some of our

B. capensis, referred to as form 6 (see e.g. Fig. 43A-H, 47A-F) by their

strong, crescentic dorsolateral tubercles. Even though some representatives of

B. capensis and B. codyensis appear identical, the populations as a whole are

very distinctive. No known specimens of B. capensis ever develop as strong and

regular crescentic lateral ribbing as in typical B. codyensis, and no forms of

B. codyensis ever develop longitudinally elongated lateral tubercles and a slight

depression at mid-flank as in typical B. capensis.

An as yet unnamed baculitid fauna from the Upper Turonian—Lower Coni-

acian of Mossamedes, Angola, collected by Dr M. R. Cooper and now in the

collections cf the S.A. Museum, is of interest in apparently linking smooth

CRETACEOUS FAUNAS FROM SOUTH AFRICA

B. yokoyamai and ornate B. codyensis. These specimens all have dorsolaterally

situated, closely spaced, crescentic tubercles (Figs 131N-R, 132A-F). These

are very similar to our form 3 or B. schencki, and seem to suggest that acquisi-

tion of lateral ornament took place in a similar manner in both B. codyensis and

{

Pe COREL 4

Fig. 56. Baculites incurvatus Dujardin, 1837. Numbers 13a-d: copy of

Dujardin (1837, pl. 17 (figs 13a-d)). Numbers 8-10: copy of D’Orbigny (1842,

pl. 139 (figs 8-10)). Note the apparent ventral keel in both figure 13c of

Dujardin and figure 9 of D’Orbigny.

92 ANNALS OF THE SOUTH AFRICAN MUSEUM

B. capensis, albeit at different times. Lateral ornament in the B. codyensis

lineage first appears in the Lower Coniacian, whereas that of B. capensis occurs

later, in the Middle or Upper Coniacian as shown by the Zululand and the

Madagascan Mahagaga fauna.

C. European Region. Less easy to resolve are the relationship between the

Indo-Pacific and the European nodose baculitids. In contrast to the Western

Interior and the Indo-Pacific regions, baculitids are not as common in the

European Tethys and are poorly preserved, and consequently the species are

poorly known.

The relevant species are B. incurvatus Dujardin, 1837, and B. brevicosta

Schluter, 1876. The former name has, on occasions, been used indiscriminately

for European Baculites with widely spaced nodes; the latter has been used for

specimens with closely spaced, slightly oblique tubercles near the dorsolateral

edge of the flanks.

Baculites incurvatus was recently reviewed by Immel ef al. (1982), and

Kennedy (1984), based on Austrian (Gosau) and French material respectively.

Dujardin based this species on a series of fragments. The largest, a curved body

chamber, was designated lectotype by Immel et al. (1982: 27). The lectotype

(MNHP R1025a) and paralectotypes (MNHP R1025b-c (?e in WJK p. 143))

were refigured by Kennedy (1984, pl. 33 (figs 4-6, 15, 19-22)).

It is difficult to reconcile Dujardin’s original figures (herein Fig. 56—13a-—d)

and descriptions on the one hand, and the type material on the other. Dujardin

(1837: 232) gave the following diagnosis: ‘Testa vaginaeformi, compresa,

versus basim aliquantum incurvata; dorso nunc carinato, nunc rotundo, rugoso

aut laevi, utrinque tuberculis evanescentibus instructa’. In his illustrations, two

whorl sections are figured. The smaller (pl. 17 (fig. 13d)) is distinctly ovoid, but

the larger (pl. 17 (fig. 13c)) shows distinct lateral sulci on either side of the

venter, suggesting either a ventral keel, or a row of ventral (siphonal)

tubercles—although the latter seems to be unlikely. If there were ventral

tubercles on the body chamber, surely these would have shown up in the lateral

view of the body chamber. The lectotype, MNHP R1025a, however, shows no

trace whatsoever of a ventral keel.

Fig. 57 (see facing page). Baculites menabensis Collignon, 1969. Plaster cast of the

holotype, GD 12036, the original of Collignon (1969, pl. 518 (fig. 2036)) from the Lower

Campanian of gisement 307, km 15600 Coupe Ampolypoly-Antsirasira-Behamotra (Belo sur

Tsiribihina), Madagascar. x 1.

Fig. 58 (see overleaf). A-C. Baculites falcatus Collignon, 1969. Plaster cast of the

holotype, GD 12045, the original of Collignon (1969, pl. 520 (fig. 2045)) from the Lower

Campanian of gisement 303, km 15000 Coupe Ampolypoly-Antsirasira-Behamotra (Belo sur

Tsiribihina), Madagascar. D-F. Baculites ventroplanus Collignon, 1969. Plaster cast of the

holotype, GD 12048, the original of Collignon (1969, pl. 520 (fig. 2048)) from the same

locality as above. Both x 1.

Fig. 59 (see overleaf). A-C. Baculites antsirasiraensis Collignon, 1969. Plaster cast of the

holotype, GD 12040, the orginal of Collignon (1969, pl. 519 (fig. 2040)) from the Lower

Campanian of gisement 304, km 15200 Coupe Ampolypoly-Antsirasira-Behamotra (Belo sur

Tsiribihina) Madagascar. D-F. Baculites subtilis Collignon, 1969. Plaster cast of the

holotype, GD 12042, the original of Collignon (1969, pl. 519 (fig. 2042)) from the same

locality as above. Both x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA

ie Aare

"Re BRM

sy rs

Fig. 57

ANNALS OF THE SOUTH AFRICAN MUSEUM

gc 314

CRETACEOUS FAUNAS FROM SOUTH AFRICA

Fig. 59

25)

96 ANNALS OF THE SOUTH AFRICAN MUSEUM

A B

Fig. 60. Baculites sparsinodosus Collignon, 1969. Plaster cast of the holotype, GD 12052,

the original of Collignon (1969, pl. 521 (fig. 2052)) from the Lower Campanian of

gisement 302, km 14200, Coupe Ampolypoly-Antsirasira-Behamotra (Belo sur Tsiribihina),

Madagascar. x 1.

D’Orbigny’s (1842: 564, pl. 139 (figs 8-10)) (herein Fig. 56-8-10) sub-

sequent description and illustration of B. incurvatus is better known than that of

Dujardin. Again, D’Orbigny’s figure shows a baculitid with a curved body

chamber, and a whorl section that is ovoid on the phragmocone, but distinctly

carinate on the body chamber. Two specimens in D’Orbigny’s collection, pre-

sumably used in his reconstruction of B. incurvatus, survive and were refigured

by Kennedy (1984, pl. 33 (figs 1-3, 16-18)). Neither shows any trace of a

ventral keel. We suspect that D’Orbigny’s figure of B. incurvatus may have

been based more on Dujardin’s original figures than on his own material...

CRETACEOUS FAUNAS FROM SOUTH AFRICA 97

LW)

A L

0 10 mm

piers eae ee

E en

@) 10 mm

ibe ase Se ey

E L

. |

0 5 mm

pee ee

Fig. 61. Suture lines. A. Baculites sparsinodosus. Collignon, 1969 (see

Fig. 60). B. Baculites falcatus Collignon, 1969 (see Fig. 58A-C).

C. Baculites subtilis Collignon, 1969 (see Fig. 5S9D-F). Scale bar for size.

Subsequent interpretations of B. incurvatus are confused. Meek (1876: 392)

divided the genus Baculites into two groups. His group b was defined as

follows: ‘?b. Shell straight posteriorly, but with the non-septate part gently

arcuate; aperture a little oblique; appendage of siphonal side of lip arching

slightly with the general curvature of the non-septate part, but not curving over

the aperture—(B. incurvatus Dujardin)’. Spath (1926: 80) subsequently

introduced the new baculitid genus Euhomaloceras as follows: ‘the new genus

Euhomaloceras gen. nov. is proposed for Meek’s group b (Invertebr. Cret. and

Tert. Fossils, U.S. Geol. Surv. Territ., vol. ix, 1876, p. 392) with B. incurvatus

98 ANNALS OF THE SOUTH AFRICAN MUSEUM

E L

mre 3

C O 5 mm

[ea ae bee

Fig. 62. Suture lines. A. Baculites ambatryensis Collignon, 1971 (see

Fig. 128). B. Baculites antsirasiraensis Collignon, 1969 (see Fig. 59A-C).

C. Baculites menabensis Collignon, 1969 (see Fig. 57). Scale bar for size.

Dujardin (in D’Orbigny, Pal. Franc., Terr. Cret., vol. 1, 1842, p. 564, pl. 139

(fig. 8)) as genotype.’ Spath’s concept of Euhomaloceras is difficult to interpret.

Both Dujardin and D’Orbigny’s figures show a curved body-chamber, and it

was this feature that defined Meek’s group b; Meek did not refer to the whorl

section at all. Wright (1957: L218) provided the first, and only diagnosis of

Euhomaloceras as: ‘Bodychamber gently curved with distinct rounded siphonal

and laterodorsal tubercles.’ The mention of siphonal tubercles is based on a mis-

interpretation of the cross section in the figures of Dujardin and D’Orbingy.

Fig. 63 (see facing page). Baculites sulcatus Baily, 1855. A-B. BMNH C35625, the

lectotype, and impression of paralectotype, from an unrecorded horizon at locality 1,

Pondoland, Mzamba Formation—presumably from the upper parts of the section,

Campanian ?J. x5. Photograph courtesy of Natural History Museum, London.

CRETACEOUS FAUNAS FROM SOUTH AFRICA

100 ANNALS OF THE SOUTH AFRICAN MUSEUM

Subsequent examination of Baculites has shown that curvature of the body

chamber is a very variable and inconsistent feature, and is certainly not of

generic significance. The name Euhomaloceras is thus unnecessary as discussed

by Immel ef al. (1982: 27-28).

As mentioned above, none of the surviving specimens on which Dujardin

and D’Orbigny based their figures shows any indications of a ventral keel on the

body chamber. Why both Dujardin and D’Orbigny figured a ventral keel on the

body chamber of B. incurvatus remains a mystery. We suspect that Dujardin

may have erred, and that D’Orbigny’s artist merely copied the error.

In France, B. incurvatus first occurs in the Middle Coniacian in Touraine

and Aquitaine, and persists through the Upper Coniacian into the Santonian, a

stratigraphic range nearly identical to that of B. capensis. In Austria it occurs in

the Lower Santonian. |

Younger Campanian records of B. incurvatus, e.g. Holzapfel (1887: 64,

pl. 4 (figs 5-6), pl. 5 (fig. 10)) from the Vaals Formation at Aachen, Germany;

Muller & Wollemann (1906: 4, pl. 2 (figs 2-5)) from Braunschweig and

Broitzem in northern Germany, and Collignon (1938: 88, pl. 6 (figs 4-5)) from

the Campanian of Andimaka in Madagascar, are all probably misidentifications.

The specimens referred to B. incurvatus by Holzapfel and Muller & Wollemann

were considered the same as Baculites sp. 1 by Kennedy (1986b: 110, pl. 17

(figs 7-9, 13-15, 21-23), pl. 18 (figs 18-22), pl. 23 (figs 1, 7), text-fig. 8A—C),

from the Upper Campanian of France. However, Muller & Wollemann’s speci-

mens show distinct longitudinally to obliquely elongated tubercles, whereas

Kennedy’s Baculites sp. 1 has transversely elongated tubercles. In this respect,

the German specimens seem closer to some of the baculitids from the Lower

Campanian of Madagascar described by Collignon (1969); all are probably

synonyms of B. menabensis Collignon or B. tanakae Matsumoto & Obata,

1963, as discussed below (p. 109).

The Campanian B. incurvatus of Collignon (1938) has a distinct trigonal

whorl section and is closer to B. bassei Besairie or B. nibelae sp. nov., to be

described below.

Fig. 64 (see facing page). Baculites sulcatus Baily, 1855. Typical forms. A. SAM-7043.

Specimen on back of block with Hauericeras figured by Klinger & Kennedy (1980, fig. 5b),

collected by T. Gevers from the top bed, T2. B-D. SAM-PCP5695. E-G. SAM-PCP5684.

H-I. BMNH C35625, the lectotype. J. SAM-PCP8420. K. SAM-PCP8656. L-N. SAM-

PCP5684. A-G, J-N, all from the top beds at Mzamba Cliff, Pondoland, Mzamba

Formation, Campanian I. A-D, H-N x 1; E-G ~x 2.

Fig. 65 (see overleaf). Baculites sulcatus Baily, 1855. All weakly ornamented forms.

A-C. SAM- PCP8653. D-F. SAM-PCP8361. G-H. SAM-PCP8656. I-K. SAM-—

PCP8153b. L-N. SAM-PCP8422. O-Q. SAM-PCP8153. R. SAM-PCP8660. All from

the upper beds at the type section of the Mzamba Formation, Campanian I. All x 1.

Fig. 66 (see overleaf). Baculites sulcatus Baily, 1855. Early forms—Van Hoepen

collection, Transvaal Museum. A-B. TM540g. C-E. TM548c. F. TM540f. G-I.

TM 548a (microconch with part of aperture preserved). J-L. TM540a. M-O. TM 540b.

O-P. TM 540d. All from an unspecified horizon at locality 1, Pondoland, Mzamba

Formation, presumably below the level of typical forms figured in Figures 64-65.

axitll Se Ay

101

CRETACEOUS FAUNAS FROM SOUTH AFRICA

. 64

102 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 65

CRETACEOUS FAUNAS FROM SOUTH AFRICA 103

Fig. 66

104 ANNALS OF THE SOUTH AFRICAN MUSEUM

It is difficult to distinguish satisfactorily between B. capensis and

B. incurvatus. Curvature of the body chamber, initially used to separate

B. incurvatus (as Euhomaloceras) from the other baculitids, is of little

significance, and indeed is shown only by the lectotype and by one other

specimen figured to date (Kennedy 1984, pl. 33 (figs 4-6, 7-9)). Curved and

straight body chambers occur in both species; occurrence of curved body

chambers is apparently random.

As Woods (1906: 342) had already noticed, typical B. capensis differs from

B. incurvatus by the longitudinal elongation of the tubercles, and by the pres-

ence of a shallow sulcus at mid-flank. A single specimen of B. incurvatus

figured by Kennedy (1984: 143, pl. 33 (12-14)) has doubled tubercles and a

Shallow lateral groove, which, apart from being smaller, is very similar to

Collignon’s (1966: 7, pl. 457 (fig. 1865)) B. malagasyensis, here regarded a

synonym of B. capensis s.s. Apart from this specimen, which was regarded by

Kennedy as possibly being pathologic, no specimens of B. incurvatus are as yet

known with typical B. capensis ornament.

However, the majority of specimens of B. capensis with rounded, conical or |

slightly crescentic tubercles are indistinguishable from contemporary

B. incurvatus.

Woods (1906: 342), Spath (1921: 257-258) and Matsumoto (1959: 125) all

commented on minor differences in suture lines of B. capensis and B. incurv-

atus. We can see absolutely no significant difference between the sutures of

B. capensis and B. incurvatus (Kennedy 1984, text-fig. 42F-H).

Immel et al. (1982: 28) also considered the whorl sections to be different—

that of B. capensis being more elliptical, with wide venter and flat flanks.

Again, typical forms of B. capensis do have elliptical whorl sections, but many

have whorl sections indistinguishable from those of known B. incurvatus (see

lees 30), SIL).

It is interesting to note that some specimens of B. incurvatus are virtually

smooth and lack tubercles, and show a similar range of variation in this respect

as B. capensis.

