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

Annals and Magazine of Natural History (13) 2: 309-320.

KOHN, A. J. 1960b. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.

Bulletin of the Bingham Oceanographic Collection, Yale University 17 (4): 1-51.

THIELE, J. 1910. Mollusca. B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische

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ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 107 Band

January 2001 Januarie

Part ~ 1" Deel

STRATIGRAPHIC AND GEOGRAPHIC

DISTRIBUTION, PHYLOGENETIC TRENDS AND

GENERAL COMMENTS ON THE AMMONITE

FAMILY BACULITIDAE GILL, 1871

(WITH AN ANNOTATED LIST OF SPECIES

REFERRED TO THE FAMILY)

HERBERT CHRISTIAN KLINGER

WILLIAM JAMES KENNEDY

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STRATIGRAPHIC AND GEOGRAPHIC DISTRIBUTION,

PHYLOGENETIC TRENDS AND GENERAL COMMENTS ON

THE AMMONITE FAMILY BACULITIDAE GILL, 1871

(WITH AN ANNOTATED LIST OF SPECIES REFERRED

TO THE FAMILY)

HERBERT CHRISTIAN KLINGER

Division of Earth Sciences, South African Museum, Cape Town

WILLIAM JAMES KENNEDY

Geological Collections, Oxford University Museum of Natural History

(With 197 figures)

[MS accepted 30 January 1998)

ABSTRACT

Systematics in the ammonite family Baculitidae, especially in the genus Baculites, are

complicated by wide intraspecific variation and apparent strong endemism. Several aspects of

the stratigraphic and geographic distribution, phylogenetic trends in ornamentation, suture

lines, coiling, absolute size, dimorphism and whorl section are investigated to determine

whether morphological ‘spikes’ can be identified to improve the biostratigraphic potential of

the family. General comments on aptychi, ammonitellas, mass occurrences, muscle scars,

microsculpture, predation and parasitism are provided, as well as an annotated list of species

referred to the family. Some of these are figured photographically.

CONTENTS

PAGE

BALROG ICE OM weer er ee erence ee ioe titi o chia sie aisle Ss oar oe Siete era ioriercloles S.aefee/e Nuln’s dio satis Selene 2

(CLESRITICAGION » wins soceecatedaoedeReuts coeto tors Oncol nA Steen SAC aTOL aarti enna ar 2

Geographic and stratigraphic distribution .................cccceccceccceesccceccceeceseeseseeess 4

GEMUSPETD AGILE S are ec sane sce tictilan eel eere Aaee ecco wa oa Tine see asl dasaeeduloensnas 4

GEMS TEC CHILES ate ictaepai as Sok LSE = SOR Saeco diele wills Sn SS ueameued debe 7

GETMUST SCH IOMOCCT OS enero are cS oS eels ee eS Tal as aS BER ie sin BOAR aie Naee ETERS 8

GEMUSPBOCHITECS: AN Gv OUNETS 8 serie eoys ccs ascs ore Toor Seto woe scinla Siloro vie bio sfosjerdin wees slealone’s SMe 9

INI ROMIAT Me cee erree neceec sere cmc te ne ccc ccc: sociien nosonnes ea eaeen BA eet Capaae clan ORE 9

COMA Lame ese oa ie Tee cle isi aes Societe aes elders ine Shae ae Oule ac deleniowe ne declares 11

SEIMONTENY £As acdc aaaGhs ab AGGRO AGACTES ood Rupee Sea aN ane nn Iyer tn Seino a a 11

CO ANN PANT AINE eee soto Fe ee A rt RS eh dtigso gases jeu nitdwenetebinn oeee 15

IMI AAS tre GAM memes as wince Sans teers Mine vo ARMS S os ais ons lero uewnn Nee sla Mes swhceceeeenes 25

WWINORIESCCHOM epee re sere soon eesccecn. cic caren nes cates dedi tence eviecad se stds auc mcate usmnsseaes 33

SU CUNC MUNG Peremne Meer RGR erence ea sse stir clsnee cs vsces es ede gies pecseesatate seelseateenenes 35

Oinlamenmtawon pence oe cece sess eee ese Saws water loa ne soled Ne acid aaewaweNcenvas ebdaaeewlainade Ss 51

Absolute size, degree of taper, dimorphism and shape of the aperture ................... 59

PAD SOMILE SIZE err cree ee ee aac Cae ae ow ne oe aetotictcouse Fan sateekuinudscedecaee aja eee ses 59

ID CRTCCOIR Ape teen sce eect eee tae Rae ae aisles os.c cist ha sicte Sesie's vineieie dicen dates Sacleied 63

Dimorphism and shape Of the aperture .o..c2 o2. 2. . eee cmeeeceswseseacseecscocsiecsacees 64

SUUINIEM Ag eee Ee oe eo nice SE late cP esi Sein mies sonee dee see ete veemecineseee se Tk

General commentsyon Baculitidae <6. 2 ..sscce.<sc cece oheewee dec cvesceweseesdemocccbeeueeress a2

2 ANNALS OF THE SOUTH AFRICAN MUSEUM

APtyCh © oi.5 Sos aaie acenesinccnmactie vounse wed saceneenere ian snenine sein aeaarele sinc eco ose eee GZ

Mass! OCCUIFENCES fied st Teh eide cd eee ee oe CT coe re oS oe eee 79

Ammonitella and early ontogeny’ cic) b2..2005- 3. pace oes acne «ed esis ane eit ene eee 82

Muscle SCAtS! sc écatsen hos Pst locute eee ees oe Wak ons Sonal eb Senin CL eee 88

Drag bands and related features ai.0.5. occ ce sans cen iek ace ninweie + eb wesisle’s one ciotae eee 90

Microscul ptureS 3. scssciscee conceals an esese manos eaeiea eames ence hee a eee 94

Feather structure os os 55ceo.as sent oaecen tase en tie oo aio on oo oe bos is Ba anne eee Cee EEE 97

ParaSitiSi oo sna pasice sis aise ee ee EO oS oe esis Re 97

PYECALION «5 dieiacs geisiedec of eos os sist noe Sea a a Oeslete TIA Sei areinte oles is sintare oe eee 102

Annotated list of species referred to the family Baculitidae ......................0seeeeee: 112

Genus: BACULILCS sos cic5 Se ees ea sme aio eae OT eos A EEE 113

GENUS BOCHMOCETAS: ais bss sca voees vorcsk le coeaw es NO 218

Genus. CrioDGeUlites 56sec oy OC de eT ss ss 222

GeNUS EUDGCULILES MestiicseccekGh coccishs Meictn oe eR See ds 4 NO oe 222

GENUS FTES VET GT ee Oe, os a eae ne ten nae ee Rival aVOe ARCA EEE EERE 239

GENUS LOCH IES 2 sca hie Sa oh aloes Sas A ook EOS Felon Eee 241

Genus PSCUCODGCUIUES ccc eso nis eee eee ee a eb eS So Sli ao cis T Ode Uae e ECE 251

Genus | SCIPONOCEIAS 5. oo ccadshsaein a tenn cnc eaass Sawsnnne oo ie cia sansa ee See eee 255

Genus Trachybactlites” was eek aes sae tote sone te ses naa os Saas Soe 267

ACKNOWLEGREMEMS | .5 5. cs oscu, Hioe sa cadeudec ss oe os deealc tee eacrateme clas wocecre sar oelae eee 268

REfETENCES he ee a a ee a eee 269

INTRODUCTION

Following our description of the South African representatives of the genus

Eubaculites Spath, 1926 (Klinger & Kennedy 1993), and the other members of

the ammonite family Baculitidae Gill, 1871 (Klinger & Kennedy 1997), it

became clear that the members of the family exhibit wide intraspecific variation,

especially as far as ornament is concerned, and that several apparently endemic

lineages are present in the most species-rich genus, Baculites. Coupled with the

conservative shell morphology, these factors all severely limit the global

biostratigraphic potential of the family—especially of the genus Baculites, the

most common and stratigraphically longest-ranging representative of the family.

This is to be regretted, as representatives of the family are locally very

common (and may even be rock-formers) and have a wide geographic

distribution. The potential of the Baculitidae for biostratigraphic zonation is

clearly shown in the Campanian-lowest Maastrichtian of the US Western

Interior. Here, zonation is based predominantly on Baculites species, some of

which are estimated to have had life spans of as little as 500 to 900 thousand

years (Kennedy & Cobban 1976: 69). This zonation, in turn, is linked to the

most precise numerical time scale known for rocks of Cretaceous age

(Obradovich & Cobban 1975; Obradovich 1993).

From a consideration of the geographic and stratigraphic distribution,

morphology and phylogeny of the Baculitidae, we seek to identify certain

‘spikes’, patterns or common trends that may be used to enhance the

biostratigraphic potential of the family, especially of the genus Baculites.

CLASSIFICATION

Details of the systematics of the family were discussed earlier (Klinger &

Kennedy 1997) (Fig. 1) and we here only list the genera referred to the family

Baculitidae.

THE AMMONITE FAMILY BACULITIDAE 3

Hamites

: ; : a>)

Lechites Criobaculites fae}

| =

Sciponoceras Trachybaculites ‘es!

| Z

| ©

| 5)

| e-]

| >

L. (Tuberolechites) | | Fresvillia OQ

Jay s

i i | =

Baculites Lf =

| ! a

| |

SS aes =

Pseudobaculites |

| ] Q

| Boehmoceras | Eubaculites

Fig. 1. Classification and phylogeny of the family Baculitidae (from Klinger &

Kennedy 1997).

Genus and subgenus Lechites s.s. Nowak, 1908. Type species Baculites gaudini

Pictet & Campiche, 1861.

Subgenus Lechites (Tuberolechites) Cooper & Kennedy, 1977. Type species

Lechites (Tuberolechites) regifex Cooper & Kennedy, 1977.

Genus Sciponoceras Hyatt, 1894. Type species Hamites baculoides Mantell,

1822.

Genus Baculites Lamarck, 1799. Type species Baculites vertebralis Lamarck,

1799.

Genus Pseudobaculites Cobban, 1952. Type species Pseudobaculites nodosus

Cobban, 1952.

Genus Boehmoceras Riedel, 1931. Type species Ancyloceras krekeleri Wegner,

1905.

Genus Eubaculites Spath, 1926. Type species Baculites vagina Forbes var.

ootacodensis Stoliczka, 1866.

Genus Fresvillia Kennedy, 1986a. Type species Fresvillia constricta Kennedy,

1986a.

Genus Criobaculites Klinger & Kennedy, 1997. Type species Hamites trabeatus

Morton, 1834.

4 ANNALS OF THE SOUTH AFRICAN MUSEUM

Genus Trachybaculites Cobban & Kennedy, 1995. Type species Baculites

columna Morton, 1834.

GEOGRAPHIC AND STRATIGRAPHIC DISTRIBUTION

The ammonite family Baculitidae (as a whole) has a world wide distribution,

from the Arctic Circle to Antarctica, but individual genera show differing

geographic distributions.

For the purpose of this discussion, we recognize the following, partly

arbitrary and overlapping geographic regions where Baculitidae occur:

1. Europe—this includes both Boreal and Tethyan regions; baculites do not

appear to have been affected by the Boreal/Tethyan Realm division.

2. Indo-Pacific—this is a very large area and includes the Indian Ocean areas

(Zululand, Madagascar, India, Australia) and the Pacific Ocean areas (e.g.