Clearly, on the basis of the disparate sizes in population of B. capensis—

numbering several hundred specimens, and B. incurvatus—numbering perhaps

30 specimens, no final decision can be taken as to whether B. capensis should

be regarded as a junior synonym of B. incurvatus. However, given their similar

ages, but apparent geographic separation, it might eventually be feasible to

regard them as a subspecies (of B. incurvatus).

Fig. 67 (see facing page). A-D, H-N. Baculites sulcatus Baily, 1855. Early forms.

Van Hoepen Collection, Transvaal Museum. Smooth to weakly ribbed specimens.

A-B. TM548a. Microconch with aperture. C-D. TM548e. H-J. TM548d. Microconch

with complete aperture. Note long, slightly inwardly-curved linguiform ventral rostrum and

short, outwardly-curved dorsal rostrum. All from an unspecified horizon at locality 1,

Pondoland, Mzamba Formation, presumably below the level of typical forms figured in

Figure 64. E-G. Baculites furcillatus (Blanckenhorn, 1905). SAM -PCI8573 from the

Campanian of Israel (ex Z. Lewy collection). K-Q. Baculites bailyi Woods, 1906.

K-L. SAM-PCP6766. M. SAM-13103. Specimen from Geological Commission collection,

one of Woods’ paratypes of B. bailyi. N. SAM-PCP6765. O-Q. SAM-PCP8729, all from

the basal beds at locality 1, Mzamba Formation, Santonian II. A-J, M x 1; K-L, N-Q x 2.

105

CRETACEOUS FAUNAS FROM SOUTH AFRICA

Fig. 67

106 ANNALS OF THE SOUTH AFRICAN MUSEUM

; \

( \

ew \

D O 10 mm

ee S|

aa CLE

Fig. 68. Baculites sulcatus Baily, 1855. Typical form. Whorl sections,

suture line and ornament. A. SAM-PCP8421. B. SAM-PCP8420.

C. SAM-PCP5685. D. SAM-PCP5684., E. SAM-PCPS42

F. SAM-PCP8419. Venter in whorl sections pointing downward.

Scale bar for size.

The other European baculitid that falls within the morphological limits of

B. capensis is B. brevicosta Schliiter. Unfortunately, the only unequivocal

record of this species is the original description from northern Germany given

by Schliter (1876: 141, pl. 39 (figs 9-10)). Schliiter based this species on

several specimens (not just two as claimed incorrectly by Kennedy 1984: 146).

The original of Schliiter (1876, pl. 39 (figs 9-10)) was designated lectotype by

CRETACEOUS FAUNAS FROM SOUTH AFRICA 107

L U

O 5 mm

A | ba aa

L U |

O 5 mm

B pe pear ai

Fig. 69. Baculites sulcatus Baily, 1855. Typical form. Suture lines.

A. SAM-PCP8152. B. SAM-PCP8153. C. SAM-PCP8661.

Scale bar for size.

Kennedy (1984: 146). Unfortunately, neither the lectotype nor any of the other

paralectotypes could be traced by us in the collections in Bonn, and we have to

assume that they are lost.

Subsequent records of B. brevicosta all have to viewed with caution. The

specimens from the Lower Senonian of Sweden (with Actinocamax quadrata)

from Eriksdal, Sweden, described and figured by Moberg (1885: 37, pl. 4

(figs 5-6)) have very weak, crescentic lateral ribs, rather than dorsolaterally

situated tubercles as in B. brevicosta, and they do not seem to belong to this

species.

The specimens from Mkweyane (Umkwelane Hill), Zululand, described and

figured by Spath (1921: 146, pl. 24 (fig. 5)) as Baculites cf. brevicosta are

identical to Schluter’s figures, as also noted by Spath (1921: 260) ‘since it

probably only represents a variety of B. capensis . . . its exact agreement with

Schliter’s species may be a case of heterochronous homoeomorphy’. The

108 ANNALS OF THE SOUTH AFRICAN MUSEUM

relationship of this (Zululand) specimen to typical B. capensis is probably the

same as that of European B. brevicosta to B. incurvatus. Their relative strati-

graphic occurrences in France (Kennedy 1984) seem to confirm this.

Kaplan & Kennedy (1994: 59, pl. 40 (figs 15-19)) described and figured

alleged B. brevicosta from the Coniacian of Westphalia. These show some

B. brevicosta to have distinct ventral ribbing and weak lateral ornament, which

places it closer to B. undulatus than to B. capensis. Nevertheless, it is still a

rare species and the extent of variation is not fully known.

A closer analogue to B. brevicosta is B. schencki Matsumoto, 1959, as

discussed above (see p. 87). Baculites schencki, here regarded as an early form

of B. capensis, was said to differ from B. brevicosta merely by the strength of

the tubercles (Matsumoto 1959: 117). The association of B. schencki and

B. capensis is the same as that of B. brevicosta and B. incurvatus, thus further

suggesting that the relationship of B. capensis and B. incurvatus is very close,

and that separation at subspecific level only is probably justifiable.

In Zululand and Pondoland, the typical B. capensis lineage ends in the

Middle Santonian, although some atypical forms (form 11, B. sparsinodosus?)

may occur in the Lower Campanian of Zululand. These last representatives

include typical B. capensis forms with longitudinally elongated tubercles, with

the middle part of the flanks flat or slightly depressed and with a round or

fastigiate venter.

In Pondoland, B. capensis is succeeded by B. sulcatus in the Lower

Campanian, and in Zululand by B. increscens—B. vanhoepeni in the Middle

Campanian—both with typical rib-like or auricular lateral ornament. Morpho-

logically these seem far removed from the B. capensis type of ornament.

Some of the baculitids from Pondoland, described as B. sulcatus by Van

Hoepen (1921: 18, pl. 3 (figs 7-8)) and later referred to Baculites vagina var.

vanhoepeni by Venzo (1936: 116 [58], pl. 10 [6] (figs 11-12)) (see Fig. 66),

could possibly be seen as a link between B. capensis and B. sulcatus + B. van-

hoepeni as is to be discussed below (p. 150).

Fortunately, Collignon (1969) has described a rich baculitid fauna from

the Lower Campanian of Madagascar that seems to continue the B. capensis

lineage. This fauna includes: B. menabensis Collignon (1969: 15, pl. 518

(figs 2036-7)) (herein Fig. 57); B. antsirasiraensis Collignon (1969: 18, pl. 519

(figs 2040-2041)) (herein Fig. 59A-C); B. subtilis Collignon (1969: 18, pl. 519

(figs 2043-2044)) (herein Fig. S9D-F); B. falcatus Collignon (1969: 20, pl. 520

(figs 2045-2046)) (herein Fig. 58A-C); B. ventroplanus Collignon (1969: 20,

pl. 520 (figs 2048-2050)) (herein Fig. 58D-F) and B. sparsinodosus Collignon

(1969: 23, pl. 521 (figs 2052-2054)) (herein Fig. 60). With the exception of

B. sparsinodosus, all these species have dorsoventrally compressed, longitudi-

nally elongated tubercles of the B. capensis type. They differ from typical

B. capensis mainly in that the tubercles are more pinched, slightly oblique and

situated nearer to the dorsum. Also, the whorl section becomes more com-

pressed, and in some, e.g. B. ventroplanus, the venter becomes distinctly

fastigiate. The ornament of these Lower Campanian Madagascan species is

nearly identical to that of form 9 or ‘umsinenensis’ type of B. capensis, which

makes its first appearance in the Upper Coniacian of Zululand.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 109

E fi U

= L

O 5 mm

CS es ae eee!

Fig. 70. Baculites sulcatus Baily, 1855. Suture lines. A. SAM-PCP12019.

B. SAM-PCP8361. C. TM540m—note similarity to Woods’ figured suture

line of B. bailyi (1906, pl. 44 (fig. 5)). Scale bar for size.

With the possible exception of B. sparsinodosus, all these Lower Campanian

Madagascan Baculites are probably synonyms; as first revising authors, we

select B. menabensis as the name of this species.

As far as lateral ornament is concerned, Madagascan B. menabensis is indis-

tinguishable from similarly dated specimens described as B. incurvatus from

Braunschweig in Germany by Miller & Wollemann (1906: 4, pl. 2 (figs 2-5))

and the Vaals Formation on the Belgian—Dutch border by Holzapfel (1887: 64,

pl. 4 (figs 5-6)). More and precisely located material is needed to resolve the

true identity of these European specimens.

Baculites tanakae Matsumoto & Obata (1963: 51, pl. 13 (fig. 4), pl. 16

(figs 1-5), pl. 17 (figs 1-5), pl. 18 (figs 1, 3-4), pl. 19 (figs 1, 4), text-figs 97-

113, 115)) from the Lower Campanian of Hokkaido is probably a senior syn-

onym of B. menabensis as here interpreted. According to Matsumoto & Obata

110 ANNALS OF THE SOUTH AFRICAN MUSEUM

OBEY

D E

O 5 mm

F ' ee ee |

Fig. 71. Baculites sulcatus Baily, 1855. Early form. Whorl sections.

A-B. TM540g. C. TM540h. D. TM540a. E. TM540b. F. TM 540c.

Venter pointing downward. Scale bar for size.

(1963: 54), B. tanakae occurs above the beds with B. capensis in Hokkaido.

The roof-shaped (fastigiate) venter and the shape of the tubercles that varies

from ‘asymmetrically crescentic or obliquely elongated to longitudinally

elongated, as in B. capensis’ (Matsumoto & Obata 1963: 52-53) in B. tanakae

is identical to that of the Madagascan B. menabensis. We can see no valid

grounds for maintaining B. tanakae and B. menabensis as separate species.

Baculites tanakae differs from B. capensis by the fastigiate venter, and the

predominance of obliquely or longitudinally elongated tubercles that are situated

higher up on the flanks than in typical B. capensis. As mentioned above, this

CRETACEOUS FAUNAS FROM SOUTH AFRICA hit

style of ornament is very similar to that of form 9 or var. ‘umsinenensis’ in

B. capensis. This seems to indicate that the first appearance of B. tanakae-

B. menabensis type of ornament already occurs in the Upper Coniacian, but that

dominance of this type of ornament only takes place much later, in the Lower

Campanian. Unfortunately, we have no records of B. capensis from the Upper

Santonian of Pondoland or Zululand, but we suspect that in Hokkaido, where

the species occurs throughout the Santonian, specimens with this type of orna-

ment should be found. Alternatively, the occurrence of ‘umsinenensis’ type of

ornament has to be regarded as iterative; we cannot say for sure.

Occurrence

Mainly Middle Coniacian to Middle Santonian of Zululand, but possibly

also Lower Campanian, and Upper Coniacian to Lower Santonian of Mada-

gascar, Middle Santonian of Pondoland, Santonian of Hokkaido and California,

Upper Santonian of U.S. Gulf Coast, and, doubtfully, Lower Campanian in

Angola.

Baculites sulcatus Baily, 1855

Figs 63-66, 67A-D, H-N, 68-77, 78C

1855 = Baculites sulcatus Baily, p. 457, pl. 11 (fig. 5c only, non 5a-b) (= B. bailyi

Woods).

1906 = Baculites sulcatus Baily; Woods, p. 341, pl. 44 (fig. 4).

1921 Baculites sulcatus Baily; van Hoepen, p. 18, pl. 3 (figs 7-8).

non 1921 ~~ Baculites sp. cf. sulcatus Baily; Spath, p. 260 (= B. capensis Woods).

1922 Baculites sulcatus Baily; Spath, p. 146.

? 1930 Baculites Bailyi Woods; Besairie, p. 223, pl. 21 (fig. 6 only).

non 1931 ~~ Baculites sulcatus Baily; Collignon, p. 36, pl. 5 (figs 3, 3a, 4, 4a, 5, 5a, 13,

13a), pl. 9 (fig. 15). (= ?B. yokoyamai Tokunaga & Shimizu).

non 1963 Baculites n. sp. (?) aff. B. sulcatus Baily; Matsumoto & Obata, p. 46, pl. 12

(fig. 6), text-figs 94, 130.

1977 = Baculites sulcatus Baily; Klinger & Kennedy, p. 75, figs 3B-E, J-L.

Type

Baily (1855) apparently based this species on more than one specimen—

Woods (1906: 341) refers to (the) * . . . only specimens seen are the types

.... Matsumoto & Obata (1963: 46) claimed that Woods did not designate a

lectotype. They accordingly designated the specimen figured in ventral view by

Baily (1855, pl. 11 (fig. 5c)) as lectotype. However, Woods (1906, pl. 44

(fig. 4)) referred to this same specimen (which he figured in lateral view) in the

plate explanation as ‘The Type, Museum of the Geological Society of London’.

In our view, this is a perfectly valid lectotype designation.

The lectotype, in the collections of the Natural History Museum, London,

BMNH C35625 from an unspecified horizon at the type section of the Mzamba

Formation at Mzamba Cliff is here refigured as Figures 63A-B, 64H-I.

Material

In addition to the lectotype, we have SAM-PCP5684, 5686, 8152-8154,

8361la, 8419-8422, 8653-7, 8660, SAM-7043, all from Bed A15 at locality 1

i ANNALS OF THE SOUTH AFRICAN MUSEUM

(see Klinger & Kennedy, 1980 figs 1-2), Pondoland, Transkei, Mzamba Forma-

tion, Campanian I; NMB D1663 from, an unspecified horizon at the same

locality. In addition, we also have examined the material collected by Van

Hoepen in 1919 and described in 1921, in the collections of the Transvaal

Museum, TM 540a~g, 548a-c, all from an unspecified horizon at this locality.

Recent excavations for a car park at the Wild Coast Casino a few kilometres

north of the Mzamba Estuary, have yielded additional specimens (Cooper

collection, MRC 1-39).

Dimensions

Spec. MxWb MxWh Wb/Wh MnWb MnWh Wb/Wh D_ Ti Ri

PCZ8660 4.0 Dal 0.7 1.5 1.6 0.9 IA MAS) 2 ==

PCZ8361 6.2, 8.6 OZ 4.1 5/5 0.74 32) OR Das)

PCZ5684 6.9 8.7 0.79 523 7.4 OZ LT 26 25

PCZ8656 TOF NOVO 0.7 5.0 6.9 0.73 31 OO as

PCZ8654 70 9.0 0.78 4.7 6.4 0.73 23 ihe

PEZ8i532.0 5 726) Ow 0.76 6.5 8.1 0.8 19° TiOe es

PCZ8421 Tad 9.8 0.79 1 8.0 0.84 18 10.0 —

PCZ8152 sol NAS 0.7 7.4 10263 07 19° 10:0) 23

PCZ8420 9.0 11.0 0.82 TD IOS OWS 24 4.2 3

PCZ8422 9.0 14.0 0.64 8.8 IZ OT 16 ~ 10:09 5

PEZ8AN9 ile le 0.73 8.3 Le Oeral 38 yar _-

RECZSi53b) 12a nO 0.72 9.1 SO OL 7/ 39. 10.2025

D1663 OW P1459 OZ 8.0 WO A078 34 Lies

TM 540e 120 8.4 0.83 Doll dell 0.8 36> -326 —

T™M 540c WV 9.8 0.74 5.6 7.9 0.71 28 6.9 —

TM 540a 8.0 10.6 OMS 6.0 Ut 0.83 38 89 2

TM 540g 0) SI OWS 7.0 9.0 0.78 2 Gee 2

TM 540b VAR to) 0.75 1007 1323 s0ei6 AZ Ss OD

MRC 6 POE AG) 0.64 — 9.0 — 34. 329 D

MRC 28 8.0 10.0 0.8 6.0 120 0.86 24. 25s

MRC 13 93, “1320 Oma Tell Os On 72 26n abes 2

MRC 12 EO Use) 0.81 — — — —_- — —

MRC 8 MMOe Maes 0.79 8.0 [120 0273 35 8.6 0.5

MRC7 14.0 19.0 0.74 WO SIO Osa 46 4.3 0

MRC 1 19.0 24.0 0.79 ARO) IO 0). 72! 67. Is PME

Description

Lectotype. The lectotype is a small individual, 22 mm long with a whorl

height of approximately 7 mm, partially embedded in soft, green silty matrix

Fig. 72 (see Hae page). Baculites sulcatus Baily, 1855. ?Late forms. A. MRC 24.

B. MRC6. C. MRC9. D. MRC 18. E-F. MRC17. G-H. MRC7. Note slight curvature

in A and E. a from excavation for new car park at Wild Coast Casino, Pondoland,

Mzamba Formation, Campanian. All x 2

CRETACEOUS FAUNAS FROM SOUTH AFRICA

2 12

114 ANNALS OF THE SOUTH AFRICAN MUSEUM

(Figs 63, 64H-I). The main features are the strong lateral ornament. The ribs

are crescentic, strongest on the dorsolateral part of the flanks, weaker on the

lower half and sweep forwards over the venter, forming a distinct chevron. The

impression of another (paralectotype) individual is present in the block of

matrix.

Topotype material. The initial discovery of B. sulcatus by Capt. Garden was

serendipitous. Detailed collecting by us yielded identical specimens only in the

topmost beds at the Mzamba Cliff. However, the underlying beds are difficult to

reach at this locality, and it is quite possible that B. sulcatus already occurs

lower down in the section, probably above the Santonian—Campanian boundary.

The following description is based on material from these topmost beds exposed

at the Mzamba Cliff, collected by us and the late Prof. T. Gevers, respectively.

SAM-7043 is part of a concretion with a complete specimen of Hauericeras

on the one side (Klinger & Kennedy 1980, fig. 5B), and a large body chamber

and part of a phragmocone of B. sulcatus on the other side (Fig. 64A). The

phragmocone fragment is identical to the lectotype, and we have no doubts in

identifying B. sulcatus from this bed. Another phragmocone fragment from the

same concretion, SAM-7043a shows the transition from the smooth to the

laterally ribbed stage at a whorl height of 7.5 mm.

Ornament in the rest of the material varies considerably—from extremely

robust, circumperipheral ribbing to virtually smooth. Ornament associated with

typical B. sulcatus is shown in PCP5684 (Fig. 64E-G), PCP5695 (Fig. 64B-D)

and PCP5684 (Fig. 64L-N), consisting of almost circumperipheral ribbing; ribs

strongest on the dorsal half of the flanks, and, in:sweeping forwards over the

ventral half of the flanks, bifurcate or occur with intercalatories, forming dis-

tinct chevrons and loops over the venter (Fig. 68D). Dorsally the ribs weaken

and curve broadly forward.

On the body chamber, as shown in SAM-7043 (Fig. 64A), ornament seems

to weaken, especially on the ventral half of the flanks. Lateral ornament consists

of two to three crescentic ribs per whorl height.

Other specimens have less prominent ornament, e.g. PCP8656 (Figs 64K,

65G-H), and only the upper half of the crescentic lateral ribs is visible and less

prominent than in the above-mentioned specimens. D1663 (Fig. 76D-E), a body

chamber fragment, also has very weak ribbing.

In the last group, e.g. PCP8361 (Fig. 65D-F) and PCP8653 (Fig. 65A-C),

ornament is very faint. PCP8153b (Fig. 65I-K) is a body chamber fragment

with part of the last septum preserved. Here, lateral ornament is practically

absent and only visible under very oblique, low lighting.

Transvaal Museum, Van Hoepen collection. Van Hoepen (1921: 18, pl. 3

(figs 7-8)) described and figured as B. sulcatus part of a collection of 22 speci-

mens. Unfortunately, we do not know exactly where in the Mzamba Cliff these

specimens were collected. Judging by the preservation, we presume them to be

Fig. 73 (see facing page). Baculites sulcatus Baily, 1855. ?Late forms. A-C. MRC 23.

D. MRC 22. E. MRC 34. F-H. MRC 28. I. MRC14. J. MRC30° Ko Mkee

L. MRC 19. M. MRC39. All from excavations for new car park at Wild Coast Casino,

Pondoland, Mzamba Formation, Campanian. All x 2.

115

CRETACEOUS FAUNAS FROM SOUTH AFRICA

ig. 73

116 ANNALS OF THE SOUTH AFRICAN MUSEUM

from below the topmost beds with typical B. sulcatus, but also well above the

basal beds with B. capensis. These specimens differ from typical B. sulcatus

mainly in that the lateral ribs form a more or less distinct tubercle on the dorsal

half of the flanks. In addition, in some of these specimens the whorl section

appears to be more elliptical, with the venter as wide as the dorsum and with

parallel flanks, reminiscent of B. capensis rather than ovoid as in typical

B. sulcatus.

However, as in typical B. sulcatus described above, strength and presence

of ornament is extremely variable. TM 540a (Fig. 66J-—L) and TM 540d

(Fig. 66O-P) have extremely strong ornament; that of TM 540g (Fig. 66A-B) is

much weaker, whereas TM 540e, TM 548a (Fig. 66G-I) and TM 548e

(Fig. 67C-D) are all practically smooth, save for ventral ribbing. TM 548c

(Fig. 67E-G) and TM 548d (Fig. 67H-J) are completely smooth. TM 548d

(Fig. 67H-J) and TM 548a (Fig. 66G-I) have parts of the aperture preserved.

This consists of a prominent dorsal rostrum, raised by a slight step, a very deep

lateral sinus, and a very prominent and long, spoon-shaped ventral rostrum.

Part of another aperture is preserved in TM 548b. These are all microconchs.

TM 540b (Fig. 66M-O), a body chamber, is a macroconch.

The suture lines of several specimens are exposed, e.g. Figs 68E, 69A-C,

7OA-C. These show some variation, but the saddles and lobes are distinctly

more incised and complex than those of B. capensis; in some, the bases of the

saddles and lobes are slightly constricted, resulting in a subtriangular shape.

The suture of one of the specimens, TM 540m (Fig. 70C), has rather narrow

saddles and resembles the suture of B. bailyi—as figured by Woods (1906,

pl. 44 (fig. 5)) and as mentioned above (p. 41).

Material from excavations at the Wild Coast Casino. Excavations for a new

car park at the Wild Coast Casino, north of the Mzamba River estuary,

yielded—amongst others—numerous baculitids best referable to B. sulcatus.

Again, ornament is quite variable, but in the majority of specimens consists of

distinct, widely spaced, crescentic ribs. These are strongest on the upper part of

the flanks, decrease in strength and width ventrally, and often cross the venter

as distinct lirae. In addition to these latter, intercalatory ribs or lirae may cross

the venter, some branching near the venter, forming distinct looped chevrons

over the venter (Figs 72-75, 76A-C).

In a few specimens, e.g. MRC 39 (Fig. 73), lateral ornament is as strong as

in typical B. sulcatus. In another group of about five specimens, lateral orna-

ment consists of obliquely elongated tubercles and ventral ribs, e.g. MRC 12_

(Fig. 75A-C), MRC 9, and MRC 4 (Fig. 73K). In others, e.g. MRC 24

(Fig. 72A), MRC 6 (Fig. 72B), MRC 20 (Fig. 74A) and MRC 16 (Fig. 74F),

lateral ornament consists of auricular, dorsolateral tubercles, projected ventrally

into ribs of variable strength. In a few specimens, e.g. MRC 35 (Fig. 74G),

ornament is completely absent.

Fig. 74 (see facing page). Baculites sulcatus Baily, 1855. ?Late form. A. MRC 20.

B. MRC 25. C. MRC la. D. MRC13. E. MRC38. F. MRC16. G. MR@35> Allizem

excavations for new car park at Wild Coast Casino, Pondoland, Mzamba Formation,

Campanian. All x 2, except C x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 7

Fig. 74

118 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 75. Baculites sulcatus Baily, 1855. ?Late form. A-C. MRC 12. D. MRC8, body

chamber specimen with aperture. Note distinct curvature of latter. Both from excavations

for new car park at Wild Coast Casino, Pondoland, Mzamba Formation, Campanian.

All ole

Two nearly complete apertures are preserved: MRC 1 (Fig. 76A-C) and

MRC 2 at maximum whorl heights of 20 mm and 24 mm, respectively. Most of

the specimens show a slight to pronounced curvature on the body chamber, e.g.

MRC 32, 33, 8 (Fig. 75D), MRC 24 (Fig. 72A), MRC 25 (Fig. 74B), MRC 2,

MRC 13, MRC 20 and MRC 1 (Fig. 76A-C). At first we thought that this was

due to diagenetic deformation, but it seems to be a primary feature. In others,

however, e.g. MRC 12 (Fig. 75A-C), the body chamber is perfectly straight.

For descriptive purposes, the specimens in the Transvaal Museum may be

regarded as early forms of B. sulcatus; those from the top of Mzamba cliff as

typical forms, and those from the Casino site as ?late B. sulcatus.

The suture lines and whorl sections of the latter are shown in Figure 77.

Discussion

Klinger & Kennedy (1977: 76) mentioned that B. sulcatus was a relatively

rare and poorly understood species. So far it is only definitely known from the

Campanian of the Mzamba Formation in Pondoland and from borehole material

in Richards Bay, Zululand. As yet, we have not found typical B. sulcatus in

outcrop in Zululand. Reports of B. sulcatus from Zululand (Mkweyane

(Umkwelane Hill)) by Spath (1921: 260) are incorrect; these are smooth forms

of B. capensis. Records of B. sulcatus from Madagascar (Collignon 1931: 36,

pl. 5 (figs 3-5, 13), pl. 9 (fig. 15)) are also wrong. These Madagascan

specimens are of late Coniacian age and probably variants of B. besairiei (=

B. yokoyamai).

We (Klinger & Kennedy 1975: 280) initially thought that B. vanhoepeni

Venzo, best known from Zululand, was a synonym of B. sulcatus. Indeed,

CRETACEOUS FAUNAS FROM SOUTH AFRICA 119

Fig. 76. Baculites sulcatus Baily, 1855. A-C. ?Late form. MRC 1, from excavations

for car park at Wild Coast Casino, Pondoland, Mzamba Formation, Campanian.

D-E. NMBD1663, ex Van Hoepen collection. Typical form from an unrecorded horizon

at locality 1, Pondoland, Mzamba Formation, Campanian. Both x 1.

Venzo (1936: 116 [58]) included the specimens described by Van Hoepen

(1921: 18, pl. 3 (figs 7-8)) as B. sulcatus in the synonymy of B. vanhoepeni.

Part of the problem in distinguishing between these two species lies in the

disparate sizes. Almost all our specimens of B. sulcatus are small whereas

nearly all our specimens of B. vanhoepeni are large. Very few specimens of

both species are of equal size for comparison. Small specimens of B. van-

hoepeni, e.g. Figure 109H, show lateral ornament comparable to that of typical

forms of B. sulcatus, but none shows as strong ventral or dorsal ribbing as in

typical B. sulcatus. Adult forms are easily distinguished: B. vanhoepeni has

typical widely spaced, auricular, lateral tubercles and grows to much larger size

than any known B. sulcatus. Lateral ornament in typical B. vanhoepeni is very

similar to that of ?late B. sulcatus, but again, none of these late forms of

120 ANNALS OF THE SOUTH AFRICAN MUSEUM

OOO

UQ9

O 5 mm

ea Sea ete

Fig. 77. Baculites sulcatus Baily, 1855. Late form. Whorl sections

and suture line. A-B. MRC7. C. MRC6. D. MRC3. E. MRC21.

F. MRC 23. E. MRC20. Venter in whorl sections pointing downward.

Scale bar for size.

B. sulcatus grows to as large a size as B. vanhoepeni, nor is ornament ever as

prominent. Also, the two species appear to differ in age: B. sulcatus appears

restricted to the Lower Campanian, whereas B. vanhoepeni is a typical Middle

or even partly Upper Campanian species, occurring in association with

Australiella and Menabites.

The origins of B. sulcatus are not quite clear. The early forms of B. sul-

catus, with distinct lateral nodes and elliptical whorl section, are very remi-

niscent of B. capensis. The occurrence of the latter in the basal beds of the

Mzamba Formation would support the derivation of B. sulcatus from

B. capensis. It is possible that the transition from B. capensis to B. sulcatus

took place via some of the forms described from the Lower Campanian of

CRETACEOUS FAUNAS FROM SOUTH AFRICA 12

Madagascar by Collignon (1969)—as B. cf. tanakae (Collignon 1969: 23,

pl. 521 (fig. 2055)) or B. sparsinodosus Collignon (Collignon 1969: 23, pl. 521

(figs 2056-8 only)), with crescentic dorsolateral tubercles.

Looking at typical forms of B. sulcatus, however, especially the weakly

ornamented forms, it would alternatively seem possible to derive the former

from late forms of B. bailyi. Some specimens of B. bailyi from the basal beds of

the Mzamba Formation, e.g. SAM-PCP8729 (Fig. 670-Q), look remarkably

like weakly ornamented B. sulcatus.

The specimens of ?late B. sulcatus from the Casino excavations bear a strik-

ing resemblance to an Upper Santonian baculitid fauna recently described from

the U.S. Gulf Coast region by Kennedy & Cobban (1991b) as B. capensis and

Boehmoceras arculus. Kennedy & Cobban used the name B. capensis (1991b:

182, figs 6: 4, 8: 1-8; 10: 7-10, 12-14; 12: 2.5) for straight baculites with

dorsolateral nodes, and Boehmoceras arculus (1991b: 182, figs 6: 2, 8; 8: 9-15,

18-22; 9: 1-2, 11-52; 10: 20-21, 24-26; 12: 3) (a senior synonym of Boehmo-

ceras loescheri Riedel, 1931) for predominantly curved specimens with distinct,

crescentic to auricular lateral bullae that extend across the dorsolateral half to

two thirds of the flanks.

As far as lateral ornament is concerned, the Pondoland specimens are indis-

tinguishable from the Gulf Coast material. MRC 1 (Fig. 76A-C) or MRC 8

(Fig. 75D) are indistinguishable from specimens assigned to B. arculus by Ken-

nedy & Cobban (19915, fig. 9: 50-52 or fig. 9: 46), but none of our specimens

has as distinct curvature as the Gulf Coast material to merit assignation to the

genus Boehmoceras.

It is obvious that in both assemblages we are dealing with a baculitid popu-

lation morphologically transitional between Baculites and Boehmoceras, with

both having their origins in B. capensis. As shown above (p. 71), some

specimens of B. capensis have slightly curved body chambers. In the majority

of the Pondoland specimens, ornament is of the Boehmoceras arculus or

Baculites sulcatus type, but curvature is restricted to the body chamber, as in

B. capensis, and they still belong to Baculites s.s. In the Gulf Coast material,

lateral ornament is the same, but the curvature is more distinct and already

occurs on the phragmocone, and they are referable to Boehmoceras. In both

populations, more or less straight specimens with B. capensis-like ornament

occur associated with the curved specimens, indicating an origin in the latter

species. It is tempting to link B. capensis to the Gulf Coast Boehmoceras fauna

via the Pondoland specimens but, lacking a precise date for the latter

assemblage, this is not advisable at present. Affinities of ?late B. sulcatus with

typical forms of B. sulcatus, rather suggests that the Pondoland and Gulf Coast

faunas have a common origin in the Upper Santonian in B. capensis, and

subsequently acquired similar ornament, but developed in parallel. The Gulf

Coast fauna became progressively more curved and gave rise to true

Boehmoceras, whereas the Pondoland fauna remained in the strict Baculites

lineage.