Japan, west coast of North and South America), as well as the Subantarctic

Peninsula and Patagonia. The Austral and Neuquén basins of Argentina are also

included in this region.

3. Gulf-Atlantic region, i.e. Gulf Coast and Atlantic Seaboard regions of North

America and the Afro-American South Atlantic coastal regions (e.g. Angola,

Nigeria, Cameroon and Brazil), excluding the Austral and Neuquén basins of

Argentina.

4. The US Western Interior region, as defined by Cobban & Reeside (1952),

including its extension into Canada and, at least in the pre-Late Campanian, into

West Greenland (Birkelund 1965: 47).

5. North Africa and the Middle East.

As far as stratigraphic distribution is concerned, we must emphasize that the

stage and substage boundaries of the different bio-geographic regions can only

be regarded as approximate temporal equivalents. It became perfectly clear

during the 1995 Symposium on Stage Boundaries in Brussels that, as yet, there

is no consensus on most of the boundaries. The stratigraphic ranges of the

various taxa listed and shown in the illustrations below are taken from various

literature sources, and we are fully aware of the fact that the stage and substage

definitions from the various regions probably do not coincide and can only be

regarded as approximate.

Genus Eubaculites

The genus Eubaculites is well known, mainly as a result of recent

comprehensive studies by Klinger (1976), Henderson et al. (1992), Kennedy &

Henderson (1992), and Klinger & Kennedy (1993). It is therefore appropriate to

discuss the geographic and stratigraphic distribution of the genus first, and

compare the distribution of other genera with this pattern.

Eubaculites first appeared in the Upper Campanian, as E. occidentalis

(Meek, 1862), became locally common in the Maastrichtian, and ranged into the

upper Upper Maastrichtian as E. carinatus (Morton, 1834) in the Pétites-

Pyréneés in southern France (Kennedy 1986e), the Biscay Region (Ward &

Kennedy 1993), and the U.S. Gulf coast ((Kennedy, unpublished data). Apart

THE AMMONITE FAMILY BACULITIDAE 5

from E. occidentalis and E. carinatus, four other well-defined species are

referred to the genus—E. labyrinthicus (Morton, 1834), E. vagina (Forbes,

1846), E. simplex (Kossmat, 1895), and E. latecarinatus (Brunnschweiler,

1966).

Eubaculites is most common in the Gondwanan Region (Fig. 2) and, in

places, is the dominant Maastrichtian heteromorph ammonite. It occurs in great

numbers in Western Australia, South India, Zululand, Madagascar, southern

Chile, the Neuquén Basin of Argentina, and locally in the U.S. Gulf coast

region. A few specimens are also known from deposits off the southern Cape

coast of South Africa (Klinger & Kennedy 1980) and from Mozambique (Crick

1924).

Some of the material described by Haas (1943) as Baculites anceps from

Angola may be Eubaculites (see Klinger & Kennedy 1993: 218). If it is a

Eubaculites, this is the only record of the genus from the west coast of Africa.

Given our extensive knowledge of Nigerian Maastrichtian faunas, the absence

of Eubaculites is conspicuous. Equally, there are, as yet, no records from

Brazil, New Zealand or Antarctica. The Baculites cf. vagina recorded by

Woods (1917: 36, pl. 20 (fig. 5a-d)) from Amuri Bluff, New Zealand, is too

poorly preserved for positive identification (see Henderson 1970: 24-25),

Marshall (1926: 155) thought that it ‘almost certainly belongs to’ B. rectus.

Occurrences in the Northern Hemisphere are scattered compared to those of

the Gondwanan Region. In North America, Eubaculites is unknown from the

US Western Interior seaway. It occurs in California and British Columbia, in

the Gulf Coast Region in the Corsicana Formation of Texas, the Owl Creek

Formation of Missouri and Mississippi, the Prairie Bluff Chalk of Alabama,

Severn Formation of Maryland, Navesink Formation of New Jersey, and

reworked into the overlying Palaeocene Hornerstown Formation.

In Europe, Eubaculites occurs in the Lower Maastrichtian of Austria, and

the Upper Maastrichtian of south-eastern France, the Biscay Region, northern

Spain (unpublished data), and a single specimen is known from Maastricht in

the Netherlands. This latter occurrence is North Tethyan; the associated fauna

includes rudists.

Again, the absence of Eubaculites from well-documented Maastrichtian

deposits of northern Europe, Greenland, North Africa, the Middle East, and

Central and Eastern Asia is striking. No typical Maastrichtian Eubaculites has

as yet been recorded from Japan.

The distribution of the genus in the Maastrichtian (Fig. 2) shows that

Eubaculites is latitudinally restricted. It is common in the southern temperate

regions, rare in the northern temperate regions, and absent from the equatorial,

Arctic and Antarctic regions, and from the west coast of Africa.

The geographic and stratigraphic distribution of the individual species

varies.

The oldest, E. occidentalis occurs in the Upper Campanian-Lower

Maastrichtian of California and British Columbia.

Eubaculites carinatus has the widest geographic, and longest stratigraphic

distribution. It occurs in both the Lower and Upper Maastrichtian, and has been

recorded from Austria, south-eastern France, the Biscay Region, the

Netherlands, North America (California, Texas, Missouri, Mississippi,

ANNALS OF THE SOUTH AFRICAN MUSEUM

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THE AMMONITE FAMILY BACULITIDAE 7

Alabama and New Jersey), Neuquén Basin of Argentina, Quiriquina Island,

Chile, Zululand, Mozambique, Madagascar, South India and Western Australia.

From the available published data it appears that E. carinatus is common in

Western Australia, Zululand, Chile, Argentina, the Owl Creek Formation of

Missouri and Mississippi, uppermost Corsicana Formation (Kemp Clay) in the

Brazos River section, Texas (Kennedy, unpublished data), and apparently in the

Biscay Region. In contrast to. the mass occurrences in Zululand and Western

Australia, it is rare in Madagascar and South India.

Eubaculites labyrinthicus is only known by a few specimens from the

Maastrichtian Prairie Bluff Chalk of Alabama, the Navesink Formation of New

Jersey, and the St Lucia Formation of Zululand. In contrast, Collignon (1971:

15) recorded it (as B. simplex) as being abundant in the Lower Maastrichtian of

Madagascar, where it also grows to large sizes.

Eubaculites latecarinatus first occurs in Maastrichtian a, and is very

abundant in Maastrichtian I and II of the St Lucia Formation of Zululand. It is

also known from offshore deposits on the Alphard Group off the southern Cape

coast. Henderson et al. (1992) described only eleven specimens from Western

Australia. It is rare in Madagascar, where apparently only two specimens are

known (Klinger & Kennedy 1993, fig. 48), and none from South India or South

America. Kennedy et al. (1997a: 20, figs 15a-d, k, 1, 16g, h, 19) described

three specimens from the Upper Maastrichtian Severn Formation of Maryland.

Eubaculites simplex is known only by the lectotype from India, a single

specimen from Madagascar (as Baculites occidentalis in Collignon 1971: 15,

pl. 645 (fig. 2390)), but it is common in Western Australia (132 specimens). In

Zululand about 20 specimens are known and in Santa Cruz Province, Patagonia,

five specimens (as Baculites sp. in Hunicken 1965: 64, pl. 2 (figs 3-4), pl. 3

(figs 5-6), pl. 5 (fig. 5), pl. 6 (figs 6-7), pl. 8 (figs 2-5)).

Eubaculites vagina is the dominant species of Eubaculites in South India,

rare in Madagascar, and as yet unknown elsewhere. Fatmi & Kennedy (1999)

have shown that Baculites binodosus Noetling, 1897, should be referred to the

genus Eubaculites. Apart from the holotype, only six other specimens are

referred to this species, all from the Upper Maastrichtian of Baluchistan; India.

To summarize, Eubaculites thus seems to comprise a mixture of widely

distributed and endemic or geographically restricted, and long- and short-

ranging species; its absence from the US Western Interior seaway and West

Africa is conspicuous, as is its absence in high latitudes and equatorial regions.

With these data as a basis, it is now possible to see if the other baculitid genera

follow the same patterns, and, if not, where and to what extent they differ.

Genus Lechites

Lechites is a short-lived genus restricted to the Upper Albian. It is widely

distributed in Western and Central Europe, including southern England, France,

Switzerland, Austria, Hungary, Romania, Spain, Sardinia, Central Iran, North

Africa, Madagascar, Zululand, South India, Hokkaido, Mexico, Texas and

Antarctica. Again, the absence of the genus from West Africa and Brazil is

conspicuous.

8 ANNALS OF THE SOUTH AFRICAN MUSEUM

It is difficult to determine the distribution of the individual species because

of lack of uniform taxonomy. Lechites gaudini (Pictet & Campiche, 1861)

(which includes L. communis Spath, 1941, L. raricostatus Breistroffer, 1947,

L. italicus Wiedmann & Dieni, 1968, L. campichei Renz, 1968, L. vraconensis

Renz, 1968, and L. fasciatus Scholz, 1971, according to Cooper & Kennedy

1977: 644-645), has the widest geographic and longest stratigraphic

distribution. It has been recorded from the Upper Albian of Western and

Central Europe, Sardinia, North Africa, Madagascar, Zululand, South India,

Hokkaido and doubtfully Mexico.

Lechites moreti Breistroffer, 1936, occurs in the Upper Albian of southern

England, France, Switzerland, Spain, Sardinia, Hungary and North Africa, and

possibly Texas (as L. comanchensis Adkins, 1920 in Clark 1965, pl. 3 (figs 2,

6)) (fide Cooper & Kennedy 1977: 652). Lechites comanchensis is only known

from the Upper Albian of Texas. Lechites antanimangaensis Collignon, 1964,

from the Lower Cenomanian of Madagascar, is known from the holotype only,

and is based on a large body chamber fragment. It has sufficient characters in

common with macroconchs of Sciponoceras baculoides as to suggest it to be a

Sciponoceras (see Wright & Kennedy 1995: 314). Lechites varicostatus Chiriac,

1981, is only known from the Upper Albian of Romania.

Lechites thus also includes both cosmopolitan and endemic, and long- and

short-lived forms, but, in contrast to Eubaculites, it occurs in high southern and

northern latitudes. Given the extensive descriptions of Albian faunas from

Angola, the absence from this region is conspicuous, as is that from Nigeria.

Genus Sciponoceras

Sciponoceras is widely known from the Lower Cenomanian to Upper

Turonian in both hemispheres, but again is essentially absent from West Africa;

a single fragment has been recorded from Angola (Cooper 1978: 70,

fig. 12b-d). The specimen figured by Solger (1904: 101, figs 3-4) as Baculites

cf. gracilis is of doubtful affinities (cf. Reyment 1955: 15).

Some species have wide distribution. Sciponoceras roto Cieslinski, 1959,

which occurs in the Lower Cenomanian of southern England, France, Germany,

Switzerland, Spain, Mangyschlak, Iran, Algeria, Tunisia, Tanzania,

Madagascar, Zululand, and is said to range through the whole of the

Cenomanian of Poland; S. baculoides (Mantell, 1822) is common (sometimes in

flood abundance) in the lower part of the Middle Cenomanian of England and

France, and ranges to the lower Upper Cenomanian; other records are from

West and Central Europe, North Africa, South India, Mozambique, Zululand,

Madagascar, California and Hokkaido (as a distinct subspecies perhaps),

Washita Group, Texas and doubtfully Argentina (Riccardi & Aguirre Urretta

1988: C381); S. gracile (Shumard, 1860) is most common in the eponymous

Upper Cenomanian zone of the US Western Interior (where it ranges to the top

of the Cenomanian), and also occurs in California, Texas, northern Mexico,

southern England, northern France, southern Germany and doubtfully Angola.