Baculites increscens Collignon (1970: 3, pl. 607 (figs 2266-2268)) from the

Middle Campanian of Zululand and Madagascar resembles ?late B. sulcatus, but

lateral ornament in the former is more prominent, and transitional to the

prominent auricular B. vanhoepeni type.

122 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 78. A-B. Baculites bailyi Woods, 1906. A. SAM-PCZ8358, concretion with B. bailyi

and Premuniericeras from locality 98, Zululand, St Lucia Formation, Santonian II.

B. SAM-PCZ7214, concretion crammed with juvenile B. bailyi from locality 73, Zululand,

St Lucia Formation, Coniacian IV or V. C. B. sulcatus Baily, 1855. SAM -PCP836l,

concretion from locality 1, Pondoland, Mzamba Formation, Campanian I. ©

CRETACEOUS FAUNAS FROM SOUTH AFRICA 123

Strongly ornamented forms of typical B. sulcatus bear superficial

resemblance to Trachybaculites columna (Morton, 1834) (see Cobban &

Kennedy 1992c for a recent review). Trachybaculites columna, however, is a

Maastrichtian species known from the Prairie Bluff Chalk of Alabama, the

Corsicana Marl of the Navarro Group of Texas, Trail City Member of the Fox

Hills Formation in South Dakota, and also from the Ganzas Formation of the

San Joaquin Valley, California (Matsumoto 1959: 163). Ribbing seems to be

more uniform, and stronger on the dorsum and venter than in B. sulcatus. Also,

the suture is distinctly simple compared to that of B. sulcatus.

Baculites furcillatus (Blanckenhorn, 1905, pl. 6 (fig. 2)) (herein Fig. 67E-F)

has ornament similar to that of some B. sulcatus. This is a rare species in the

Maastrichtian of Israel, and more material is needed for definite comment.

According to Picard (1929: 441), B. furcillatus ‘is only a strongly ribbed form

of B. palestinensis’. Ribbing in B. furcillatus is stronger over the venter and

dorsum than in B. sulcatus, and thus seems closer to 7. columna.

Baculites thomi Reeside (1927b: 13, pl. 12 (figs 9-14)) from the Elk Basin

Sandstone member of the Telegraph Creek Formation of the U.S. Western

Interior, is superficially similar to some typical B. sulcatus in the strong, cir-

cumperipheral ribbing. However, a recent review of the species by Cobban &

Kennedy (1991la: C1-C8, pls 1-2) showed that Reeside’s original figures of the

holotype were retouched. According to Cobban & Kennedy (1991a), B. thomi

first appears in the Upper Santonian, where it reaches its peak, and persists into

the Lower Campanian. Ornament in B. thomi is far more regular than in

B. sulcatus, with very regular ventral corrugations developing in typical forms.

Also, the suture line (Cobban & Kennedy 1991a, fig. 2A—B) is less complex.

Some representatives of two baculitid species from the Coniacian of

the U.S. Western Interior are very similar to B. sulcatus. These are B. sweet-

grassensis Cobban, 1951 (see Kennedy & Cobban 1991la: 70, pl. 14

(figs 24-25, 29-34, 38-42)) and B. codyensis Reeside (see Kennedy & Cobban

1991a: 72, pl. 15 (figs 1-30), pl. 16 (figs 1-13), pl. 17 (figs 1-8), text-fig. 25F).

Baculites sweetgrassensis is a Middle Coniacian species, and, in typical

forms, has widely spaced, crescentic bullae that are projected ventrally into

sharp, forwardly direct ribs. Superficially some of these (e.g. Kennedy &

Cobban 1991a, pl. 14 (fig. 40-41)) resemble early forms of B. sulcatus; others,

(e.g. Kennedy & Cobban 1991a, pl. 14 (figs 30, 38)) resemble ?late B. sulcatus.

Baculites codyensis first appears in the Middle Coniacian, and ranges up to

the Middle Santonian. Typical forms have regular, concave, concentric ribs,

and these, together with the age difference clearly separate B. codyensis from

B. sulcatus. Some of the more coarsely ornamented forms of B. codyensis (see

e.g. Kennedy & Cobban 199la, pl. 16 (fig. 12)) resemble ?late forms of

B. sulcatus, but again, this is mere homoeomorphy.

Some specimens of B. ovatus Say, recently figured by Kennedy & Cobban

(1993c, fig. 15. 9-12), are remarkably similar to our ?late B. sulcatus, but

B. ovatus is a younger species, and typically has weaker ornament than

B. sulcatus.

Baculites oberholzeri Bohm (1909: 52, pl. 1 (fig. 9a—b)), imprecisely dated

from the Senonian of Switzerland, appears to have similar lateral ornament, but

this species is based on a crushed fragment, about 11 mm long, and is best

regarded as a nomen dubium.

124 ANNALS OF THE SOUTH AFRICAN MUSEUM

Some specimens described and figured by Birkelund (1965: 58, pl. 8

(fig. 1), pl. 9'iigs 1-3), pl. 10: Gig. 1), pl. 11) Gigs 1-2), pl. 12 Gigs Ie 2)alans

(figs 1-2), pl. 14 (fig. 1), text-figs 47-52) as Baculites obtusus Meek from the

Lower Campanian of West Greenland resemble B. sulcatus, but none of the

specimens develops as strong ornament as typical representatives of the latter.

Typical forms of B. sulcatus are also nearly identical with some

B. leopoliensis Nowak (see e.g. Kennedy 1986e, pl. 2 (figs 11—12)); this,

however, is an Upper Campanian species, and typical forms have more regular,

thin, crescentic ribs.

Occurrence

To date, B. sulcatus has only been recorded from the Lower Campanian of

Pondoland and in subsurface deposits at Richards Bay, Zululand.

Baculites increscens Collignon, 1970

Figs 79A-L, N-O, 80A-B, 82

1970 ~~ Baculites increscens Collignon, p. 3, pl. 607 (figs 2266-2268), p. 5, pl. 608

(fig. 2269).

1970 —_ Baculites tanakaeformis Collignon, p. 2, pl. 607 (figs 2263-2265).

1970 — Baculites androtsyensis Collignon, p. 5, pl. 608 (figs 2270-2272).

1970 = Baculites mamillatus Collignon, p. 7, pl. 609 (figs 2273-2274).

Type |

Holotype, by original designation, the specimen figured by Collignon (1970,

pl. 607 (fig. 2266)) from the Middle Campanian, Zone of Pachydiscus

grossouvrei, subzone of Eupachydiscus lamberti, gisement 177, Coupe

d’ Ankilizato (Belo sur Tsiribihina), Madagascar, housed in the collections of the

Institut des Sciences de la Terre, Université de Dijon, GD 12266, here refigured

as Figure 82.

Material

SAM-PCZ11986-11996, all from locality 102, Zululand, St Lucia

Formation, Campanian II.

Description

All of our material is fragmentary, but the specimens range in size from

juveniles to large body chamber sections, covering all the growth stages.

PCZ11992 (Fig. 79I-J) is a juvenile and shows distinct crescentic,

dorsolateral nodes. PCZ11995 at similar whorl height has less conspicuous

nodes but shows a distinct trigonal whorl section. PCZ11996, at whorl height

Fig. 79 (see facing page). A-L, N-O. Baculites increscens Collignon, 1970.

A-D. SAM-PCZ11988. E. SAM-PCZ11991. F-H. SAM-PCZ11997. I-J. SAM—

PCZ11992. K-L. SAM-PCZ11987. N-O. SAM-PCZ11993, all from locality 103,

Zululand, St Lucia Formation, Campanian ?IJ. M. Baculites vanhoepeni Venzo, 1936.

SAM-PCZ11998 from locality 110, Zululand, St Lucia Formation, Campanian II or III.

Al x A:

CRETACEOUS FAUNAS FROM SOUTH AFRICA

125

Fig. 79

126 ANNALS OF THE SOUTH AFRICAN MUSEUM

O 10 mm

ie eet sterner |

Fig. 80. Whorl sections and suture lines. A-B. Baculites increscens Collignon,

1970. A. SAM-PCZ11995. B. SAM-PCZ11986. C-D. Baculites duharti

Hiinicken, 1965. C. SAM-PCZ7685. D. SAM-PCZ7685. Venter in whorl

section pointing upward. Scale bar for size.

12 mm, shows a similar rounded, trigonal whorl section. PCZ11993

(Fig. 79N—-O) and PCZ11986 show typical phragmocone ornament consisting of

prominent, dorsolaterally situated auricular nodes, projected ventrally into thin

lirae that cross the venter, accompanied by intercalated lirae. Slightly more than

two nodes occur per whorl height. The whorl section is compressed ovoid

(Fig. 80A—C)with the dorsum flat to slightly rounded, and the venter narrowly

rounded. PCZ11990 is a fragment, and has large, rounded, instead of crescentic

dorsolateral nodes.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 127

UPPER CAMPANIAN

Zone of Delawarella subdelawarensis ae taylorensis Adkins

and B. ankilizatensis sp. nov.

(153-159)

Australiella australis B. rectangulatus sp. nov.

(156-157)

Sub- B. ankilizatensis sp. nov.

oe (first: 153-159)

Pachydiscus bassae Foe ae Nowak

Zone of

B. coagmentatus sp. nov.

(177-156)

B. increscens

(last: 177)

B. androtsyensis sp. nov.

Eupachyoiscus 180 & 327-328-329)

B. mamillatus sp. nov.

lamberti (181-326)

B. increscens

(first: 180-181)

B. tanakaeformis sp. nov.

(326-327 & 180-181)

Pachydiscus grossouvrei Sub-zone of

= 4

Top of Lower Campanian:

Sub-zone of Termiericeras /enticulare

LOWER CAMPANIAN

Fig. 81. Biostratigraphic zonation of the Middle Campanian and distribution of

Baculites spp. at Menabe, Madagascar after Collignon (1969).

PCZ11991 (Fig. 79E), PCZ11988 (Fig. 79A-C) and PCZ11989 are body.

chamber fragments. Lateral ornament varies from closely spaced crescentic

ribs, to low and rounded nodes. Some fragments appear to have been smooth.

Discussion

Baculites increscens is but one of several baculitids described by Collignon

(1970) from the Middle Campanian of Madagascar. Our material fits the

description of B. increscens best, but several features ascribed to the other

Madagascar species are also present, and it is necessary to comment on the

latter.

According to Collignon’s (1970) stratigraphy of the Middle Campanian of

Menabe, Madagascar (Fig. 81) the majority of Baculites species occur in the

lower part of the substage, in the Zone of Pachydiscus grossouvrei, Subzone of

Eupachydiscus lamberti. These include B. coagmentatus Collignon (1970: 7,

ANNALS OF THE SOUTH AFRICAN MUSEUM

128

on, 1969. Plaster cast of the holotype, GD12266, the

2266)) from the Middle Campanian of

Fig. 82. Baculites increscens Collign

ur Tsiribihina), Madagascar. * 1

original of Collignon (1970, pl. 607 (fig.

gisement 177, Coupe d’ Ankilizato (Belo s

CRETACEOUS FAUNAS FROM SOUTH AFRICA 129

pl. 609 (figs 2275-2276)) (herein Fig. 88); B. increscens Collignon (1970: 3,

pl. 607 (figs 2266-2268)) (herein Fig. 82); B. androtsyensis Collignon (1970: 5,

pl. 608 (figs 2270-2272)) (herein Fig. 83), B. mamillatus Collignon (1970: 7,

pl. 609 (figs 2273-2274)) (herein Fig. 84A-C) and B. tanakaeformis Collignon

(1970: 2, pl. 607 (figs 2263-2265)) (herein Fig. 84D-F).

The order of their listing below suggests that this is the order of their occur-

rence, but this is not strictly so. If they are arranged according to their

occurrence at the different localities or horizons (Collignon’s ‘gisements’), it is

clear that these species are contemporary to a large extent.

‘Gisement’

183 Baculites cf. B. taylorensis

153 Baculites ankilizatensis

Baculites coagmentatus

156 Baculites rectangulatus

Baculites leopoliensis

LS7 Baculites rectangulatus

159 Baculites ankilizatensis

Nyy Baculites leopoliensis

Baculites coagmentatus

Baculites increscens

180 Baculites androtsyensis

Baculites increscens

Baculites tanakaeformis

181 Baculites mamillatus

Baculites increscens

Baculites tanakaeformis

329 Baculites androtsyensis

328 Baculites androtsyensis

32] Baculites androtsyensis

Baculites tanakaeformis

326 Baculites tanakaeformis

Baculites mamillatus

According to Collignon (1970: 2), B. tanakaeformis occurs at the base of the

Middle Campanian, and B. androtsyensis is the oldest of the series of baculitids

that dominates the Middle Campanian (Collignon 1970: 5). Both occur together

at gisement 327; at gisement 181 they occur together with B. increscens; and at

326 and 181 B. tanakaeformis occurs with B. mamillatus.

On the basis of their co-occurrence, it would seem logical to regard the

above-mentioned four species, B. tanakaeformis, B. increscens, B. androt-

syensis and B. mamillatus as a single, variable species. However, Collignon

130 ANNALS OF THE SOUTH AFRICAN MUSEUM

based these species on large assemblages—B. tanakaeformis (87 specimens),

B. androtsyensis (50 specimens) and B. increscens (50 specimens), and they

cannot summarily be dismissed as synonyms.

Fig. 83. Baculites androtsyensis Collignon, 1969. Plaster cast of the holotype, GD12270,

the original of Collignon (1970, pl. 608 (fig. 2270)), from the Middle Campanian

of gisement 329, Coupe Ampolypoly-Antsirasira-Behamotra (Belo sur Tsiribihina),

Madagascar. x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 131

All four species have a subtriangular whorl section in common, with faint

indications of longitudinal depressions on either side of the venter, indicating an

incipient ventral keel. Lateral ornament varies from strong crescentic to

mammiform in B. increscens, weaker in B. androtsyensis (with more

compressed whorl section), distinct mammiform in B. mamillatus, conical to

obliquely elongated in B. tanakaeformis. The suture lines of B. mamillatus and

B. androtsyensis (Figs 85A, C) are very similar.

We do not doubt that these four broad morphological groups can be

identified in large collections, and that some morphotypes probably are more

common at certain localities than others in Madagascar. However, our Zululand

material from locality 102 shows that even though the majority of specimens

are identifiable with B. increscens, some specimens could be referred to either

of the other three species, e.g. PCZ11993 (Fig. 79N-O) and PCZ11986 have a

more compressed whorl section as in B. androtsyensis; PCZ11988

(Fig. 79A—-D) has rounded tubercles as in B. mamillatus; those of PCZ11991

(Fig. 79E) are as in B. tanakaeformis. Because of this, we doubt if it is justi-

fiable to maintain formal specific names for these different, yet overlapping

and co-eval morphotypes and, as first revising authors, we select

Baculites increscens as the name of this species.

Baculites coagmentatus Collignon (1970: 7, pl. 609 (figs 2275-2276))

(Fig. 88) appears to be slightly younger than most of the baculitids of the group

of B. increscens, but co-occurs with the last appearance of B. increscens at

Collignon’s locality 177. The whorl section is still subtriangular as in

B. increscens, but the ornament of B. coagmentatus consists essentially of thin,

closely spaced, crescentic ribs—a mode of ornament found in some of the

Durban specimens of B. vanhoepeni to be described below. Baculites

leopoliensis Collignon (1970: 10, pl. 610 (figs 2277-2278) non Nowak) (herein

Fig. 86) has a more ovoid whorl section, but similar lateral ornament as in

B. coagmentatus and they are probably synonyms. Baculites leopoliensis Nowak

(1908: 328, pl. 14 (figs 1-5, 10, 211), text-figs 1-5 on p. 329, ?text-figs 5-10

on p. 331) is a typical Upper Campanian species (see Hancock & Kennedy

1993: 165) and not Lower Maastrichtian as recorded earlier by, for example,

Kennedy (1986c: 1013). The ornament consists of lateral ribs that break down

into riblets and striae on the venter, whereas intercalated ribs develop over the

venter. This style of ornament is quite characteristic and unlike that of the

Madagascar material. Baculites leopoliensis Collignon non Nowak is here

renamed B. collignoni nom. nov. Baculites coagmentatus and B. collignoni have

a style of ornament similar to some forms of B. vanhoepeni (see e.g.

Fig. 111A-—C), and these two species can probably be regarded as ?early forms

(and junior synonyms?) of B. vanhoepeni.

Fig. 84 (see overleaf). A-C. Baculites mamillatus Collignon, 1970. Plaster cast of the

holotype, GD12273, the original of Collignon (1970, pl. 609 (fig. 2273)) from the Middle

Campanian of gisement 181, Coupe d’Ankilizato (Belo sur Tsiribihina), Madagascar.

D-F. Baculites tanakaeformis Collignon, 1970. Plaster cast of the holotype, GD12263, the

Original of Collignon (1970, pl. 607 (fig. 2263)) from the Middle Campanian of

gisement 326, Coupe Ampolypoly-Antsirasira-Behamotra (Belo sur Tsiribihina), Madagascar.

Both x 1.

132

ANNALS OF THE SOUTH AFRICAN MUSEUM

CRETACEOUS FAUNAS FROM SOUTH AFRICA 133

l é EN

ql b§ Bide

E L

U I

O 10 mm B

ob ie

Mt Slae

een fs

Fig. 85. Suture lines. A. Baculites mamillatus Collignon, 1970. GD12273.

(see Fig. 84A-C). B. Baculites ankilizatensis Collignon, 1970. GD12282

(see Fig. 89). C. Baculites androtsyensis Collignon, 1970. GD12270 (see

Fig. 83). Scale bars for size.

Baculites rectangulatus Collignon (1970: 12, pl. 611 (figs 2279-2281))

(herein Fig. 87) is a late representative of the group of B. increscens as far as

Ornamentation is concerned. The whorl section, however, is more compressed

and subrectangular. Baculites ankilizatensis Collignon (1970: 13, pl. 612

(figs 2282-2284)) (herein Fig. 89) is a large baculitid with reduced lateral

ornament.

In Zululand, B. increscens appears to occupy an intermediate position

between B. sulcatus, especially the material from the Casino excavations, and

B. vanhoepeni. Juvenile specimens of B. increscens are virtually

indistinguishable from ?late B. sulcatus. Baculites increscens differs from

B. sulcatus mainly in having stronger lateral ornament in the adult stage, and

from B. vanhoepeni in being weaker ornamented. To be quite frank, the main

reason for separating B. increscens in Zululand is probably because of its

isolated occurrence at locality 102.

134 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 86. Baculites collignoni nom. nov. Plaster cast of GD12277, the original of Baculites

leopoliensis non Nowak in Collignon (1970, pl. 610 (fig. 2277)) from gisement 156, Coupe

d’Ankilizato (Belo sur Tsiribihina), Madagascar. x 1. )

CRETACEOUS FAUNAS FROM SOUTH AFRICA 135

Fig. 87. Baculites rectangulatus Collignon, 1970. Plaster cast of the holotype, GD12279,

the original of Collignon (1970, pl. 611 (fig. 2279)) from the Middle Campanian of

gisement 157 of Coupe d’Ankilizato (Belo sur Tsiribihina), Madagascar. x 1.

136 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 88. Baculites coagmentatus Collignon, 1970. Plaster cast of the holotype, GD12275,

the original of Collignon (1970, pl. 609 (fig. 2275)) from the Middle Campanian of

gisement 177, Coupe d’Ankilizato (Belo sur Tsiribihina), Madagascar. x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 137

Fig. 89. Baculites ankilizatensis Collignon, 1970. Plaster cast of the holotype, GD12282,

the original of Collignon (1970, pl. 612 (fig. 2282)) from the Middle Campanian of

gisement 153, Coupe d’Ankilizato (Belo sur Tsiribihina), Madagascar. x 1.

138 ANNALS OF THE SOUTH AFRICAN MUSEUM

Baculites nibelae sp. nov., described below (p. 162) is younger than typical

B. increscens, lateral ornament is similar, but the whorl section is character-

istically wedge-shaped,trigonal, with a flat dorsum and a narrow venter.

Occurrence

Middle Campanian of Zululand and Madagascar.

Baculites vanhoepeni Venzo, 1936

Figs 79M, 91-115

1936 Baculites vagina Forbes var. van Hoepeni Venzo, p. 116 [58], pl. 10 [6]

(figs 11-12).

1973 ~Baculites sp. group of Baculites capensis Woods; Kennedy & Klinger, p. 100,

pl. 4 (figs 1-5), pl. 5 (fig. la-d), pl. 6 (figs 4-5).

1975 — Baculites sulcatus (non Baily); Kennedy & Klinger, p. 280.

? 1970 —Baculites coagmentatus Collignon, p. 7, pl. 609 (figs 2275-2276).

v 1970 Baculites leopoliensis non Nowak, Collignon, p. 10, pl. 610 (figs 2277-2278)

(= B. collignoni nom. nov.)

1977 —Baculites vanhoepeni Venzo; Kennedy & Klinger, p. 73, figs 2G-K, 3A, H-I,

4A-C, 5C.

Kwa-Nibela

109B—>>*

KOSIE—=—|

Hell's Gate

1000 2000 Metres

Fig. 90. Localities on Nibela Peninsula referred to in text, but

not included in Kennedy & Klinger (1975).

CRETACEOUS FAUNAS FROM SOUTH AFRICA 139

Type

Venzo (1936: 116 [58]) based this species on several specimens. Klinger &

Kennedy (1977: 73) designated the larger of the two specimens figured by

Venzo (1936, pl. 10 [6] (fig. 1la—b)) from ‘False Bay’, Zululand, as lectotype.

It is housed in the Department of Geological Sciences, University of Bologna,

no. | GO243.

Material

We have more than 150 specimens from localities on the Nibela Peninsula

(Fig. 90), Zululand, St Lucia Formation, Campanian II-III.

Dimensions

A list of dimensions is given in the appendix.

Max Wb (mm) MxWb/MxWh MnWb/MnwWh Ti Ri

10-14.9 0.71 0.69 5.32 0.81

fs—19.9 0.71 O73 4.7 0.8

20-24.9 O77 0.76 4.9 ferles

25-27.9 0.77 0.79 328 1.28

30-34.9 Doe Osi 6.2 l

37 0.80 _ — _

Description

This species is conspicuous by its large size; the largest available specimen

(D1302b, Fig. 103) has a whorl height of 55 mm. Unfortunately, the early

stages are poorly known—almost all our specimens are large. This is probably

due to taphonomic and/or diagenetic factors.

Early stages of growth are shown in PCZ7714 (Fig. 109H). Here the whorl

section is more or less ovoid, with a narrow venter. Ornament consists of

dorsolaterally situated crescentic ribs which are projected ventrally and

aperturally over the venter, numbering about 2.5 per whorl height.

All the characteristic features of the species are shown in PCZ8764

(Fig. 91A-C). The bullae are typically crescentic to auricular, with sharp to

rounded lateral crests, and are situated on the dorsal half of the flanks.

Ventrally, they are projected acutely forwards and over the venter as thin

riblets. In addition, a variable number of intercalatories arises on the ventral

half of the flanks, and these also cross the venter with distinct, albeit variable

corrugation.

Fig. 91 (see overleaf). Baculites vanhoepeni Venzo, 1936. A-C. SAM-PCZ8764.

D-F. SAM-PCZ7717. G-I. SAM-PCZ7415. J-L. SAM-PCZ7707. M-O. SAM-

PCZ7091. Note the irregular occurrence of ventral corrugations. All from locality 110,

Zululand, St Lucia Formation, Campanian II-?1II. All x 1.

Fig. 92 (see overleaf). Baculites vanhoepeni Venzo, 1936. A-C. NMBD1467.

D. SAS Z1911. E-F. NMBD1467a. G-H. SAS Z1863e. Note the aperture in D and H. All

from locality 110, Zululand, St Lucia Formation, Campanian II-?III. All x 1.

ANNALS OF THE SOUTH AFRICAN MUSEUM

140

Wy “yg

Vy j

CRETACEOUS FAUNAS FROM SOUTH AFRICA 14]

Fig. 92

142 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 93. Baculites vanhoepeni Venzo, 1936. A-C. SAS A2035. Large, adult specimen

with typical prominent auricular lateral tubercles. From locality 110, Zululand St Lucia

Formation, Campanian II-?III. x 1.

Fig. 94 (see facing page). Baculites vanhoepeni Venzo, 1936. A-C. SAS A2032.

D. SAS A477. E. SAM-PCZ7411. F. SAM-PCZ8765. All from locality 110, Zululand,

St Lucia Formation, Campanian I]-?111. All x 1. iy

143

CRETACEOUS FAUNAS FROM SOUTH AFRICA

? Fh 3

ema

Fig. 94

144 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 95. Baculites vanhoepeni Venzo, 1936. A-C. SAM-PCZ7635. D-F. SAS A2034.

Both from locality 110, Zululand, St Lucia Formation, Campanian II-?III. Both x 1.

Fig. 96 (see facing page). Baculites vanhoepeni Venzo, 1936. A. SAM-PCZ7640.

B. NMBD1302b. C-E. NMBD1302a. All from locality 110, Zululand, St Lucia

Formation, Campanian II-?1II. All x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 145

Fig. 96

146 ANNALS OF THE SOUTH AFRICAN MUSEUM

The dorsal projection of the bullae is striking. Viewed dorsally, their outline

is pointed, mammiform and, together with the dorsal ribbing, forms an open-

ended, inverted V. The whorl section varies from ovoid at the smaller end to

elliptical at the larger.

The essential features of this species are the widely spaced, dorsolateral

auricular bullae. In the adult stage, they are distinctly auricular in lateral view

and conical mammiform in dorsal view. (Figs 96D, 99C). These are generally

widely spaced, about one per twice the whorl height, but in some specimens,

especially on the body chamber, they may be more closely spaced—up to three

per whorl height (e.g. SAS A477—Fig. 94D), or even more widely spaced

(e.g. PCZ9404, Fig. 102C), but this is exceptional.

There is some variation in the shape of the bullae. At the one extreme they

are reduced to thin riblets on the flanks, e.g. Fig. 111A—C; at the other they are

low and rounded to bullate, rather than transversely elongated, e.g. SAS A2034

(Fig. 9SD-F), PCZ7640 (Fig. 96A), and Figures 110A-C, 112A-C, D, E-F.

Specimens from locality 109A (Fig. 114A-—C) have more widely spaced, smaller

and rounded tubercles. We suspect that these are slightly older than typical

forms at locality 109 and 110. These could also possibly be referred to as ?early

forms of B. vanhoepeni, or, given more material, possibly be separated

specifically.

The presence of riblets on the ventral half of the flanks and accompanying

corrugation on the venter is an extremely variable feature and of no apparent

taxonomic value. In some examples, e.g. PCZ8764 (Fig. 91A-C), PCZ7717

(Fig. 91D-F), PCZ1860h (Fig. 101D), PCZ1911 (Fig. 101F), these are vqume

distinct, but in the majority of specimens they are absent or very faint. The

majority of specimens with ventral ribbing are small, but as shown in Figure 91,

other specimens of similar size totally lack these ventral ribs. In short, ventral

ribbing appears to be a variable character, sometimes, but not always associated

with early stages of growth, and taxonomically of no significance in

B. vanhoepeni.

The aperture is preserved in several specimens, e.g. Z1911 (Fig. 92D),

Z1863e (Fig. 92G-H), Z2077 and consists of a short, rounded dorsal rostrum, a

prominent lateral sinus and a longer ventral rostrum. Distribution of maximum

whorl height (Fig. 113) suggests that B. vanhoepeni is dimorphic, but that some

overlap in size occurs between macro- and microconchs. In one specimen,

SAM-PCZ7091 (Fig. 91M-O) part of the body chamber is distinctly flared.

This is very similar to a specimen of B. obtusus figured by Birkelund (1965,

pl. 9 (fig. 3a—c)).

Fig. 97 (see facing page). Baculites vanhoepeni Venzo, 1936. A. SAS Z1860a.

B. SAS A2029b. C. SAM-PCZ7655. D. SAS Z1860b. All from locality 110, Zululand,

St Lucia Formation, Campanian II-?III. All x 1.

Fig. 98 (see overleaf). Baculites vanhoepeni Venzo, 1936. A. SAS Z1190. B. SAS Z1860c.

Both from locality 110, Zululand, St Lucia Formation, Campanian II-?III. Both x 1.

Fig. 99 (see overleaf). Baculites vanhoepeni Venzo, 1936. A. SAS Z1911. Specimen with

typical adult lateral ornament. D. SAS A2029a. E. SAM-PCZ12020. F. NMBD1302a.

All from locality 110, Zululand, St Lucia Formation, Campanian IJ-?11I. All x 1.

CRETACEOUS FAUNAS FROM SOUTH AERICA

147

Fig. 97

148 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 98

CRETACEOUS FAUNAS FROM SOUTH AFRICA 149

Fig. 99

150 ANNALS OF THE SOUTH AFRICAN MUSEUM

The suture line of B. vanhoepeni (Figs 104-108) is slightly more complex

than that of the majority of the Lower and Middle Campanian baculitid species

from Madagascar (Figs 61-62, 85) and quite variable. The lateral lobe (L) tends

to become constricted. In some, e.g. Z1923 (Fig. 108A), the suture is highly

incised and dendritic, with narrow-stemmed saddles and a prominent,

asymmetrically trifid, constricted lateral lobe, whereas in others (e.g.

SAM-PCZ7706—Fig. 104A) it is relatively simple with quadrate saddles and

lobes with open (unconstricted) lobes.

Discussion

This species was originally described as B. vagina var. van Hoepeni by

Venzo (1936: 116 [58], pl. 10 [6] (figs 11-12)). Venzo incorrectly included the

Pondoland specimens described and figured by Van Hoepen (1921: 18, pl. 3

(figs 7-8)) as B. sulcatus in the synonymy of this species. Baculites vagina is a

typical Maastrichtian Eubaculites, and totally different from the present species,

bituberculate, with a tabulate venter (see e.g. Klinger & Kennedy 1993).

Baculites sulcatus, as here interpreted, is an older species, occurring in the

Lower Campanian of Pondoland, and typically lacks the strong lateral ornament

of B. vanhoepeni.