Other species are more restricted geographically: Sciponoceras bohemicum

(Fritsch, 1872) is only known from the Turonian of England, northern France,

southern Germany and Bohemia; S. cucullatum Collignon, 1964, from the

THE AMMONITE FAMILY BACULITIDAE 5

Lower Cenomanian of Madagascar and Lower or Middle Cenomanian of

Zululand; S. glaessneri Wright, 1963, Middle Cenomanian of Bathhurst Island,

S. intermedium Matsumoto & Obata, 1963, Upper Turonian of Hokkaido;

S. orientale Matsumoto & Obata, 1963, Lower and Middle Turonian of

Hokkaido, and S. subbaculoides (Geinitz, 1875), Cenomanian of Bohemia and

Hungary; S. kossmati (Nowak, 1908), Turonian of India (fide Kennedy 1988:

108-109); S. santacrucense Leanza, 1970, Turonian of Patagonia, and

S. matsumotoi Inoma, 1980, from the Cenomanian of Hokkaido.

The virtual absence of the genus from West Africa is of note, as is the near-

cosmopolitan distribution of the typical US Western Interior species S. gracile,

in contrast to the marked endemism of the later Baculites species of the region.

Genus Baculites

The geographic and stratigraphic distribution of most of the well-known,

and/or adequately defined species of Baculites, as well as of Eubaculites,

Boehmoceras, Fresvillia, Criobaculites and Trachybaculites is illustrated in

Figures 3-19. The exact stratigraphic range of individual species is often poorly

constrained, and there are problems in global correlation of stage boundaries

and substages as indicated above (p. 4). Unless otherwise indicated, the strati-

graphic ranges of the species in the diagrams can only be considered as

approximate.

TURONIAN

Fig. 3

Only four definite Baculites species are known from the Turonian:

B. yokoyamai Tokunaga & Shimizu, 1926, B. undulatus d’Orbigny, 1850,

B. calamus Morrow, 1935, and B. embaensis Balan, 1982. Two forms in open

nomenclature have been recorded from Romania (Szasz 1986). Neal Larson has

informed us that Dr W. A. Cobban identified a nodose baculitid from the Carlile

Formation of Wyoming as cf. B. boulei. We suspect that B. embaensis, with

deep constrictions may be a Sciponoceras and we omit this species from the

diagram.

Baculites yokoyamai is best known from the US Western Interior, where it

ranges throughout the stage. Summesberger (1992: 124, pl. 8 (figs 10-11))

recently recorded B. yokoyamai from the Middle Turonian of Austria. Baculites

calamus is rare, and only known from the Middle Turonian Collignoniceras

woollgari Zone of the US Western Interior, and, according to Neal Larson,

based on one specimen only.

Baculites undulatus is best known from the Upper Turonian of Western

Europe and Hokkaido, and was also recently recorded from Trans-Pecos Texas

(Kennedy et al. 1989), and, according to Dr W. A. Cobban is rare in the Upper

Turonian of New Mexico. It may possibly already be present in the Lower

Turonian of Romania (Szdsz 1986: 120, pl. 1 (figs 1-2)), accompanied by a

questionable baculitid with ornament very similar to that of B. calamus.

The first nodose baculitid (Klinger & Kennedy 1997, fig. 132) may possibly

occur in the Upper Turonian of Angola and is the oldest definite record of the

10 ANNALS OF THE SOUTH AFRICAN MUSEUM

TURONIAN

B. undulatus

Q Ws B. yokoyamai

Pea Woe SY

B. ? sp. [Szasz 1988]

Se Sa

Of) yp

qdOdna

JITOVd-OGNI

~ -B. undulatus

Mek eee

B. sp. nov. [Angola]

B. yokoyamai

B. calamus

Ol) WD)

Fig. 3. Geographic and stratigraphic distribution of species of the genus

Baculites in the Turonian.

JIINWILV-ATOAO

“INT LSHM °S°

THE AMMONITE FAMILY BACULITIDAE 11

family from Angola. Dr W. A. Cobban has informed us that nodose baculitids

also occur sparingly in the Upper Turonian of the US Western Interior.

Turonian Baculites are a rarity and species diversity is low. Even though

B. yokoyamai and B. undulatus are predominantly US Western Interior and

Indo-Pacific-European species respectively, some faunal migration and mixing

seems to have taken place during the Turonian.

CONIACIAN

Fig. 4

Baculites yokoyamai survives into the Lower Coniacian in the US Western

Interior, the Indo-Pacific and the Gulf-Atlantic regions, but all the other species

recorded from the Coniacian appear to be geographically restricted.

The baculitid faunas of the US Western Interior are predominantly endemic

from the Middle Coniacian and remain so until the Maastrichtian (see Cobban

1994 for a detailed discussion). Only a few typical US Western Interior species

have been recorded outside this region, and vice versa. These include: two

Campanian and one Maastrichtian species recorded from Belgium by Kennedy

(1993), Baculites undatus Stephenson, 1941, a typical Gulf Coast and Atlantic

Seaboard species recorded from the Western Interior by Cobban (1973),

B. haresi, a typical Lower Campanian Western Interior species recorded from

the Atlantic Seaboard by Kennedy & Cobban (1993e), B. reduncus, another

typical US Western Interior species recorded from Arkansas by Kennedy &

Cobban (1993a), B. crickmayi Williams, 1930, from the Annona Chalk of

Arkansas by Kennedy & Cobban (1993a), Baculites gr. of aquilaensis in

northern Texas by Cobban & Kennedy (1992a), and B. mclearni Landes, 1940,

also in Texas by Cobban & Kennedy (1993). The noded variant of B. scotti in

Cobban (1958: 662, pl. 90 (figs 5-9)) (= B. texanus Kennedy & Cobban, 1999)

has also been recorded from Texas, New Jersey, South Dakota and Colorado.

Pseudobaculites, with its complex suture has no equivalents outside the Western

Interior in the Coniacian, and again in the Upper Campanian and Lower

Maastrichtian. Baculites mariasensis Cobban, 1951, and B. sweetgrassensis

Cobban, 1951, are analogues, but not equivalents of B. yokoyamai of the Indo-

Pacific region. Baculites codyensis Reeside, 1927a, is comparable, but not

identical to the tuberculate group of B. incurvatus Dujardin, 1837, and

B. brevicosta Schliter, 1876, of Europe, and B. capensis Woods, 1906, of the

Indo-Pacific. Smooth B. bailyi Woods, 1906, from the Indo-Pacific has no

equivalents in the Coniacian of Europe.

SANTONIAN

Figs 5-6

Nodose to ribbed forms of the Coniacian groups of Baculites codyensis,

B. incurvatus and B. capensis persist into the Middle Santonian of the US

Western Interior and Europe and Lower Campanian of the Indo-Pacific

respectively. Baculites capensis has recently been recorded from the Gulf Coast

by Kennedy & Cobban (1991b). Even though some individuals of these three

species from these regions are morphologically inseparable, populations are

ANNALS OF THE SOUTH AFRICAN MUSEUM

CONIACIAN.

B. incurvatus

qAdOWNa

B. boulei B."schencki" B."boulei"

E L U I

B. yokoyamai

B. capensis

JIAIOVd-OdNI

“INI ISHM °S°O | OLINWITLVAW1INOD

Fig. 4. Geographic and stratigraphic distribution of species of the genera Baculites and

Pseudobaculites in the Coniacian.

THE AMMONITE FAMILY BACULITIDAE 13

SANTONIAN |

a oan

Boehmoceras

AdoOwna

B. capensis

omy rye

B. princeps

OL) | 8 ear

B. pseudobaculus

eo)

an)

-Q

B. delvallei

Fig. 5. Stratigraphic distribution of species of the genera Baculites and Boehmoceras in the

Santonian in Europe and the Indo-Pacific.

14 ANNALS OF THE SOUTH AFRICAN MUSEUM

SANTONIAN

B. capensis

ESS

JLINV TLV-ATOAS

LSVal “ddr

7 VOTHAV °N

Fig. 6. Stratigraphic distribution of species of the genera Baculites and Boehmoceras in the

Santonian in the Gulf-Atlantic Region, US Western Interior and Greenland, North Africa,

and the Middle East.

THE AMMONITE FAMILY BACULITIDAE 15

significantly distinct. Smooth B. bailyi also persists up to the Lower Campanian;

smooth B. nugssuaquensis Birkelund, 1965, from the Santonian of Greenland is

an analogue. Baculites thomi Reeside, 1927b, has no equivalents outside the US

Western Interior.

Several smooth forms occur in the Middle and/or Upper Santonian of the

Indo-Pacific region, e.g. B. bailyi, B. uedae Matsumoto & Obata, 1963, and

B. delvallei Riccardi, 1980. These are analogues of European B. fuchsi

Redtenbacher, 1873. Baculites kirki Matsumoto, 1959, with a ventral keel, is

endemic to the Santonian of California. Boehmoceras, as B. arculus (Morton

1834) (=B. loescheri Riedel, 1931) and B. krekeleri (Wegner, 1905) are rarities

in the Upper Santonian of Europe, but are abundant in the US Gulf Coast as

B. arculus.

To summarize, similarities may still be seen in some Santonian baculitids

from the different regions, but most of the species appear to be endemic,

especially those of the US Western Interior.

CAMPANIAN

Figs 7-14

In contrast to the Turonian to Santonian stages, radiation of Baculites in the

Campanian appears explosive. However, if the duration of the stage is taken

into account, the rate of evolution does not seem excessively accelerated.

The best documented succession is that of the US Western Interior

(Figs 7-9). Lower Campanian species include B. aquilaensis Reeside, 1927b,

B. haresi Reeside, 1927b, and B. thomi, the latter a survivor from the

Santonian. The Middle and Upper Campanian succession consists of a series of

ribbed and smooth species (see Scott & Cobban 1965, map I-439), many of

them restricted to the Western Interior, with others also known from the Gulf

Coast region. The difference between US Western Interior and Gulf Coast

Baculites faunas (see discussion above, p. 11) may be more apparent than real.

The Gulf Coast ammonite record is limited to a small number of levels only,

and most of these include Western Interior taxa (Ozan, Annona, Yancy, Wolfe

City Sand, spiniger fauna, etc.).

The succession in the Indo-Pacific is not quite as clear. Lower Campanian

smooth species include Baculites bailyi and B. delvallei, both survivors from the

Santonian and probably synonyms, and B. chicoensis Trask, 1856; nodose

species include descendants of B. capensis, B. tanakae Matsumoto & Obata,

1963, B. menabensis Collignon, 1969, and B. sparsinodosus Collignon, 1969;

all probably synonyms of B. tanakae. Baculites sulcatus Baily, 1855, is

restricted to Pondoland and subsurface Zululand. In the Middle Campanian of

Zululand and Madagascar, B. vanhoepeni Venzo, 1936, and B. increscens

Collignon, 1969, are generally strongly ornamented, and homoeomorphic with

the B. obtusus—B. asperiformis series of the US Western Interior. Baculites

regina Obata & Matsumoto, 1963, from Honshu could possibly belong to this

group. Baculites nibelae Klinger & Kennedy, 1997, succeeds this group in

Zululand and equivalent forms in Madagascar and possibly the Middle East.