Baculites vanhoepeni is easily identified by the distinct, auricular

dorsolateral ornament, especially in the adult stage, and, most conspicuously,

by its large size, which distinguishes it from all the older Lower and other

Middle Campanian baculitid species of Zululand and Madagascar.

Juvenile stages of B. vanhoepeni are rare, probably for taphonomic and/or

diagenetic reasons. Available specimens (Fig. 109H) are indistinguishable from

similarly sized specimens of B. increscens (Fig. 79I-J) or B. sulcatus (Fig. 731).

This suggests a phylogenetic lineage starting with B. capensis, via B. sulcatus

and B. increscens or similar forms, to B. vanhoepeni. The adult stages,

however, are clearly different—both in terms of ornament and maximum adult

SIZe.

As mentioned above, there is some variation in lateral ornament. Typical

specimens have widely spaced, auricular bullae but, in some, the bullae are

reduced to thin lateral riblets (Fig. 111A-D), or the bullae are more closely

spaced and sharp crested (Fig. 94D), whereas in some the bullae are large and

rounded and mammiform (Figs 95D-F, 96A). Some of these morphologies are

already present in some of the Lower and Middle Campanian baculitids of

Madagascar described by Collignon (1969, 1970).

Fig. 100 (see facing page). Baculites vanhoepeni Venzo, 1936. A-B. SAS Z1860d.

C-E. SAM-PCZ7645. Both from locality 110, Zululand, St Lucia Formation,

Campanian II-?HI. Both x 1.

Fig. 101 (see overleaf). Baculites vanhoepeni Venzo, 1936. A. SAM-PCZ7786.

B. SAM-PCZ7690. C. SAS A2036a. D. SAS Z1860h. E. SAS A2036. F. SAS Z1911.

G. SAS Z1923. H. SAS Z1860f. All from locality 110, Zululand, St Lucia Formation,

Campanian II-?1T. All x 1.

Fig. 102 (see overleaf). Baculites vanhoepeni Venzo, 1936. A-B. SAS Z1860b.

C. SAM-PCZ9404. Both from locality 110, Zululand, St Lucia Formation,

Campanian II-?1II. Both x 1. ,

151

CRETACEOUS FAUNAS FROM SOUTH AFRICA

OO! “SI

ANNALS OF THE SOUTH AFRICAN MUSEUM

lay

153

CRETACEOUS FAUNAS FROM SOUTH AFRICA

Fig. 102

154 ANNALS OF THE SOUTH AFRICAN MUSEUM

MB D1320b. One of the largest. specimens

ation, Campanian i-7l, a

Venzo, 1936. N

0, Zululand, St Lucia Form

Fig. 103. Baculites vanhoepeni

of this species from locality 11

CRETACEOUS FAUNAS FROM SOUTH AFRICA 155

Large, mammiform nodes already occur in Lower Campanian B. sparsi-

nodosus (Fig. 60) and are well developed in B. mamillatus (Fig. 84A-C) in the

Middle Campanian. Thin, rib-like ornament, albeit very closely spaced, occurs

in B. coagmentatus (Fig. 88) and B. leopoliensis Collignon non Nowak

(= Baculites collignoni nom. nov.) (Fig. 86) in the Middle Campanian of

Madagascar.

We initially thought (Klinger & Kennedy 1977: 74) that B. tanakae

Masumoto & Obata (1963: 51, pl. 13 (fig. 4), pl. 16 Gigs 1-5), pl. 17

(figs 1-5), pl. 18 (figs 1, 3-4), pl. 19 (figs 1, 4), text-figs 97-113, 115) was a

synonym of B. vanhoepeni, but now rather regard it as being closer to

B. capensis than to B. vanhoepeni as discussed above (p. 109), and probably a

senior synonym of B. menabensis Collignon.

0) 5 mm

(oe ee

Fig. 104. Baculites vanhoepeni Venzo, 1936.

Suture lines. A-B. SAM-PCZ7706.

Scale bar for size.

Baculites taylorensis Adkins (1929: 204, pl. 5 (figs 9-11)), recently revised

by Kennedy & Cobban (1993a: 93, figs 10.1-10.9, 10.11-10.12, 10.16,

10.18-10.19; 11.1-11.2; 1993d: 143, pl. 6 (figs 1-9), pl. 7 (figs 1-6, 10-13),

text-fig. 8b, d), from the Middle Campanian Taylor Formation of Texas and the

Annona Chalk in Arkansas, is indistinguishable from B. vanhoepeni on the basis

of the original descriptions and figures of Adkins. The specimens figured by

156 ANNALS OF THE SOUTH AFRICAN MUSEUM

Kennedy & Cobban, however, all seem to be consistently more weakly

ornamented than B. vanhoepeni and more like B. sparsinodosus.

Matsumoto & Obata (1963: 55) also noted the parallelism between the

lineages of B. capensis—B. tanakae in the Indo-Pacific region, and B. codyensis

(as B. asper)-B. taylorensis in the Gulf—Atlantic region. In both lineages, which

cover more or less the same time interval, there is a progressive increase in

size, and coarsening and wider spacing of ornament.

O 10 mm

Fig. 105. Baculites vanhoepeni Venzo, 1936. Suture lines.

A-B. NMBD1302. C. SAM-PCZ7640.

Scale bar for size.

Baculites sp. (in Stephenson 1941: 407, pl. 76 (figs 7-8)) has similar lateral

ornament, but the dorsum seems more flattened—as in B. nibelae sp. nov.

According to Matsumoto (1959: 126), the former may be a representative of

B. lomaensis Anderson (1958: 191, pl. 48 (figs 5-6)). The latter, however,

apparently occurs in the Lower Maastrichtian of California.

CRETACEOUS FAUNAS FROM SOUTH AFRICA [57

ee,

0 5 mm

Fig. 106. Baculites vanhoepeni Venzo, 1936. Suture lines.

A-B. NMBD1302. C. SAS Z1191.

Scale bar for size of B and C.

A similar convergent lineage occurs in the Middle Campanian of the U.S.

Western Interior, consisting of B. obtusus-B. maclearni-B. asperiformis. The

latter part of this lineage, B. asperiformis, is remarkably similar to

B. vanhoepeni; compare e.g. B. asperiformis in Cobban (1962, pl. 106

(figs 10-11)) and Figure 91. The lateral ornament is identical, as is the

occurrence of ventral ribbing. The sutures are also similar, although some

specimens of B. vanhoepeni may have a more complex, dendritic suture.

However, the early part of this lineage, B. obtusus, has closely spaced

158 ANNALS OF THE SOUTH AFRICAN MUSEUM

dorsolateral crescentic ribs and a strongly corrugated venter—quite unlike any

of the Indo-Pacific material. Baculites vanhoepeni can perhaps be considered

analogous to B. obtusus, but there are sufficient differences to separate them.

0) 5 mm

ee eee

rad

Fig. 107. Baculites vanhoepeni Venzo, 1936. Suwtunewinnes

A-D. SAS Z1923. Specimen with complex suture line and constricted

lateral lobe (L). See also Fig. 108.

Scale bar for size.

As far as the ornament is concerned, small specimens of B. vanhoepeni

show a remarkable similarity to Boehmoceras arculus from the U.S. Gulf Coast

described by Kennedy & Cobban (1991b), already referred to above (p. 121).

Apart from the lack of the distinct curvature, specimens of B. vanhoepeni, e.g.

Figure 91A-O are indistinguishable from B. arculus in Kennedy & Cobban

(e.g. 1991b: fig. 9.11-52). Some specimens of B. vanhoepeni, e.g. PCZ8764

(Fig. 91A-C), do show slight curvature, but nowhere as pronounced as in

CRETACEOUS FAUNAS FROM SOUTH AFRICA 159

0) 5 mm

Oo 10mm

O 10 mm

Fig. 108. Baculites vanhoepeni Venzo, 1936. Suture lines.

A. SAS Z1923. Extremely complex, dendritic suture compared to other

specimens. B. SAS A2029. C. SAS A2034.

Scale bar for size.

Boehmoceras arculus. Apart from the curvature, B. arculus is an older species,

occurring in the Upper Santonian, whereas B. vanhoepeni is definitely younger,

occurring in the Middle to Upper Campanian of Zululand.

It thus appears that three, apparently unrelated baculitid lineages in the Indo-

Pacific, the Gulf Atlantic and the U.S. Western Interior, respectively, gave rise

in the Middle Campanian to remarkably similar forms with prominent, widely

spaced arcuate bullae, viz. B. vanhoepeni, B. taylorensis and B. asperiformis.

Another lineage gave rise to similar ornament in Boehmoceras in the Upper

Santonian. This is a remarkable example of both isochronous and hetero-

chronous homoeomorphy. It also illustrates the difficulties in trying to identify

baculites on isolated specimens on the basis of ornament alone. A fuller

discussion on these aspects of baculitid evolution is covered in Klinger &

Kennedy (in press).

160 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 109. Baculites vanhoepeni Venzo, 1936. A-C. SAS Z1868. D-E. SAM-PCZ7705.

H. SAM-PCZ7714, all from locality 110, Zululand, St Lucia Formation, Campanian II-III.

F-G. From the collections of the Geology Department, University of Natal, from locality 4,

Durban, Mzamba Formation, Campanian II. All x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 161

Fig. 110. Baculites vanhoepeni Venzo, 1936. A-I. Three unregistered specimens in the

collections of the Geology Department, University of Natal, from locality 4, Durban,

Mzamba Formation, Campanian IJ. All x 1.

162 ANNALS OF THE SOUTH AFRICAN MUSEUM

Kennedy (1986b: 110, pl. 17 (figs 7-9, 13-15, 21-23), pl. 18 (figs 18-22),

pl. 23 (figs 1, 7), text-fig. 8A, C) informally described a baculitid from the

Upper Campanian of France as Baculites sp. 1. Baculites vanhoepeni was

regarded as being closest to this informally named baculitid. In addition,

Holzapfel’s (1888: 64, pl. 4 (figs 5-6)) and Miller & Wollemann’s (1906: 4,

pl. 2 (figs 2-5)) Baculites incurvatus were regarded as belonging to this species.

As discussed above, B. incurvatus is a Coniacian—-Santonian species, and the

German records from the Lower Campanian are obvious misidentifications.

They seem closer to the Madagascan B. menabensis, which in turn is probably a

synonym of B. tanakae.

Some of the larger French specimens figured by Kennedy (e.g. 1986b,

pl. 18 (figs 18-22)) have ornament comparable to that of B. vanhoepeni, but

more material is necessary for definite comparisons.

Occurrence

Baculites vanhoepeni typically occurs with a Middle Campanian texanitid

fauna of Australiella and Delawarella, but we suspect that it ranges into the

Upper Campanian by its association with Hoplitoplacenticeras howarthi, both at

locality 110 in Zululand (Klinger & Kennedy 1989), and at Somtseu, Durban

(Kennedy & Klinger 1973).

Baculites nibelae sp. nov.

Figs 116-120

? 1970 = Baculites cf. taylorensis Adkins; Collignon, p. 13, pl. 612 (fig. 2285).

a) 1970 Baculites sp. (nov.?) cf. aquilaensis Reeside; Collignon, p. 81, pl. 639

(fig. 2358).

Type

Holotype is SAM-PCZ9148b from locality 111, cliff section just east of the

Nibela Peninsula, Zululand, St Lucia Formation, Campanian III.

Material

Paratypes are SAM-PCZ7570, 7578, 7625, 7629, 7644, 7698a-e, 7702,

7850, 9155, 9163 and 9166, all from locality 109C; SAM-PCZ7569 from

locality 1O9A, SAM-PCZ763la, 7699, 7630, 7632, 7700 and 7701, from

locality 109D, all on the western shores of the Nibela Peninsula, Zululand, St

Lucia Formation, Campanian III; and SAM-PCZ9146, 9147, 9148a-e,

9149a-f, 9150a-g and 9151-9154, all from locality 111, St Lucia Formation,

Campanian III.

Fig. 111 (see facing page). Baculites vanhoepeni Venzo, 1936. Two unregistered specimens

in the collections of the Geology Department, University of Natal, from locality 4, Durban,

Mzamba Formation, Campanian II. Note similarity of A-C to Baculites collignoni nom.

nov. (sce Fig. 96). Allx i

Fig. 112 (see overleaf). Baculites vanhoepeni Venzo, 1936. Three unregistered specimens in

the collections of the Geology Department, University of Natal, from locality 4, Durban,

Mzamba Formation, Campanian II. Note similarity to Baculites mamillatus

(see Fig. 84A-C). All x 1. ;

CRETACEOUS FAUNAS FROM SOUTH AFRICA

163

Figs 111

164 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 112

CRETACEOUS FAUNAS FROM SOUTH AFRICA 165

10 Size distribution in Baculites vanhoepeni

Ojindicates distribution of specimens with preserved

apertures.

NUMBER OF SPECIMENS

20 30 40 50

MAXIMUM WHORL HEIGHT IN MM

Fig. 113. Histogram illustrating maximum size distribution in Baculites

vanhoepeni Venzo, 1936. Open circles indicate specimens with the aperture

preserved.

Dimensions

Specimen MxWb MxWh Wb/Wh MnWb MnWh Wb/Wh D_ Ti Ri

Porvs5'6 26.0 34.0 0.76 — — ~—

PCZ9148b 22.0 29.0 0.76 1La0: 26.0) 070 96y 3.1 2,

PCZ7698a 21.0 29.0 0:72 — — — —_- — —

PoZ1699 = =«©21.0 29.0 0.72

PCZ9148a_ 20.0 27.0 0.74 1720 220 OT COm Wet ~

meets) 20.0 30.0 0.67 19.0> 228.0) (0:68 63 7.1 214

PCZ9149b 18.0 23.0 0.78 16:0) 23.0" 10:7 52. — Pap

meee 16.0 25.00 0.72 — — — —- — —

eegese 17.5 24.0. 0.73 — — — —- — —

PCZ9149a — 5.0 — — 12.0 — 540550 22

meevisse 10.0 14.0 0.71 9.0 135.0); 0-61 60 1.7 pM ps

Diagnosis

Baculitid with subtrigonal whorl section and arcuate lateral ornament.

Description

The maximum observed whorl height is 34 mm. In typical forms the whorl

section is subtrigonal with a distinctly flattened dorsum and flanks converging to

a rounded venter (Fig. 118B-G). In juvenile specimens the whorl section is

distinctly ovoid to tear-shaped with the venter much narrower than the dorsum.

166 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 114. Baculites vanhoepeni Venzo, 1936. ?Early forms. A-B. SAM-PCZ9255.

C. SAM-PCZ9252. Both from locality 109A, Zululand, St Lucia Formation, Campanian II.

Fig. 115 (see facing page). Baculites vanhoepeni Venzo, 1936. A-C. Typical form,

SAM-PCZ9284 from locality 110. D-F. SAM-PCZ9131, ?early form, from locality 109A.

Both x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 167

168 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 116. Baculites nibelae sp. nov. The holotype, SAM-PCZ9148, from locality Pos

St Lucia Formation, Campanian III. x 1.

Fig. 117 (see facing page). A-D, F-I. Baculites nibelae sp. nov. A. SAM-PCZ9148a.

B. SAM-PCZ9149a. C-D. SAM-PCZ9150e. Specimen with part of aperture preserved.

F-H. SAM-PCZ9153. I. SAM-PCZ7631. All from locality 111, St Lucia Formation,

Campanian III. E. Baculites duharti Hinicken, 1975. SAM-PCZ9873c, from concretion at

base of cliff at locality 110, Zululand, St Lucia Formation, Campanian II.

Allee.

169

CRETACEOUS FAUNAS FROM SOUTH AFRICA

wee

Fig. 117

170 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 118. Baculites nibelae sp. nov. Partial suture line and whorl sections.

A. SAM-PCZ7630. B. SAM-PCZ7631. C. SAM-PCZ7701.

D. SAM-PCZ7644. E. SAM-PCZ7/630. EF. SAM-=PCZ7Goae

G. SAM-PCZ7698. Venter in whorl section facing upward.

Scale bar for size.

Fig. 119 (see facing page). Baculites nibelae sp. nov. A. SAM-PCZ7702. B-D. SAM-

PCZ736la. E-F, I. SAM-PCZ7578. G. SAM-PCZ7698. H. SAM-PCZ9153. All from

locality 109, Zululand, St Lucia Formation, Campanian III. All x 1.

A. SAM-PCZ7630. B-D. SAM-

Fig. 120 (see overleaf). Baculites nibelae sp. nov.

All from

PCZ12021. E. SAM-PCZ9149. F. SAM-PCZ7698. G. SAM-PCZ9148e.

locality 109, Zululand, St Lucia Formation, Campanian III. All x 1.-

171

CRETACEOUS FAUNAS FROM SOUTH AFRICA

Fig. 119

LZ ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 120

CRETACEOUS FAUNAS FROM SOUTH AFRICA 173

Lateral ornament consists of crescentic ribs of variable strength,

which cover the dorsal two thirds of the flanks—occurring at a frequency of

2-3 per whorl height. In some, e.g. PCZ7630 (Fig. 120A) and PCZ7701,

the flanks are virtually smooth; in others, e.g. PCZ7702 (Fig. 119A), the ribs

are quite prominent. In the latter, the costal whorl section is conspicuously

trigonal to cuneate. In some specimens the venter appears to be slightly

undulating.

The aperture is preserved in PCZ9150e (Fig. 117C-D). The dorsal rostrum

is short, and the ventral rostrum only a little longer, and distinctly spoon-

shaped.

The suture is only partially exposed in a few specimens, e.g. PCZ9150h and

PCZ7630 (Fig. 118A). It shows narrow lateral saddles and a splayed lateral

lobe (L).

Fig. 121. A-C. Baculites sp. (?nov.) cf. B. aquilaensis Reeside. Plaster cast of GD12358,

the original of Collignon (1970, pl. 639 (fig. 2358)), from the Upper Campanian, zone of

Hoplitoplacenticeras marroti of gisement 227-2, Mokotibe (Antsalova), Madagascar.

D-E. Baculites bassei Besairie, 1930. Copy of the lectotype illustrated in Besairie (1930,

pl. 22 (fig. 8-8a)), from the Upper Campanian or Lower Maastrichtian, of north of

Trangahy, Madagascar. Both x 1.

Discussion

The subtrigonal whorl section clearly distinguishes B. nibelae from

B. vanhoepeni, the latter having a typically elliptical whorl section. The shape

of the lateral ribs in both species is similar but those of B. vanhoepeni are much

more robust, and the latter species grows to a much larger size than B. nibelae.

The occurrence of B. nibelae at locality 111, east of 110, suggests that it is

younger than B. vanhoepeni, but still in the Campanian.

174 ANNALS OF THE SOUTH AFRICAN MUSEUM

The subtrigonal whorl section of B. nibelae places this species close to the

group of baculitids described by Collignon (1970) from the lower part of the

Middle Campanian of Madagascar. The closest match is probably B. rectangu-

latus Collignon (1970: 12, pl. 611 (figs 2279-2281)) (herein Fig. 87), but the

whorl section of B. nibelae is more prominently triangular and, in typical

forms, it has stronger lateral ornament.

The specimen described by Collignon (1970: 13, pl. 612 (fig. 2285)) from

the upper part of the Middle Campanian of Madagascar as Baculites cf.

taylorensis is probably the same as B. nibelae. Baculites sp. (nov.?) cf.

B. aquilaensis in Collignon (1970: 81, pl. 639 (fig. 2358)) (Fig. 121A—-C) from

the Upper Campanian of Madagascar has stronger ornament than any of our

B. nibelae, but the whorl section suggests that it is closely allied. Baculites

bassei Besairie (1930: 222, pl. 22 (fig. 8, 8a)) (herein Fig. 121D-E), from the

Upper Campanian or Lower Maastrichtian of north of Trangahy (Maintirano) in

Madagascar, is known from a single specimen only. It is also very similar to

B. nibelae. Given more and precisely located material from Madagascar,

B. bassei may prove to be a senior synonym of B. nibelae.

The Campanian baculitid from Andimaka described by Collignon (1938) as

B. aspero-anceps (Collignon 1938: 89, pl. 6 (fig. 7)), also has a trigonal whorl

section similar to that of B. nibelae, and may belong to this species.

In terms of lateral ornament and to a lesser extent, the whorl section,

B. nibelae is similar to B. subanceps Haughton (Figs 130, 131A—H) (see also

Howarth 1965: 368, pl. 5 (fig. 3), pl. 6 (figs 6-7), pl. 7 (fig. 1), text-figs 4,

13-15). In the latter species, however, the venter tends to become flat and, in

some specimens, even forms an incipient tabulate ventral keel as in typical

Eubaculites.

The whorl section and ornament of B. nibelae are very similar to those of

Eubaculites labyrinthicus (Morton), recorded from the Lower Maastrichtian of

Madagascar by Collignon (1971: 18, pl. 646 (fig. 2395)) (as Eubaculites

otacodensis Stol.). As yet undescribed baculitids from the Upper Campanian of

Israel, with even more angular trigonal whorl section and prominent lateral

ribbing, are even closer to E. labyrinthicus. Baculites lomaensis Anderson

(1958: 191, pl. 48 (figs 5, 5a-6)) (see also Matsumoto 1959: 126, pl. 34

(figs la-c, 2a-c), text-figs 35-38, 39-41), from the Lower Maastrichtian of

California, has a similar trigonal whorl section, but apparently weaker

dorsolateral bullae. We doubt if B. nibelae, B. lomaensis and the Israeli

baculitids are ancestral to E. labyrinthicus, but the style of ornament and whorl

section are remarkably similar.

The evolution of baculitids with a*trigonal whorl section, as in E. laby-

rinthicus, appears to be a wide-spread feature near the Campanian-

Maastrichtian boundary, as already noticed by Lewy (1986: 5) (see also Klinger

& Kennedy in press).

Fig. 122 (see facing page). Baculites duharti Hiinicken, 1975. A-D. SAM-7683 from

locality 109H, Zululand, St Lucia Formation, Campanian II. E-G. SAM-PCZ9873a from

locality 110, Zululand, St Lucia Formation, Campanian II.

A, E-G x 1; B-D x 0.75.

CRETACEOUS FAUNAS FROM SOUTH AFRICA Ys)

176 ANNALS OF THE SOUTH AFRICAN MUSEUM

Cc D E

Fig. 123. Baculites duharti Hiinicken, 1975. A-B. SAM-PCZ7703a. C-F. SAM-

PCZ7685, both from locality 109G, Zululand, St Lucia Formation, Campanian II. Both x 1.

Fig. 124 (see facing page). Baculites duharti Hinicken, 1975. Concretion, SAM-

PCZ9873, from base of cliff at locality 110, Zululand, St Lucia Formation, Campanian II.

All x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA

178 ANNALS OF THE SOUTH AFRICAN MUSEUM

Occurrence

Campanian III of Zululand.

Baculites duharti Hinicken, 1975

Figs 122-127

1975 Baculites duharti Hunicken, in Hinicken et al. 1975, p. 116, pl. 1 (figs 1-4),

pl. 2 (figs 1-2), pl. 3 (figs 5-8), text-figs 2a-d, 3a-c, 4-5.

1980 Baculites duharti Hunicken; Hiinicken, Charrier & Lahsen, p. 224, pl. 1

(figs la—b, 2), pl. 2 (figs la-c, 2, 3a-b, 4-5, 6a-b), text-figs 3-9.

Type

Holotype is the specimen figured by Hunicken (in Hunicken et al. 1975,

pl. 1 (figs 1-4), text-figs 2c-d, 4a) from the Middle and/or Upper Campanian of

Member ‘e’ of the Cerro Matero Formation, at Rio Sur, Tierra del Fuego,

Chile, CPC D1127.

Material

More than 100 specimens from a single concretion, SAM-PCZ9873,

9873a-v, found at the base of the section at locality 110, Nibela Peninsula,

Zululand, St Lucia Formation, Campanian III; SAM-PCZ7683, 7685, and

7703, from locality 109G; and SAM-PCZ7689 from locality 109H, Nibela

Peninsula, Zululand (Fig. 90), St Lucia Formation, Campanian II.

Dimensions

Spec. MxWb MxWh Wb/Wh MnWb MnWh Wb/Wh D_ Ti

PCZ987 3a) 12209 1720) 0 8.67 | 12-07 0572 48 10.4

PCZ7689 22.0 31/0") 2071 20s 280 a Ors 64 4.69

PCZ7685.. 22:0' 29:0 O56) 9 2030) eo Ue Oy) 57 26

PCZ7683 «34.0 43.0 079 23.5. 38.0 0:76.) 200siane

PCZ7703:. 135.0") 58:0 4) 70:60 — = ae

Description

A large concretion from the base of the section at locality 110 is crammed

with more than 100 specimens of a smooth baculitid (Figs 124-126), differing

markedly from the heavily ornamented B. vanhoepeni from the higher parts of

the section. Four other smooth baculitids from the south-western part of the

Nibela Peninsula at localities 109G and 109H (Figs 122A-D, 123A-F) are much

larger than those from locality 110, but we assume they belong to the same

species—being of more or less the same age.

Fig. 125 (see faing page). Baculites duharti Hiinicken, 1975. Concretion, SAM-PCZ9873,

from the base of cliff at locality 110, Zululand, St ‘Lucia Formation, Campanian iL.

Note slight wavy venter on top right. x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 179

Fig. 125

180 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 126. Baculites duharti Hiinicken, 1975. Concretion, SAM-PCZ9873, from base of

cliff at locality 110, Zululand, St Lucia Formation, Campanian I]. -

Note wavy venter at top left. x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 181

The whorl section (Figs 123B, F, 127) is distinctly elliptical, to ovoid, with

the venter only very little narrower than the dorsum. Lateral ornament consists

of very fine striae only, but in some specimens, e.g. PCZ9873a (Fig. 122E-G),

low, fold-like ribs develop over the ventral half of the flanks. In some of the

larger specimens, where the shell is preserved (Figs 125, 126), the venter is

ornamented with scale-like undulations.

A slight constriction is visible on the venter of SAM-PCZ7685

(Fig. 123C-F). The species grows to a large size—the maximum whorl height

measured is 60 mm (Fig. 123A-B). The suture is finely incised with open

saddles and lobes (Fig. 127A-C).

E iE

O 10 mm

Yop Po .

E U C

| a

D 0) 5mm G a

Fig. 127. Baculites duharti Hiinicken, 1975. Suture lines and whorl sections.

Venter facing upwards. Scale bar for size.

182 ANNALS OF THE SOUTH AFRICAN MUSEUM

a gs :

Fig. 128. Baculites ambatryensis Collignon, 1971. Plaster cast of the holotype, GD12392,

the original of Collignon (1971, pl. 645 (fig. 2392)) from the Lower Maastrichtian of

gisement 400, Cote d’Ambatry (Betioky), Madagascar. All x 1.

Discussion

The elliptical whorl section and relatively simple suture show that this

baculitid belongs to the group of B. vanhoepeni. However, the total absence of

lateral ornament in more than a hundred specimens clearly shows that this is

indeed a consistently smooth baculitid species and not merely a smooth variant

of B. vanhoepeni.

As discussed above, identifying or separating smooth Baculites species is

very difficult unless the whorl section and sutures are very distinctive, or if the

exact age is known.

The closest match we could find both in morphology and age is B. duharti

first described from Tierra del Fuego by Hiinicken (in Hiinicken et al. 1975:

116, pl. 1 (figs 1-4), pl. 2 (figs 1-2), pl. 3 (figs 5-8), text-figs 2a—d, 3a—c), 4-5)

and imprecisely dated as Campanian—Maastrichtian. Later, Hunicken ef al.

(1980: 224, pl. 1 (figs la-b, 2), pl. 2 (figs la—-c, 2, 3a—b, 4-5, 6a-b),

Fig. 129 (see facing page). Baculites asperoanceps Lasswitz, 1904. Plaster cast of the

lectotype, nr 3045 s (k), MGUWr, from Austin, Texas, in the collections of the Henryk

Teisseyre Geological Museum of the Institute of Geological Sciences of the University of

Wroclaw. x l.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 183

184 ANNALS OF THE SOUTH AFRICAN MUSEUM

text-figs 3-9) described additional material, also from Tierra del Fuego, but

more precisely dated as Middle to Late Campanian. The drawings of the suture

lines by Hunicken ef al. (1980, text-figs 3-9) show a suture more complex and

phylloid than ours, but photographs of the sutures (e.g. their pl. 2 (figs 3-6)) are

much simpler and comparable to the Zululand material. From a palaeobio-

geographical point of view, the record of B. duharti from Zululand also is

acceptable.

Baculites duharti differs from the older B. bailyi mainly in being much

larger and having a predominantly elliptical, rather than ovoid, whorl section.

We do not think that B. duharti is derived from the B. bailyi lineage; instead we

regard it as a smooth offshoot of the B. capensis—B. vanhoepeni lineage.

Baculites duharti clearly differs from B. rectus in having a much simpler

suture.

The only Middle Campanian Madagascan baculitid species with which

B. duharti could possibly be compared, is B. ankilizatensis (Collignon 1970: 13,

pl. 612 (figs 2282-2284)) (herein Fig. 89), especially as far as maximum size is

concerned. However, some specimens of B. ankilizatensis show distinct, albeit

low, lateral ornament.

Some specimens of B. subanceps in the collections of the South African

Museum, e.g. SAM-6829a (Figs 130D-F) are indistinguishable from

B. duharti, but this species includes forms with distinct lateral ornament, and

the whorl section is quite distinctive in some specimens, tending to form a

tabulate ventral keel.

Our material also resembles Baculites smooth species of Cobban (1962:

714, pl. 108 (figs 1-4), text-figs li-j) from the Middle Campanian of eastern

Wyoming. This species was renamed B. cobbani by Khakimov (in Atabekian &

Khakimov 1976: 98, pl. 11 (figs 2-7)) for material from the Lower Campanian

of Central Asia.

The largest of our specimens (Fig. 123A-B) is superficially similar to

B. knorrianus Desmarest, 1817 (recently reviewed by Kennedy &

Summesberger 1987: 32, pl. 4 (figs 4-6), pl. 5 (figs 1-14), text-fig. 2;

Birkelund 1993: 52, pl. 13 Ciigs 12-14) and’ Kennedy 1993-3109 3sgiees

(figs 13-22), pl. 6 (figs 11-13, 18-23), text-fig. 5a-c). However, B. knorrianus

is restricted to the Lower Maastrichtian, and has a far more compressed whorl

section and a more complex suture line.

Fig. 130 (see facing page). Baculites subanceps Haughton, 1925. A-C. SAM-6824, the

lectotype. D-F. SAM-6829a. G. SAM-6829b. All from the Upper Campanian-Lower

Maastrichtian of Carimba, Angola.