Baculites duharti Hiinicken, 1975, is a large, smooth baculite of the Middle to

Upper Campanian of Zululand and Patagonia. Baculites kotanii Matsumoto et

16 ANNALS OF THE SOUTH AFRICAN MUSEUM

LOWER CAMPANIAN

B. sp. weak flank ribs

7 )

“3

)

Baculites sp. smooth

OES

Fig. 7. Stratigraphic distribution of species of the genus Baculites in the Lower Campanian

in the US Western Interior.

THE AMMONITE FAMILY BACULITIDAE

MIDDLE CAMPANIAN

B. obtusus NOT TO SCALE

| i

B. mclearni

OP2

B. asperiformis

ened

QUE

B. sp. smooth

‘ae

| B. naples

ine

SW ODE

| | |

B. gilberti

roe Sees | O pean

B. pero

nf reduncus

| B. scotti

megane

YOIAALNINAALSAM’S |

Fig. 8. Stratigraphic distribution of species of the genus Baculites in the Middle Campanian

in the US Western Interior.

18 ANNALS OF THE SOUTH AFRICAN MUSEUM

UPPER CAMPANIAN

B. pseudovatus NOT TO SCALE

0 : y or P. natosini

| z

B. crickmayi |

B. rugosus |

00) Bn ne

B. corrugatus | |

fee ined

B. compressus |

NYALSAM’S 1

YOrwalni

B. aeguRrae

a =

B. jenseni

Fig. 9. Stratigraphic distribution of species of the genus Baculites in the Upper Campanian

in the US Western Interior.

THE AMMONITE FAMILY BACULITIDAE 19

al. 1980, is a small, smooth baculite apparently endemic to Japan. Baculites

rectus Marshall, 1926, is a smooth baculite, probably from the Upper

Campanian of New Zealand, Campanian s.1. of South Africa?, and Antarctica.

Baculites subanceps Haughton, 1925, is widespread in the Upper

Campanian, occurring in Angola, South Africa, California, British Columbia,

Hokkaido, Antarctica, and the Middle East; smooth forms are very similar to

Eubaculites occidentalis.

In the Gulf-Atlantic region individual species of Baculites are common to

abundant, but diversity is low. Two fragments of B. tanakae? (as B. capensis)

are recorded from the Lower Campanian of Angola by Cooper (1988), and

B. subanceps from the Upper Campanian and/or Lower Maastrichtian. Middle

Campanian B. taylorensis Adkins, 1929, is similar to B. vanhoepeni from the

Middle Campanian of the Indo-Pacific region. Baculites claviformis Stephenson,

1941, B. undatus Stephenson, 1941, and B. ovatus Say, 1821, occur in the

Upper Campanian of the Gulf Coast and Atlantic Seaboard. There are doubtful

records of B. ovatus from the Middle East, and we have seen a single specimen

from southern France. Baculites undatus has recently been recorded from the

Western Interior (Cobban 1973; Cobban et al. 1992; Larson et al. 1997), and,

with B. texanus is a Gulf Coast immigrant into that region during the

Campanian. Lower Campanian B. haresi, a typical Western Interior species,

was recorded from New Jersey by Kennedy & Cobban (1993e).

The baculites of the Middle East (Figs 192-195) are largely undescribed.

North African Baculites cf. ovatus and B. subanceps (Luger & Gréschke 1989)

probably connect with the Gulf-Atlantic and Indo-Pacific regions.

Trachybaculites? furcillatus (Blanckenhorn, 1905) (Taubenhaus 1920) is

endemic to Israel. Lefeld & Uberna (1992) mentioned Baculites from Libya, but

the material has yet to be described.

Apart from Baculites leopoliensis Nowak, 1908, the European Campanian

Baculites succession is still largely unknown, and details of the systematics still

have to be worked out. The material is generally poorly preserved and lacks key

morphological characters in most cases. Only Baculites leopoliensis Nowak,

1908, can be regarded as well-characterized. (However, see comments on the

age of B. leopoliensis in the appendix.) The remaining species are based on a

few fragments and most are best regarded as nomina dubia in the absence of

data on intraspecific variation. Some of the baculites are very similar to

B. tanakae and B. vanhoepeni of the Indo-Pacific. Kennedy & Jagt (1995)

described a new species, B. vaalsensis, from the Lower Campanian Vaals

Formation of the Aachen area (Germany, Belgium and the Netherlands); it also

occurs in southern England and New Jersey (Kennedy et al. 1997b). Santamaria

Zabala (1996) described two new species from northern Spain: B. alonsoi

Santamaria Zabala (1996: 13, pl. 3 (figs 3-6)) and B. alavensis Santamaria

Zabala (1996: 14, pl. 3 (figs 7-9)) as well as two other species in open

nomenclature. Baculites alavensis is similar to B. leopoliensis and was

described fully on the basis of additional material (Klinger & Kiichler 1998).

The recent records of Baculites aquilaensis and Baculites ‘smooth species’

from Belgium by Kennedy (1993: 110, pl. 4 (figs 22-24), p. 111, pl. 5

(figs 1-12), pl. 6 (figs 5-10), text-fig. 6, respectively), typical Lower and

Middle Campanian species from the US Western Interior are at first surprising,

20 ANNALS OF THE SOUTH AFRICAN MUSEUM

CAMPANIAN

B. sulcatus

OLD

B. menabensis

vo, gaa)

B. sparsinodosus

Crea

B. tanakae

O B. ankilizatensis

See eT, Sit:

B. delvallei ()

OL)

B. basset

Ola

B. increscens

OLZ2)

B. vanhoepeni

U B. nibelae

Mormitlovd-OdNiI

Fig. 10. Stratigraphic distribution of species of the genus Baculites in the Campanian in the

Indo-Pacific (1).

THE AMMONITE FAMILY BACULITIDAE 21

CAMPANIAN

B. chicoensis

Bae _ __B. duharti

PAPA Pts B. kotanii

B. lomaensis

E. occidentalis

| a

B. rectus

me Motes) Of

B. regina

ee LALLA

ZWOTAIOVd-OCNI

B. rex

fete VL_

Mh es ©] 0

iLL

B. inornatus

OL) fangs

Fig. 11. Stratigraphic distribution of species of the genera Baculites and Eubaculites in the

Campanian in the Indo-Pacific (2).

22 ANNALS OF THE SOUTH AFRICAN MUSEUM

CAMPANIAN

MIDDLE

B. tanakae [Cooper]

si eas

B. haresi

OID) 8 aye ae

B. taylorensis

OWIy

B. subanceps P)

a co

Shee () i is Z =

B. cLavasOnms ea

(222) | >

_B. ovatus _ |

= o

JD) |>

B. undatus Z,

etree OLD) |=

B. crickmayi Ol

E L vl ee Rig Se

2%") Of)

B. mclearni

ERS

yfts Wee] O

B. reduncus

OP)

Fig. 12. Stratigraphic distribution of species of the genus Baculites in the Campanian in the

Gulf-Atlantic Region.

THE AMMONITE FAMILY BACULITIDAE 23

CAMPANIAN

Je a ae

B. pe RS ae 1. cf. asper

° aa

B. sp. 2. cf. haresi

OLD

B. sp. 3. cf. increscens

OLZ2)

B. subanceps

[?=B. scotti in Hassan]

yy )

VW WIM

B. cf. ovatus

VDD»)

B. palestinensis

VOIATAV HLAUON F LSVA ATIACACIN

Fig. 13. Stratigraphic distribution of species of the genus Baculites in the Campanian in the

Middle East and North Africa.

24 ANNALS OF THE SOUTH AFRICAN MUSEUM

CAMPANIAN

“-- B. vaalsensis

ne/2

B. sp. 1 [Holzapfel] | _

B. sp. [Miller & Wolleman]

[EES

0) es

B. aquilaensis |

OpD

B. alonsoi

ores

B. alavensis

B. smooth sp.

©) ais

B. leopoliensis

OVA

OL)))

Fig. 14. Stratigraphic distribution of species of the genus Baculites in the Campanian in

THE AMMONITE FAMILY BACULITIDAE 25

but the poor record at other levels and localities conceal other common taxa. If

these are indeed correctly identified, it certainly alters our current ideas about

the currently perceived endemic character of the US Western Interior baculitid

faunas.

MAASTRICHTIAN

Figs 15-19

The Maastrichtian is characterized by the dominance of Eubaculites in the

southern Gondwanan regions, and by the presence of long-ranging Baculites

species in Europe.

In contrast to the Campanian, the Maastrichtian baculite species of the

Western Interior are few in number. The Campanian lineage of B. compressus

Say, 1821, ended in the basal Maastrichtian with B. eliasi Cobban, 1958. This

fauna is replaced by B. baculus Meek & Hayden, 1861, an immigrant from the

Gulf Coast region. Baculites baculus gave rise to B. grandis Hall & Meek,

1854, and B. clinolobatus Elias, 1933, all large forms with broad, rib-like

swellings on the flanks. The last baculitids of the Western Interior (Cobban &

Kennedy 1992c) are all atypical. Baculites larsoni Cobban & Kennedy, 1992c,

is very small, Trachybaculites columna (Morton, 1834) has circumperipheral

ribbing and merits generic separation (Cobban & Kennedy 1995); it has recently

been recorded from the Netherlands (Kennedy & Jagt 1998). Baculites sp. is a

curved Boehmoceras homoeomorph, probably referable to Criobaculites.

Fragments of a more typical large Baculites occur in the Jeletzkyites

nebrascensis Zone in the Pierre Shale near Verdigre, Knox County, Nebraska

(Kennedy et al. 1998).:

European baculites, are very characteristic. Eubaculites carinatus is a rarity,

as is Upper Maastrichtian Fresvillia. Baculites knorrianus Desmarest, 1817, is a

large, smooth species, typically Lower Maastrichtian but occurring in the lower

Upper Maastrichtian in Denmark. Baculites vertebralis Lamarck, 1801, and

B. anceps Lamarck, 1822, are best known from the Upper Maastrichtian; in

Denmark they persist to the end of the stage (Birkelund 1979, 1993). Very

badly preserved specimens referred to both species have been recorded from the

Lower Maastrichtian of Zumaya (Ward et al. 1991; Ward & Kennedy 1993).

Baculites baculus, a typical Gulf Coast and US Western Interior species, has

recently been recognized from Belgium (Kennedy 1993: 110, pl. 4 (figs 10,

20-21)).

As discussed above, the Indo-Pacific, especially the south Gondwanan

regions, are dominated by Eubaculites species. Several species of Baculites

appear to span the Campanian-Maastrichtian boundary (or are dated no more

accurately than the high Campanian or low Maastrichtian). These include

B. rectus, B. subanceps and B. rex Anderson, 1958. Trachybaculites vicentei

(Stinnesbeck, 1986) and B. huenickeni Stinnesbeck, 1986, are endemic to Chile,

B. ambatryensis is only known from Madagascar and Pakistan (Fatmi &

Kennedy 1999). Eubaculites binodosus (Noetling, 1897) is based on the

holotype and six other specimens and is only known from the Upper

Maastrichtian of Baluchistan (Fatmi & Kennedy 1999).

26 ANNALS OF THE SOUTH AFRICAN MUSEUM

MAASTRICHT.

NOT TO SCALE

B. eliasi

B. baculus

ODD)

B. grandis

aes:

Of) “hear

B. clinolobatus

Tees

NYA LSAM'S |

OL)

T. columna

meer ALU OPP]

B. larsoni

Rei es) ON

P. natosini Criobaculites

RPE

pina OKO

YVOIAALNI

Fig. 15. Stratigraphic distribution of species of the genera Baculites, Pseudobaculites

and Criobaculites in the Maastrichtian in the US Western Interior.

THE AMMONITE FAMILY BACULITIDAE

MAASTRICHT.

B. eB veriebralis iene

ei, | OL

B. anceps

B. knorrianus

E. carinatus

Ord»)

Adowond

B. baculus

OLD)

T. columna

Olds)

Fig. 16. Stratigraphic distribution of species of the genera Baculites, Trachybaculites

Eubaculites and Fresvillia in the Maastrichtian in Europe.

ANNALS OF THE SOUTH AFRICAN MUSEUM

MAASTRICHT.

iB. ambatryensis E L U

| B. subanceps

Tee

-- B. vicentei

OL ry ;

- B. (E.) huenickeni S

. °

ae <

ro- B. rex | Boat: B. columna >

er pins @)

OT _Joapyees [ORI :

fossrae E. latecarinatus E. vagina poe

OL 0

E. carinatus

E. labyrinthicus

VE3)

E. binodosus

Sea

E. simplex ©)

Vie

E. occidentalis

Fig. 17. Stratigraphic distribution of species of the genera Baculites, Eubaculites and

Fresvillia in the Maastrichtian in the Indo-Pacific.

THE AMMONITE FAMILY BACULITIDAE

MAASTRICHT.

B. subanceps

0 Ph ig

Wi Ml sf

posse B. anceps [Haas 1943]

| eT

:-- B. claviformis

i () DDD

CMMI ONATTTT

Gas B. teicherti

OTT) tir

---- B. ovatus

ODD) i

|--- B. undatus C. trabeatus

“OL2®) OCW

--- B. kegeli T. columna

-OUIT OWE) nf EP ALU

E L

OT NV ie Ved ta 2D

B. baculus

ODD)

E. carinatus

Psi

nk an E. oe.

me [EEE

Fig. 18. Stratigraphic distribution of species of the genera Baculites, Trachybaculite

Eubaculites, and Criobaculites in the Maastrichtian in the Gulf-Atlantic Region.

30 ANNALS OF THE SOUTH AFRICAN MUSEUM

MAASTRICHT.

NOT TO SCALE

B. "anceps" (smooth)

Of) LR esry see

B. "anceps" (ornate)

or ail

B. "ovatus"

POD) ay ae

B. libyensis

VD2P)

B? paradoxus

LSVH #IGCGCIN F VOIAAV HLAON

ODD)

Fig. 19. Stratigraphic distribution of species of the genus Baculites in the Maastrichtian in

North Africa and the Middle East.

THE AMMONITE FAMILY BACULITIDAE 31

In the Gulf-Atlantic Region, several species seem to span the Campanian-

Maastrichtian boundary: B. subanceps, B. claviformis and B. undatus. Baculites

teicherti Reyment, 1956, is endemic to Nigeria but may be a B. subanceps.

Some doubtful specimens of B. anceps (SAM collections) from the Lower

Maastrichtian of Angola may permit correlation with Europe and the Middle

East. Baculites baculus and Trachybaculites columna permit correlation with the

Western Interior. Exogastrically curved Criobaculites trabeatus is analogous to

the Western Interior Baculites ?sp. of Cobban & Kennedy (1992c: 684,

figs 1.1-1.4, 3.1).

Thus, in contrast to Eubaculites, Baculites has a wide distribution to higher

northern and lower southern latitudes and across the equator; it also occurs in

West Africa, albeit apparently not in very great numbers. The paucity of

Baculites in the Southern Hemisphere in the Maastrichtian compared to the

abundance of Eubaculites and the dominance of Baculites in the Northern

Hemisphere are striking.

COILING

Apart from the ammonitella, the shell in the family Baculitidae is more or

less straight in the majority of taxa. Criocone curvature occurs in two short-

lived lineages, in the endogastric Upper Santonian Boehmoceras and exogastric

Maastrichtian Criobaculites respectively.

As far as is known, the shell in Lechites is always straight and that of

Sciponoceras too, except at the aperture in some (see e.g. Wright 1979: 286, or

in §. cucullatum Collignon (1964, pl. 326 (fig. 1458)). Early (Turonian) forms

of Baculites appear to be straight, but some of the later species may show

various degrees of endogastric curvature of the body chamber. In the Coniacian

and Santonian, some specimens of B. incurvatus and B. capensis have slightly

curved body chambers; Meek’s 1876 ‘group b’ of Baculites included in Spath’s

(1926) genus Euhomaloceras, of which B. incurvatus is the type species. The

majority of B. capensis, however, have straight body chambers and this feature

is of no taxonomic value in these species. Kennedy & Wright (1985: 142)

suggested that these curved B. incurvatus gave rise to crioceratitid-coiled

Boehmoceras in the Upper Santonian; new data from the Gulf Coast (Kennedy

& Cobban 1991b) suggest that B. capensis could be the ancestor. An alternate,

possibly more plausible explanation is that B. incurvatus and B. capensis are

synonyms as suggested earlier (Klinger & Kennedy 1997).

In later Santonian, Campanian and Maastrichtian Baculites, perfectly

straight, as well as slightly curved species occur. A good example of a perfectly

Straight Baculites is B. thomi (in Cobban & Kennedy 1991a, pls 1-2). Amongst

the curved species, curvature may only encompass the early whorls (e.g. in

B. cuneatus (Cobban 1962a: 128), or the whole shell may be curved, more or

less like a scabbard as in B. reduncus Cobban, 1977, or only the body chamber

may be affected as in B. claviformis (Stephenson 1941: 404, pl. 1). Curvature of

the body chamber in some of the Madagascan Campanian species of Baculites is

as variable, as in the ancestral Coniacian-Santonian B. capensis, as shown by

various specimens figured by Collignon (1969). Compare e.g. B. menabensis

(Collignon 1969, pl. 518 (fig. 2036) with straight body chamber, and fig. 2037

32 ANNALS OF THE SOUTH AFRICAN MUSEUM

with curved body chamber)), B. antsirasiraensis (Collignon, 1969, pl. 519

(fig. 2040 with straight body chamber, and fig. 2041 with curved body

chamber)), B. ventroplanus (Collignon 1969, pl. 520 (fig. 2049 with straight

body chamber and fig. 2050 with curved body chamber)). All of these are

probably synonyms of B. tanakae. Slight curvature of the body chamber is also

present in some specimens of B. vanhoepeni and B. sulcatus from the

Campanian of Zululand and Pondoland, respectively. These all indicate that

curvature of the body chamber in these Baculites is very variable and of no

taxonomic significance.

HAPE

ae se WHORL SECTION

ORNAMENT | VENTER | Normal forms Compressed forms

Venter Zon ee.

fastigiate

Crescentic

lateral E. labyrinthicus ——E. binodosus

ribs :

iP APs 7 od

Ae S =

: >E

E. simplex =o

. AG

No Venter 53

lateral tabulate O&

ornament

E. latecarinatus

Bituberculate

lateral

ornament Fonte

Fig. 20. Diagram to illustrate whorl sections and ornament of Eubaculites

species in the Maastrichtian.

In the predominantly Maastrichtian genus Eubaculites, perfectly straight as

well as curved species occur. Distinct curvature of the whole shell in a

scabbard-like manner seems to be characteristic of E. vagina in India (see e.g.

the lectotype in Klinger & Kennedy 1993, fig. 8, or fig. 13) and E. labyrinthicus

in Madagascar (see e.g. Collignon 1971, pl. 645 (fig. 2388) (as Baculites

simplex); Klinger & Kennedy 1993, fig. 6). In contrast, some adult specimens of

E. latecarinatus and E. carinatus in Zululand reach gigantic size with whorl

heights of up to 90 mm and estimated lengths of 2 metres with no sign of

curvature at all.

Criobaculites trabeatus (Morton 1834: 45, pl. 15 (fig. 3)) from the

Maastrichtian Prairie Bluff Chalk of Alabama is known from the holotype only,

and is a body chamber fragment of a criocone exogastrically recoiled baculitid.

Recently, Cobban & Kennedy (1992c, figs 1.1-1.4, 3.1) described another

exogastrically curved heteromorph as Baculites? sp. from the Maastrichtian Fox

THE AMMONITE FAMILY BACULITIDAE 33

Hills Formation of South Dakota, and compared it with Hamites trabeatus; as

mentioned above, it too is probably also a Criobaculites.

WHORL SECTION

Apart from obvious ontogenetic changes, the whorl section seems to be one

of the few reliable features in baculitid systematics, especially in the genus

Baculites.

The whorl section in Lechites and Sciponoceras is remarkably conservative;

it is predominantly circular to elliptical.

The earliest (Turonian) baculitids, B. yokoyamai and B. undulatus retain the

more or less elliptical whorl section of ancestral Sciponoceras (see e.g. Matsu-

moto. & Obata 1963, text-figs 63-86). Later baculitids all have a more or less

ovoid whorl section with the venter narrower than the dorsum. Some Baculites

retain an elliptical whorl section, e.g., some Coniacian—Santonian B. capensis,

Baculites smooth species of Cobban (1962b: 714, pl. 108 (figs 1-4)) in the

Campanian, and B. vertebralis in the Maastrichtian, but these are few. A

circular or near-circular whorl section is rare in Baculites s.s., but occurs in the

Maastrichtian in the short-lived lineages of Fresvillia and Trachybaculites.

Some specimens of B. capensis, especially those from the type locality at the

Mzamba Estuary, have a shallow, longitudinal depression at mid-flank. This

appears to be a unique feature in Baculites and is probably associated with the

unique lateral ornamentation of B. capensis.

Ventral ‘keels’ of variable shape (fastigiate to tabulate) and prominence are

developed in several, apparently unrelated lineages of Baculitidae. The oldest

Baculites with a distinct, siphonal keel is B. kirki, from the Santonian of

California. According to Matsumoto (1959: 143), the keel is small and rounded,

and not clearly developed on internal moulds, where the venter may be

fastigiate. In the latter respect it resembles B. tanakae from the Campanian of

Hokkaido and Madagascar.

Incipient (tabulate) keels have been recorded in B. fuchsi (Summesberger

1979, pl. 1 (figs 2-4)) of the Upper Santonian, in B. bailyi (Woods 1906: 341)

from the Upper Santonian—Lower Campanian, and in B. androtsyensis

Collignon, 1970, B. increscens Collignon, 1970, and B. mamillatus Collignon,

1970, all from the Middle Campanian of Madagascar. None of these species,

however, develop sufficiently prominent keels to link them with true (ventral

tabulate) keeled Eubaculites of the Maastrichtian.

Baculites chicoensis from the Lower Campanian has a more or less distinct

ventral keel but, because of its apparent sutural complexity cannot be linked

with older keeled B. kirki or younger, late Campanian, E. occidentalis. The

latter is regarded by us (Klinger & Kennedy 1993) as the first true Eubaculites

because of its distinct tabulate venter.

Two types of whorl section are present in Eubaculites (Fig. 20). In the first,

and generally considered a diagnostic feature, seen in FE. vagina, E. carinatus

and E. latecarinatus, but not in the type species E. labyrinthicus, the whorl

section is pyriform, with a distinct, tabulate ventral keel, separated from the

convergent flanks by distinct longitudinal furrows, and a tabulate to slightly

concave or convex dorsum.