Fig. 131 (see overleaf). A-H. Baculites subanceps Haughton, 1925. A-C. SAM-6829c.

D-G. SAM-6829d. H. SAM-6829e. All from the Upper Campanian—Lower Maastrichtian

of Carimba, Angola.

I-M. Baculites anceps Lamarck, 1822. I-K. SAM-6145b. L. SAM-6145c.

M. SAM-6748, from the ?7Lower Maastrichtian east of Capolo. N-R. Baculites sp. from the

uppermost Turonian or basal Coniacian at Mossamedes, Angola. The oldest tuberculate

baculite. Unregistered specimens from Cooper collection, SAM.

A-C, G, I-R x 1; D-F ~x 0.5.

CRETACEOUS FAUNAS FROM SOUTH AFRICA

185

"htecnnianatatbtitetne

Fig. 130

186 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 131

CRETACEOUS FAUNAS FROM SOUTH AFRICA 187

Baculites ambatryensis Collignon (1971: 14, pl. 645 (fig. 2392)) (herein

Fig. 128) from the Lower Maastrichtian of Madagascar is also superficially

similar, but has a more compressed whorl section at comparable diameters

(Wb: Wh 19.5: 29), and coarse corrugations over the venter.

Baculites eliasi Cobban (1958: 663, pl. 91 (figs 1-11), text-figs 1f-g, i-j)

from the Lower Maastrichtian of the U.S. Western Interior also has a more

complex suture with a constricted lateral lobe (L).

A B C D

Fig. 132. Baculites sp. from the uppermost Turonian or basal Coniacian at Mossamedes,

Angola. The oldest tuberculate baculite. Cooper Collection, S.A. Museum.

A. SAM-1742. B-D. SAM-1743. E. SAM-1733. F. SAM-1745.

Allo< I:

Baculites fuchsi Redtenbacher (1873: 134, pl. 30 (fig. 15)) is difficult to

interpret (see above, p. 47) but it is older, Santonian, being associated with

B. incurvatus.

Baculites inornatus Meek (1862: 316) (see especially Matsumoto 1959: 155,

pl. 38 (fig. la-c), pl. 43 (fig. 5a—c), text-figs 72a—b, 73a—d, 74-79; Ward 1978:

1151, pl. 1 (figs 1-2), text-fig. 5) is a predominantly smooth species from the

Lower Campanian of California, British Columbia and Hokkaido. Records of

B. inornatus from the Coniacian of Venezuela by Renz (1982: 105, pl. 34

(figs 3-4, 5a—b, 6), text-fig. 80) are obvious misidentifications. The suture of

B. inornatus is complex, with phylloid folioles, clearly differing from our

material.

Baculites kotanii Matsumoto ef al. (1980: 408, figs 1-2) from the Upper

Campanian of Shikoku, Japan, is nearly smooth, with only faint subcostae on

the flanks. Again, the suture appears to be more complex with narrower saddles

and lobes than the Zululand material.

Baculites vertebralis Lamarck, 1801, is a compressed, smooth species from

the Upper Maastrichtian of Europe and North Africa, and resembles our largest

specimen. Age difference apart, it also has a far more complex suture line (see

e.g. Kennedy 1986a, text-fig. 7D-E).

188 ANNALS OF THE SOUTH AFRICAN MUSEUM

Occurrence

Middle and/or Upper Campanian of Tierra del Fuego, and Campanian III of

Zululand.

ACKNOWLEDGEMENTS

Financial assistance to H. C. Klinger by the Foundation for Research

Development (S.A.) and to W. J. Kennedy by the Natural Environment

Research Council, U.K.) is gratefully acknowledged. We also thank Sally

Dove, Jacque Blaeske and Samantha Black for their technical and artistic

assistance.

For access to, or information about collections in their institutions, we are

indebted to numerous colleagues: Dr W. A. Cobban (Denver), Prof. M. R.

Cooper (University of Durban-Westville), Dr N. Eldridge (Washington),

Dr J. Gorzyca-Skala (Wroclaw), Dr M. K. Howarth (London), Prof. H. Immel

and Dr G. Schairer (Munich), Dr K. Larsson (Lund), Dr U. Leppig (Freiburg),

Dr R. Ragaini (Pisa), Prof. H. Remy (Bonn), Prof. S. Ritzkowski (G6ttingen),

Dr Carlo Sarti (Bologna), Dr F. Thackeray (Pretoria) and Dr J. Thierry

(Dijon).

We thank the Natal Parks Board, especially Mr R. Taylor, for access to

areas under their control and for assistance with transportation around the Lake

St Lucia area during the field season of 1992.

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198

APPENDIX

List of material and dimensions of Baculites bailyi, Baculites capensis, and Baculites vanhoepeni.

Baculites bailyi

ANNALS OF THE SOUTH AFRICAN MUSEUM

MnWb = MnWwh_ Wb/Wh Ti

MxWb MxWh Wb/Wh

Specimen

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Baculites bailyi (cont.)

MnWb = MnWh_ Wb/Wh

Wb/Wh

MxWb MxWh

Specimen

CRETACEOUS FAUNAS FROM SOUTH AFRICA

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Tu/Wb

MnWb = MnWh_ Wb/Wh

MxWb MxWh ~~ Wb/Wh

Specimen

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199

200

Baculites capensis (cont.)

Tu/Wb

MnWb MnWh Wb/Wh

MxWb MxWh Wb/Wh

Specimen

ANNALS OF THE SOUTH AFRICAN MUSEUM

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si va) nv Inn mn nN WH Ta)

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ao @)sopao app co sean (M@yan ac jaa) Sy ay ahacpan, an (atjac en ee) oc) 1

201

CRETACEOUS FAUNAS FROM SOUTH AFRICA

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(é CY €7 99'0 O'€] fee: IL'0 O'+rl 0°01 ET8Pb-INVS

Cre ob 6£ LL'‘0 O'€l 0°01 890 8b] 0°01 ®//7OCTH

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Ca tee 87 Lew 0°71 76 pL'0 O'€] 9°6 C/PT11G

QM /"L 1 d UM/IM YUMUN IMU YUM/IM = UMXW IM XW uauioad¢

(WU0d) SIsuadDd SajljnIvg

202

Baculites capensis (cont.)

Tu/Wb

MnWb MnWh Wb/Wh

MxWb MxWh Wb/Wh

Specimen

ANNALS OF THE SOUTH AFRICAN MUSEUM

nm nm QO

| Piao ye | te

MANN MOAN NMANAANN NANM NN —

WOON \OmMm a LV ae al @N EL ) O Or

NMAN LV a Oa YN aN gi orn oO WY wT Oo Oa aM

S22 | AeotesS WM FN WN — 0O WH FN D © &'6aico

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On CO COmmnWY ve) ioe)

Oxo |OERh | moenoer | Rtn i meee

eo oQOoQoqoqg GoeQoqaoo eoqQoqo moos]

NOS), Soot MOONNS ,CCSSO ,OMSCO

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TFHNNNNHNNNHE HOOK OHOOOSKEONEEH

eee eee eee eee ee ees es ae

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BANANA AAA ANAAANAAAA

ete eee es ss es es a

GUS) Two =) ON C00

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SONAR amo esasertasesseags ANnNAXO

SSOGFNMMATOSSCOSOSSSCSCOONGDO=NS

ANS HOUTMMOMAANANAANAANAA Ss ]OO082 S83 zazN0

LhOAMnmeoNnOAS Lae aL Leata Aaaiaise

Baculites capensis (cont.)

Tu/Wb

MnWb = MnWh_ Wb/Wh

MxWb MxWh ~ Wb/Wh

Specimen

CRETACEOUS FAUNAS FROM SOUTH AFRICA

el lt me eanih lee

cn a a ANAAAANN ANNANA TNA

on Netsgle) ven We OlAtS man

Meco | [eee ee ee es Pe

Gon aii: g ANOF MEOH AMMAN FNM

fa '2 | | |] | wmeoceroay | | Qnetre tot

— ol 4 MNO SAE + AMOM MMO

ation THAMMA HAN =r aMr ODN

ft | fe) ft eeeorr eo | [eee eee

Ss © Slololelelelere) Seqoqcoqgneoo]oe

Se oO Oooocoon SVSleoere) a7)

Iw lw l ll leer |! lkKowrKr

— — eee es ss eee ee es st

Sr > Soma oco Ss oooCcoooco

— — ON Oe ee es se Seni aren fi seen ith seen fi cen i cen ih eee I

DBDOODONADANA BOATS TOTTO FTO TNH UM

ee Oe er er rice ee

SS) Ss sleleleloleleljoloelo) ejalole\oalelolajealelerel=)

WOOoocouoooeoonenoooocoooonNnooe

WU 00 00 ~ 00 \0 00 00. OF COM F CD OONDOONONAACOCO

ee Ot ON OS OS as

MOmooDoooooooom Nnoeooo0oooconne

ANAAKNHHNHNHKHNKHKNKNMNOKntttt¢t¢¢ttttHN

ee es a et ed

PCP8052

H201/9

PCZ8004

H203/4

H201/12a

H205/18

H202/5

H201/26

H201/21

H200/14

Z.1795d

PCZ8027

203

204

ANNALS OF THE SOUTH AFRICAN MUSEUM

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ee ee =e ss as c8 0 0 CC 081 8£08ZO0d

= 8S I S6 1 O One OST LL O SG 081 78/00CH

Gs |! Cv 08 68 0 0 0¢ OLI i) Dte€ 081 SC8P-NVS

SC 99'S 06 GLY 0 81 One vL 0 OxEC OLI SOLIZ

= DS te L8 OL 0 9°61 Ih toll (ele, LOG COI 1/cSOTd

0 WIL LC vl, 0 61 OVI 6L 0 OI? Shh O¢/SOCH

0) 8 TI be vl O 061 OVI 69 0 8 CC LSI yolld

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dM/"L WD d YM/IM = YUMYN = UIMYW YM/4M = YUMXIN = IM XW uauioads

(‘JUOD) Sisuadvd salljnovg

Baculites vanhoepeni

Tu/Wb

MnWb MnWwh Wb/Wh

MxWb MxWh Wb/Wh

Specimen

CRETACEOUS FAUNAS FROM SOUTH AFRICA

Orne tHt tAKR+t+HONO a Ya) Nt Fm

W\O x~F 0O <+F NMN\OM Sie aiGn) on! a) coo 4 CO CO \O

AOD NH OAM CO CIN st rT +4 C~ t~ nS)

Set | (Ree |scrrm (fee ler | feoer

S S'o'\o Solo’ SoS SIS) S\S) Sle

fee eee eee Sas ha) Sa eas

—_— — ws oe eo —_—_ ee — —

AMNONO =O NAG RHANHNTM ONHAN NOx x

WR RRR RRR OR KEE EE ORK EEE OF OF Ee

2 2S elersojelelele) ole) slololerealevele) ololeyeo:

OMMOOTONCOMmMOOCCCONNGTOOCCCCoCo

ONnNNn~rKo- —=ANnO— attMNtonrrwdoonnonan

ss st GN GN GNIIQN GIUGAIGCI GG GiGi Qi QI QIGAGaicn Gaicn

RRS t ee

\O t~ [~ OW SI) (ex | Ne) Orn WY Sic DY

Sa ES SOOS KH SHED S8s4¢6

Bene CR OQSKKroSSrornteSeno

SRN oe SNS CHN NER SNEED SND N ES

lal. 0 t-te e—e a

205

206 ANNALS OF THE SOUTH AFRICAN MUSEUM

= eile sea) fa pees | ea

eS = =m ONNSO — —

co | C010 eA 7 REO 1

Bet |) > eoecill ate een aoc tet ee ediaes We Ald

Q C0 ]1ND | |MMar |oeM |S | | |

~ lum Mm~tnoo |! Mw ! oO

= \O Wma NeNM N © |

= S Ss SEeS SO Sc

Slo. SoOSOe Se 6

S AN oman AAMAM MMH O-

= S SOSoe (oo ©

Sl ve ee ele ee

S N AN AN AN A

oSoooseoeoesseessooosesco

AMMAAMN NM ANMNOONNDWM OO

GB GG) GO CG) Ga)! GG) Ga) GaGa)! Ga)! Ga) Ga) Ga) sir astpostn O@ ote

Soooooooeseseosoosoescsse

WWW \O \O [~~ C~ C~ F~ CO CO COCO SH HNN

NANANAANAANIAAAAANMAMMNAMNM

MxWb MxWh Wb/Wh

Specimen

Z1860

PCZ7635

A2034

A477

A2039

109b

D1302b

Z1860b

Z1190b

PCZ8630

PCZ7417

PCZ7406

PCZ8638

A2029b

PCZ7411

Z1860d

PCZ7407b

Baculites vanhoepeni (cont.)

Z1860

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. The name of the taxon should be followed, without intervening punctuation, by the author’s name

(not abbreviated) and the year of publication; a comma must separate author’s name and year. The

author’s name and date 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 either 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 (see example 1), or 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 (see example 2). The author should adopt one style or the

other throughout a paper.

Family Nuculanidae

Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-15A

Example 1

Nucula (Leda) bicuspidata Gould, 1845: 37.

Leda plicifera A. Adams, 1856: 50.

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

Nucula largillierti Philippi, 1861: 87.

Leda bicuspidata (Gould): 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.

Example 2

1845 Nucula (Leda) bicuspidata Gould, p. 37.

1856 Leda plicifera A. Adams, p. 50.

1859 Laeda bicuspidata (Gould) Hanley, p. 118, pl. 228 (fig. 73).

1861 Nucula largillierti Philippi, p. 87.

1871 Laeda bicuspidata (Gould): Sowerby, pl. 2 (fig. 8a-b).

1950 Leda bicuspidata (Gould): Nicklés, p. 163, fig. 301.

1955 Leda bicuspidata (Gould): Nicklés, p. 110.

1964 Leda bicuspidata (Gould): Barnard, p. 234, figs 8-9.

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

mentioned in the original description are to be designated allotype (if applicable) and/or 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 .. .’, or *. . . in C. namacolus (Fig. 10). . .’

(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded by initials or full

names: e.g. Du Toit, but A. L. du Toit: Von Huene, but F. von Huene

Punctuation should be loose, omitting all not strictly necessary. Reference to the author should

preferably 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, e.g. ‘Revision of the Crustacea. Part VIII.

Amphipoda.’. A specific name must not stand alone, but be preceded by the generic name or its

abbreviation to initial capital letter (except at the beginning of a sentence or paragraph), provided the

same generic name is used consecutively. The name of new genus or species should not 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.

8. GENERAL. Once referees’ reports have been received by the editor, these will be discussed by the

editorial committee. If the paper is considered acceptable after minor or major revision, the reports

will be forwarded to the author who must then thoroughly revise in accordance with the referees’

Suggestions. Final acceptance of the revised manuscript will be considered by the editorial committee.

In the case of major revision being necessary, the committee reserves the right to consult one or more

referees regarding the revised manuscript.

HERBERT CHRISTIAN KLINGER

WILLIAM JAMES KENNEDY

CRETACEOUS FAUNAS FROM ZULULAND AND >

NATAL, SOUTH AFRICA. THE AMMONITE

FAMILY BACULITIDAE GILL, 1871

(EXCLUDING THE GENUS EUBACULITES)

VOLUME 105 PART 2 FEBRUARY 1997

OF THE SOUTH AFRICAN

MUSEUM

CAPE TOWN