34 ANNALS OF THE SOUTH AFRICAN MUSEUM

In the second group of Eubaculites, including the type species E. laby-

rinthicus, the whorl section is distinctly trigonal to cuneate, and the venter is

narrowly fastigiate, rather than broadly tabulate and the dorsum is flattened.

Eubaculites simplex is generally more compressed than any of the other

Eubaculites species, but the venter may be fastigiate or tabulate, even at

different ends of the same specimen; the species thus links the two groups

within the genus. Eubaculites binodosus (Noetling, 1897) also belongs to this

group, and differs from E. labyrinthicus by having a single, dorsolateral row of

tubercles.

Apart from E. occidentalis, there are several apparently unrelated Baculites

species in the Upper Campanian-Lower Maastrichtian from different

biogeographic regions that acquire a whorl section similar to that of Eubaculites

as already noted by Lewy (1986: 5).

The uppermost Campanian-basal Maastrichtian baculitid assemblage from

Angola described by Haughton (1925, 1926), Haas (1943) and Howarth (1965)

as B. subanceps, B. anceps and Baculites sp. illustrate the point. Baculites

subanceps normally has a broadly rounded venter; that of B. anceps is narrow,

but in both populations, specimens with tabulate ventral keels occur. Seen in

isolation, these could easily be referred to Eubaculites (see e.g. Klinger &

Kennedy 1993: 218). Eubaculites occidentalis figured by Haggart (1991, pl. 5

(fig. 5)) is indistinguishable from Angolan B. subanceps in the collections of the

S.A. Museum in this respect.

Baculites rioturbioensis Hiinicken, 1965, from the uppermost Campanian or

basal Maastrichtian of Patagonia has a distinct cuneiform whorl section as in the

second group of Eubaculites. Again it is difficult to decide if it is a Baculites or

a Eubaculites. Baculites nibelae and Baculites sp. cf. increscens in the Upper

Campanian of Zululand and Israel respectively have a trigonal whorl section and

lateral ornament very much like that of E. labyrinthicus described from

Madagascar (e.g. Klinger & Kennedy 1993, fig. 36a-d). Baculites bassei

Besairie, 1930, from the uppermost Campanian or basal Maastrichtian of

Madagascar is another baculitid with a Eubaculites-like whorl section, as is

Baculites cf. asperoanceps figured by Collignon (1938, pl. 6 (fig. 7)) from the

Campanian-—Maastrichtian of Andimaka, Madagascar. Baculites lomaensis

Anderson, 1958, from the Lower Maastrichtian of California also has a distinct

trigonal whorl section.

Baculites rugosus Cobban, 1962a, from the Upper Campanian of the US

Western Interior is a very large species. It has a broad, corrugated, slightly

tabulate venter and is superficially very similar to equally large Eubaculites

latecarinatus. Baculites cuneatus Cobban, 1962a, from the uppermost

Campanian has a cuneiform whorl section (as indicated by the name) very much

like that of the second group of Eubaculites. Baculites jenseni Cobban, 1962a,

and B. reesidei Elias, 1933, from the base of the Maastrichtian have incipient

tabulate venters as in the first group of Eubaculites. All these specimens are part

of the endemic Western Interior lineage, and the resemblance is mere

homoeomorphy.

Baculites sp. in Stephenson (1941, pl. 76 (figs 7-8)), from the Upper

Campanian of the Gulf Coast Region of the USA, has a whorl section and

lateral ornament similar to that of E. labyrinthicus in Madagascar.

THE AMMONITE FAMILY BACULITIDAE 35

In the European region, Maastrichtian B. anceps typically has a tear-shaped

whorl section, but also includes specimens with a fastigiate venter or incipient

tabulate keels. These individuals are close to contemporary E. carinatus. Study

of populations, however, shows them to be clearly different.

Boehmoceras krekeleri has a ventral keel, but it is rounded, and either

smooth, serrated or plaited (‘Zopfkiel’).

ig ey,

0 5)

————— el

Fig. 21. Variation in details of the suture line in Baculites vanhoepeni Venzo, 1936.

A. SAS-Z1191. B. SAM-PCZ13146. C. SAM-PCZ7706. D. SAS-Z1923.

SUTURE LINE

All baculitids have a quadrilobate adult suture. The external lobe (E), lateral

(L) and umbilical (U) are large and bifid; the internal lobe (I) is smallest and

trifid. References to six lobes and saddles in Baculitidae by e.g. Meek (1876:

36 ANNALS OF THE SOUTH AFRICAN MUSEUM

0 5)

Fig. 22. Variation in details of the suture line in Baculites capensis Woods, 1906.

A. SAS-H148/3. B. SAS-H13/6. C. SAS-H149/16. D. SAS-L12. E. NMBD1028g.

392) and Reeside (1927a: 3; 1927b: 9) include the total number of saddles and

lobes around the periphery. Unfortunately, studies of the early ontogeny in

Baculites by Brown (1892), Smith (1901), Reeside (1927b), Reyment (1958),

Landman (1982), Bandel et al. (1982), and Landman & Bandel (1985) have not

shown whether the early suture is truly quadrilobate or quinquelobate.. Our

material is not suitable to resolve this problem. The implications of quadri-

versus quinquelobate early sutures in the higher systematics of heteromorph

THE AMMONITE FAMILY BACULITIDAE 37

ammonites are still controversial, see e.g. Mikhailova (1983), Doguzhaeva &

Mikhailova (1982), Wiedmann et al. (1990) and Wright (1996).

The baculitid shell is very simple and, apart from lateral ornament (when

present) and the shape of the whorl section, has very few distinguishing

features. Because of this, details of the adult suture lines have received far more

attention from some authors than they perhaps deserve—especially in the

smooth or feebly ornamented forms of Baculites. Descriptions by e.g. Nowak

(1908), Elias (1933) and others, illustrate the point.

Details of the adult suture that have been emphasized include:

1. Relative complexity, i.e., degree of incision of the suture. (= Index of

sutural complexity ‘ISC’ (Ward 1980: 39)—the length of the suture line divided

by the shortest distance from the umbilicus to the venter.)

2. Mode of incision and shape of the folioles, e.g. phylloid, lytoceratine,

dendritic, etc.

3. Shape of the lobes and saddles, e.g. subquadrate, triangular, etc.

4. Relative length and width of the individual elements.

5. Symmetry of the saddles and lobes.

However, some words of caution are necessary before trying to recognize

sutural patterns in the Baculitidae. Some are obvious, others less so. The

complexity of, and degree of incision of the suture line is directly related to

size, 1.e. whorl diameter. Thus the suture lines of juvenile and adult specimens

will be very different. Suture lines of juvenile specimens are generally not very

diagnostic; they only show the quadrilobate nature of the suture. The effects of

dimorphism also have to be taken into account; the suture line of a large

macroconch may appear slightly more complex than that of a smaller micro-

conch. Thus, when comparing sutures of different species, these should ideally

be at similar ontogenetic stages. Simplification of the last few sutures before

cessation of growth must also be taken into consideration: compare e.g.

B. knorrianus in Kennedy & Summesberger (1987: 33, text-fig. 2). Intraspecific

variation of the adult suture can be extensive: compare, for example,

B. vanhoepeni (Fig. 21), B. capensis (Fig. 22), B. ovatus in Cobban (1974, text-

fig. 4), and E. occidentalis in Matsumoto (1959, text-figs 64-66), etc.

Genus Lechites

The basic baculitid sutural pattern was already established in Albian

Lechites (Fig. 23) with subrectangular lobes E, L and U, and small trifid I. The

umbilical lobe in Lechites may be asymmetrically trifid, and may even be

different on both sides of the same specimen, as shown by Wiedmann & Dieni

(1968: 63, text-fig. 36) and Scholz (1979: 14, text-fig. 5a).

Genus Sciponoceras

In the Cenomanian and Turonian genus Sciponoceras, two basic, but largely

overlapping types of suture can be distinguished (Fig. 24). In the first, the lobes

and saddles are trigonal in outline, the saddles have narrow, pinched stems, and

the stems of the lobes are constricted, with the dorsolateral edges of the saddles

nearly touching. This type of suture is present in S. baculoides (see especially

Matsumoto 1959, text-figs 2-3); S. orientale (see Matsumoto & Obata 1963,

38 ANNALS OF THE SOUTH AFRICAN MUSEUM

text-figs 45-49) and Sciponoceras sp. (Matsumoto & Obata 1963, text-fig. 4). In

the other group, the saddles and lobes are subrectangular, with the stems of the

saddles and lobes nearly as wide at their bases as near the apices. Examples

include S. kossmati (?non Nowak in Matsumoto 1959, text-figs 4-6; Matsumoto

& Obata 1963, text-figs 24-25), S. gracile (see Matsumoto 1959, text-fig. 3)

and §. intermedium (see Matsumoto & Obata 1963, text-fig. 6).

A E

L i U

E L U

Bie

C E z

Fig. 23. Suture lines of Lechites gaudini (Pictet & Campiche, 1861) to show

variation in symmetry of the umbilical lobe (U). A-B. After Scholz (1979,

text-fig. SA-B). C. After Wiedmann & Dieni (1968, text-fig. 36).

Arrow points to subtrifid umbilical lobe (U).

Differences between the two broad types of sutures in Sciponoceras appear

to be in part related to stratigraphic occurrence, and in part to shell shape. The

first group is predominantly Cenomanian and with circular to near-circular

whorl section. The second is predominantly late Cenomanian to Turonian, and

associated with an elliptical whorl section. However, it should be emphasized

that, given the variability of baculitid sutures, there is considerable overlap

between these two types of sutures and one should be cautious in using details

of the suture for specific differentiation (see also discussion by Wright &

Kennedy 1981: 114).

THE AMMONITE FAMILY BACULITIDAE 39

Bie aU

Ml ee

Cc = Cae

E I

D ie U

Edin a Unset

F eS

Fig. 24. Suture lines in Sciponoceras. A-D. S. orientale, with trigonal

saddles and lobes. E-F. S. kossmati, with subrectangular elements.

A-D. After Matsumoto & Obata (1963, text-figs 46-49); E-F. After

Matsumoto & Obata (1963, text-figs 24-25).

Genus Baculites

In the oldest Baculites, B. undulatus and B. yokoyamai, the suture lines

(Fig. 25A-C) are similar to those of the second group of Sciponoceras, with

subrectangular, little-incised saddles and lobes (although some specimens, e.g.

B. yokoyamai in Kennedy & Cobban 1991la, text-fig. 25e (herein Fig. 25C)

deviate slightly from this pattern in being more complex, thus illustrating

variation in the suture). On the basis of sutural similarities, it seems reasonable

to derive Baculites from this group of Sciponoceras.

40 ANNALS OF THE SOUTH AFRICAN MUSEUM

K 0 10

Fig. 25. Suture lines of Turonian and Coniacian Baculites. A. B. undulatus

d’Orbigny, 1850. After Matsumoto & Obata (1963, text-fig. 62).

B. B. yokoyamai Tokunaga & Shimizu, 1926. After Matsumoto & Obata

(1963, text-fig. 87). C. B. yokoyamai. After Kennedy & Cobban (1991a,

fig. 22A). D. B. mariasensis Cobban, 1951. After Kennedy & Cobban

(199 1a, fig. 25E). E. B. codyensis Reeside, 1927a. After Kennedy & Cobban

(1991a, fig. 25F). Scale bars for size.

This simple sutural pattern is retained in the Coniacian in Europe in

B. incurvatus (Fig. 26B-C), in the Indo-Pacific in B. capensis (Fig. 26D-E),

B. yokoyamai and B. bailyi, and in the US Western Interior in B. yokoyamai,

B. mariasensis (Fig. 25D), B. sweetgrassensis and B. codyensis. In contrast, the

sutures of Pseudobaculites, thus far only known from three species,

P. wyomingensis and P. nodosus (Fig. 26A) from the Upper Coniacian, and

P. natosini from the Upper Campanian and Lower Maastrichtian of the US

Western Interior, are extremely complex. The saddles are slender-stemmed and

asymmetrically dendritic. The larger size of Pseudobaculites in the Coniacian

THE AMMONITE FAMILY BACULITIDAE 41

E U I

A |

0 10

E L U I

0 1

E L U

L GT UY. L

D Bd ad Mls 2

E L U I

0 1

Fig. 26. Suture lines of Coniacian Pseudobaculites and Baculites.

A. Pseudobaculites nodosus Cobban, 1952. After Kennedy & Cobban (1991a,

text-fig. 27). B-C. Baculites incurvatus Dujardin, 1837. After

Kennedy (1984, text-fig. 42F). D-E. B. capensis Woods, 1906 (= B. boulei

Collignon, 1931). D. After Collignon, 1931, pl. 11 (fig. 14). E. After

Matsumoto & Obata (1963, text-fig. 93). Scale bars for size.

alone cannot account for this phylogenetically sudden increase in sutural

complexity—specimens of B. capensis of similar size do show a progressive

increase in overall denticulation of the suture line (Fig. 22), but the elements

remain more or less quadrate and unconstricted and quite simple when

compared to that of Pseudobaculites. Sutural complexity similar to that of

Coniacian Pseudobaculites only occurs in much younger Baculites and

Pseudobaculites natosini in the Campanian and Maastrichtian.

The relatively simple Baculites suture is retained by the majority of

baculitids in the Santonian (Fig. 28A-D), including the criocone curved genus

Boehmoceras (Fig. 27). Baculites pseudobaculus (Fig. 28D), a rather poorly

42 ANNALS OF THE SOUTH AFRICAN MUSEUM

known species seems to have slightly asymmetrical lateral saddles, vaguely

reminiscent of Pseudobaculites.

Reeside (1947: 2, pl. 3 (figs 1-6)) recorded three small fragments of an

apparent smooth Baculites from questionable Santonian sediments in Haiti. The

suture seems simplified, but, being small specimens, could be a juvenile feature

only.

The most striking changes in the sutures of Baculites take place in the

Campanian, especially in the Middle and Upper Campanian, and to a lesser

extent in the Maastrichtian. Several broad, overlapping trends can be

recognized, but their interrelationship remains obscure.

L Ch al

E L

Fig. 27. Suture lines of Santonian baculitids. A. Boehmoceras

arculus (Morton, 1834). After Kennedy & Cobban (1991),

fig. 12.3). B. B. arculus. After Kennedy (1987, text-fig. 2).

There is a general trend towards increase of the number of incisions, i.e.

general lengthening of the suture line or increase in ISC. This may be partially

due to the phyletic increase in size in some of the Baculites lineages, but it is

not the only factor. The suture line of some Campanian and Maastrichtian

Baculites is distinctly more complex than that of species of similar size of the

Coniacian and Santonian.

The rate at which increase of complexity of the suture line takes place in

post-Santonian Baculites is variable. In some lineages it is gradual, in others it

is sudden. Also, the degree of complexity does not increase linearly from the

beginning of the Campanian to the end of the Maastrichtian; it may peak and

then simplify before the end of the Maastrichtian.

Changes in sutural complexity are best documented from the Pierre Shale in

Colorado in the US Western Interior (Scott & Cobban 1965). The earliest,

Middle Campanian species, B. obtusus, B. mclearni (Fig. 29A), B. asperiformis

and B. perplexus (Fig. 29B) retain the simple suture of the Coniacian and

Santonian Baculites with subrectangular saddles and lobes. Later, Middle

Campanian species, e.g. B. gregoryensis and B. scotti (Fig. 29C), have a far

more incised, dendritic suture in which the saddles and lobes have slender

THE AMMONITE FAMILY BACULITIDAE 43

E I

E U

B L

0 me

E U I

C j

0 1

| ; ee

0 1

E L I

0 1

Cee!

Fig. 28. Suture lines of Santonian and Campanian Baculites.

A. B. nugssuagensis Birkelund, 1965. After Birkelund (1965, text-fig. 36C).

B. B. thomi Reeside, 1927b. After Cobban & Kennedy (1991a, fig. 2B).

C. B. uedae Matsumoto & Obata, 1963. After Matsumoto & Obata (1963,

fig. 92). D. B. pseudobaculus Matsumoto & Obata, 1963. After Matsumoto

& Obata (1963, fig. 166). E. B. tanakae Matsumoto & Obata, 1963.

After Matsumoto & Obata (1963, fig. 115).

Scale bars for size.

bases. The baculitids of the Upper Campanian and basal Maastrichtian, e.g.

B. compressus, B. cuneatus, B. jenseni and B. eliasi (Fig. 29D) have an even

more incised suture line, in which, according to Scott & Cobban (1965: 2) ‘the

terminal branches of the lateral lobe are constricted at their base—an entirely

44 ANNALS OF THE SOUTH AFRICAN MUSEUM

L Pee

A ;

E L

eS nel

B :

D : U sm erg!

E &®&%

io U png!

Fig. 29. Suture lines of Campanian and Maastrichtian Baculites of the US

Western Interior. A. B. mclearni Landes, 1940. After Cobban (1962), text-

fig. 1h). B. B. perplexus Cobban, 1962b. After Cobban (1962), text-fig. 1c).

C. B. scotti Cobban, 1958. After Cobban (1958, text-fig. ih).

D. B. compressus-eliasi lineage. After (Scott & Cobban 1965).

E. B. baculus-clinolobatus lineage. After Scott & Cobban (1965).

different arrangement from that of older baculites’. The remaining, later

Maastrichtian species of the Western Interior, B. baculus (an immigrant from

the Gulf Coast), and its descendants, B. grandis and B. clinolobatus have, in

contrast, a simple suture (Fig. 29E), comparable to that of the B. obtusus-

B. perplexus group. |

As mentioned above, Pseudobaculites natosini from the Upper Campanian

and Lower Maastrichtian, has a complex suture characteristic of the genus.

Details of the Baculites succession of the Indo-Pacific and European regions

are less precisely known than those of the US Western Interior. Matsumoto

THE AMMONITE FAMILY BACULITIDAE 45

Fig. 30. Suture lines of Campanian and Maastrichtian Baculites and Eubaculites.

A. Eubaculites carinatus (Morton, 1834). After Matsumoto (1959, text-fig. 85a).

B. B. subanceps Haughton, 1925. SAM-6829. Scale bars for size.

In the first identifiable group, the saddles and lobes remain more or less

quadrate or rectangular, and their bases unconstricted and open, as in the early,

Turonian, Coniacian and Santonian Baculites. The width of the individual

elements may vary, but their more-or-less angular outline is characteristic. In

addition, some of the folioles in the lobes may become phylloid. This group

typically includes B. subanceps (Fig. 30B), B. inornatus and E. occidentalis

(Fig. 31D-E) in the Upper Campanian, and B. anceps (Fig. 31A),

B. palestinensis, B. huenickeni, B. teicherti and the whole genus Eubaculites

(Fig. 30A) in the Maastrichtian. This group corresponds more or less to Ward’s

(1978: 1145) ‘phylloid’ group. However, because of the variable development

of phylloid folioles, Ward’s terminology is not recommended, and we merely

refer to it as group #1. The origins of this group are not fully established.

Matsumoto & Obata (1963: 102, fig. 218) and Ward (1978: 1148, text-fig. 4)

suggested B. bailyi as the ancestor.

Some of the other Campanian and Maastrichtian European baculitids, e.g.

B. leopoliensis, B. vertebralis (Fig. 31B) and B. knorrianus, may be referable

to this group, but we remain uncertain. Available suture lines of these species

show a more complex pattern than in the typical representatives of this group.

A second identifiable lineage includes mainly ornate, but also smooth forms.

It originated in B. capensis, and includes B. sulcatus, B. duharti,

B. vanhoepeni, and ends in B. nibelae in Zululand (Fig. 32). All the baculitids

from the Lower and Middle Campanian of Madagascar described by Collignon

(1969, 1970) (see Klinger & Kennedy 1997, figs 82-89) also belong to this

lineage, as does B. tanakae (Fig. 28E) from the Campanian of Hokkaido (which

46 ANNALS OF THE SOUTH AFRICAN MUSEUM

is probably the prior name for all the Lower Campanian baculites of Madagas-

car). In this lineage, there is a progressive increase in the degree of incision of

the suture line, accompanied by a narrowing of the bases of the saddles and the

lobes, resulting in a suture line comparable to that of the group of B. gregory-

ensis of the US Western Interior. According to Matsumoto & Obata (1963: 102,

text-fig. 218) this lineage continues into the Maastrichtian with B. lomaensis.

Some of the Campanian baculitids from France described by Kennedy (1986b,

text-fig. 8f-h) have a similar suture and probably also belong to this group, as

may specimens from Israel referred to as Baculites sp. cf. increscens.

L i

C 5

im)

L I

0 5

it jk

7 U

0 5

Fig. 31. Suture lines of Campanian and Maastrichtian Baculites

and Eubaculites. A. B. anceps Lamarck, 1822. After Kennedy (1986c, text-

fig. lld). B. B. vertebralis Lamarck, 1801. After Kennedy (1986c, text-

fig. 11A). C. B. lomaensis Anderson, 1958. After Matsumoto (1959, text-

fig. 35). D-E. Eubaculites occidentalis (Meek, 1862). After Matsumoto

(1959, text-figs 64, 65a). Scale bars for size.

THE AMMONITE FAMILY BACULITIDAE 47

0 5

Fig. 32. Suture lines of the lineage B. yokoyamai (E, D), B. capensis (C),

B. sulcatus (B), and B. vanhoepeni (A) in Zululand to illustrate increasing

sutural complexity. Scale bar for size.

The third identifiable lineage consists of smooth Baculites with extremely

complex sutures. This includes B. hochstetteri Liebus, 1902 (Fig. 33A-B),

B. rex, B. chicoensis (Fig. 34A-D) and B. rectus (Fig. 33C). This is in part

Ward’s (1978) ‘lytoceratine’ group. Unfortunately, the relationship of these

species is far from clear. All have an extremely dendritic suture line with

narrow-stemmed saddles and lobes. In B. rex and B. chicoensis the incisions of

the lobes and saddles appear to be predominantly pointed, i.e. ‘lytoceratine’ in

the sense of Ward (1978). In contrast, the sutures of B. hochstetteri and

B. rectus are more rounded ‘phylloid’. This would suggest that this is not a

homogeneous group, but rather a grouping of convergent taxa, but we cannot be

sure.

The origins of this group are uncertain. Matsumoto & Obata (1963: 102,

text-fig. 218) suggested derivation from B. bailyi and B. uedae (which we

48 ANNALS OF THE SOUTH AFRICAN MUSEUM

regard as synonyms). Whatever the origins of this group, the increase in

complexity of the suture line appears to be sudden: punctuate. In South Africa,

a similar sudden jump in sutural complexity occurs in the series presumably

originating in B. yokoyamai, through B. bailyi and ending in Baculites cf. rectus

(Fig. 35).

E L

U L

| iis Sea

OI Uae ne

1 Ry’ ane

Fig. 33. Complex suture lines of smooth Campanian-?Maastrichtian

Baculites. A-B. B. hochstetteri Liebus, 1902. After Liebus (1902,

text-fig. 2). C. B. rectus Marshall, 1926. After Henderson (1970,

text-fig. 6a).

The sutures of this third lineage, especially those of B. rectus are very

similar to those of the group of B. compressus-B. eliasi of the US Western

Interior. Furthermore, they seem to be of more or less similar age. This

suggests a parallel development of sutures in the Indo-Pacific and US Western

Interior towards the end of the Campanian, and persisting into part of the

Maastrichtian. A similar degree of sutural complexity is also achieved in

B. ovatus (Fig. 36) from the Campanian the Gulf Coast Atlantic Seaboard of the

USA.

During the Maastrichtian several small, short-lived baculitid groups

appeared in which sutural simplification or modification has taken place.

The sutures of Fresvillia are not fully known, but illustrations of

F. constricta (Fig. 37B) and F. teres show subtriangular, but not simplified

saddles and lobes, quite unlike those of contemporary Baculites or Eubaculites.

The specimen described and figured by Pervinquiére (1907: 95, pl. 4

(fig. 12a-b), text-fig. 25) (Fig. 38B) as Baculites indét. from the Upper

Senonian of Pont du Fahs, Tunisia, is minute, with a circular whorl section,

smooth surface and a constriction. The suture shows distinct triangular saddles

THE AMMONITE FAMILY BACULITIDAE 49

E If

L U

iG U

E ;

Cc Eas

0 5

E naa

L ae U

Fig. 34. Complex suture lines of Campanian and Maastrichtian Baculites.

A-B. B. chicoensis Trask, 1856. After Matsumoto (1959, text-figs 62a, 63a).

C-D. B. rex Anderson, 1958. After Matsumoto (1959, text-figs 45a, 46a).

Scale bars for size.

and lobes, which suggest polyptychoceratid rather than baculitid affinities. The

suture line and constriction are homoeomorphic with some Cenomanian

Sciponoceras. Baculites paradoxus (Fig. 38A) is another minute, doubtful

baculitid from the Maastrichtian of Draa et Tbaga, Tunisia, described by

Pervinquiére (1907: 94, pl. 4 (figs 10-11), text-fig. 24). According to

Pervinquiére (1907: 94), this species is unique in allegedly having five saddles

and lobes instead of six (around the whole periphery), a feature also noted by

Reeside (1927b: 9). An alternative interpretation could be that the internal lobe

(I) is merely very small, and as large as the median incisions of the saddles E/L

and L/U. The triangular shape of the saddles and lobes is similar to that of

Baculites indet., from which it differs by the lack of constrictions. Both are

50 ANNALS OF THE SOUTH AFRICAN MUSEUM

probably polyptychoceratids rather than baculitids as suggested earlier (Klinger

& Kennedy 1997).

Trachybaculites vicentei Stinnesbeck, 1986 (p. 203, pl. 9 (fig. 4), pl. 10

(figs 3-6), text-figs 23a—c) (Fig. 37A) from the Maastrichtian of Quiriquina

Island, Chile, has a pseudoceratitic suture line.

In the US Western Interior, the last baculites are B. larsoni (Fig. 37E),

Baculites sp., Trachybaculites columna (Fig. 37D) and an endogastrically-coiled

baculitid (Fig. 37C), probably referable to Criobaculites, all with simplified

sutures.

L U :

A |

E ik U I |

L U

(¢

L U

0 5

(ee eae]

Fig. 35. D-A. Suture lines of Coniacian to Campanian smooth

B. yokoyamai-Baculites cf. rectus lineage in South Africa to illustrate

increasing complexity. Scale bar for size.

THE AMMONITE FAMILY BACULITIDAE 51

Fig. 36. Complex suture line of Campanian-Maastrichtian B. ovatus Say,

1821. After Cobban (1974, text-fig. 4). x 2.8.

ORNAMENTATION

The following elements of ornamentation can be identified in Baculitidae:

1. Growth lines. In near-smooth, non-ornate forms there are growth lines and

striae only. These are often only visible on the shell and absent on internal

moulds. Some of the striae may be stronger (lirae) than others.

2. Lateral tubercles—mostly only one row, in very rare cases two rows

connected by ribs of variable strength. The shape, size and spacing of these

tubercles varies widely, and they often grade into:

3. Lateral ribs. These can vary from mere transversely elongated tubercles, to

distinct, circumperipheral ribs; either single or often bifurcating or with

intercalatories on the venter.

4. Ventral corrugations. These are also very variably developed—in some they

are mere undulations on the venter, and only present in shelly preservation—in

others distinct undulations; also often associated with ventral bifurcating or

intercalated ribs.

5. Constrictions. These may appear regularly, and be of taxonomic importance,

or random, sometimes as body-chamber modifications only, and of little

significance.

6. Ventral tubercles—these are extremely rare.

Ornamentation in the early representatives of the family, Lechites and

Sciponoceras is very conservative. Lechites has simple, albeit variably spaced,

circumperipheral ribbing and occasional constrictions. An aberrant offshoot in

52 ANNALS OF THE SOUTH AFRICAN MUSEUM

the Upper Albian, L. (Tuberolechites) has minute ventral tubercles on the ribs.

Cenomanian and Turonian Sciponoceras either retain the simple ribbing of

Lechites or become smooth, and constrictions become a permanent feature of

ornamentation.

In contrast, ornamentation in later, Turonian to Maastrichtian Baculites is

notoriously variable. The great number of apparently endemic Baculites species

recognized, as well as gross misidentifications all indicate how variable

ornament can be and the extent of homoeomorphy.

Most problematic are the smooth (non-ornate species). Unless they have a

very characteristic whorl section, as in e.g. some B. anceps, or a distinct sutural

E E

18 UT

0 10

\ L a

0 10

Fig. 37. Suture lines of diverse Maastrichtian baculitids.

A. Trachybaculites vicentei (Stinnesbeck, 1986). After Stinnesbeck

(1986, text-fig. 23a, c). B. Fresvillia constricta Kennedy, 1986. After

Kennedy (1986a, text-fig. 10a). C. Criobaculites sp.

D. Trachybaculites columna (Morton, 1834). E. Baculites larsoni

Cobban & Kennedy, 1991b. C-E. After Cobban & Kennedy (19915,

text-fig. 3.1-3.3). Scale bar for size.

THE AMMONITE FAMILY BACULITIDAE 53

Baculites paradoxus PErviNQuizERE 1907

Baculites indet.

Fig. 38. Sutures of Maastrichtian Baculites paradoxus

Pervinquiére, 1907, and Baculites indet. After Pervinquiére

(1907, text-figs 24, 25).

pattern, e.g. B. rectus, identifying unlocalized smooth Baculites is virtually

impossible. In addition, some predominantly smooth species, e.g. B. yokoyamai

or B. bailyi (see Klinger & Kennedy 1997) include rare forms with distinct

lateral ornament.

Ornamented forms present even more problems. Variation in Eubaculites

vagina, recently comprehensively figured by Klinger 1976, Kennedy & Hen-

derson (1992) and Klinger & Kennedy (1993) illustrates the point—lateral

ornament varies from distinctly bituberculate, through ribbed to smooth

(Figs 39-41). Our material of B. capensis (Klinger & Kennedy 1997) shows a

similar wide range of variation. Apart from simple intraspecific variation,

similar forms of ornament appear at different times (heterochronous

homoeomorphy), or in apparent unrelated baculitid lineages of more-or-less the

Same age (synchronous homoeomorphy). Thus the crescentic dorsolateral

tubercles of B. brevicosta in the Coniacian are virtually identical to those of

Maastrichtian B. lomaensis. Some Lower Campanian B. sulcatus with strong

lateral and bifurcating ventral ribbing are indistinguishable from similar Upper

Campanian B. leopoliensis. Specimens with identical lateral tubercles occur in

B. capensis and B. codyensis, although populations differ markedly. Identical

ornament consisting of distinct auricular lateral nodes occurs in the lineages of

B. vanhoepeni and B. sulcatus from the Middle and Lower Campanian of

Zululand and Pondoland respectively, B. asperiformis Meek, 1876, in the

Middle Campanian of the Western Interior, B. taylorensis in the Middle

Campanian of the Gulf Coast and, in Upper Santonian, Boehmoceras arculus.

Unlocalized specimens of these species (save the last) would be impossible to

separate.

A broad outline of the ornamentation of Turonian to Maastrichtian

baculitids, especially of the genera Baculites and Eubaculites is given in the

54 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 39. Eubaculites vagina (Forbes, 1846). Variation in lateral ornament. Specimen

with typical bituberculate lateral ornament. BMNH C77593 from the Valudavur

Formation of southern India. x 1.

sketches in the stratigraphic and geographic distribution charts in Figures 3 to

19. It is impossible to give a detailed description of the ornamentation of each

individual species, and only broad outlines are provided.

THE AMMONITE FAMILY BACULITIDAE

"I x

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jesaye] Oy]

qld YIM suouIoeds

JUSUILUIO [eIO}L] UT UOTIETIeA *(QPRT

‘saqio

‘TIVIISO HNN “O-V ‘JuoweUIO

sannsvqng “Op “3tJ

d) Duispa

56 ANNALS OF THE SOUTH AFRICAN MUSEUM

The earliest Baculites, B. yokoyamai and B. undulatus are non-tuberculate,

but the shell bears fine ribbing; that of B. undulatus is coarser than that of

B. yokoyamai. Baculites calamus, poorly known from the Middle Turonian of

the US Western Interior only, has ribbing more prominent than that of

B. undulatus. On the body chamber of both B. yokoyamai and B. undulatus,

coarse, fold-like ribs may appear similar to those of some Sciponoceras

bohemicum (Fig. 181A-F):

The first dorsolateral ornament in Baculites probably arose through

thickening of, and/or bundling of sheaves of lirae on the dorsal flanks, giving

rise to feebly crescentic or oblique tubercles (Klinger & Kennedy 1997, fig. 55).

Tubercles first appear in the uppermost Turonian or basal Coniacian of Angola

in Baculites sp. as weak dorsolaterally situated crescentic nodes. We assume

that the Baculites codyensis, B. capensis and B. brevicosta-B. incurvatus

lineages arose independently during the Coniacian from B. yokoyamai in the US

Western Interior, the Indo-Pacific Region and Europe, respectively, during the

Coniacian, albeit at slightly different times.

The baculitid succession in the US Western Interior starts with smooth

B. yokoyamai in the Lower Turonian. Ornamented B. sweetgrassensis and

Cc D

Fig. 41. Eubaculites vagina (Forbes, 1846). Variation in lateral ornament. Specimens

with lateral striations only. A-B. BMNHC51142. C-D. BMNHCS51145.

E. BMNHC73569. All from the Valudavur Formation of southern India. x 1.

THE AMMONITE FAMILY BACULITIDAE 57

A B C

Fig. 42. A. Baculites sp. from the Baculites compressus Zone, Pennington County,

South Dakota. BHI 4360. B-C. B. cuneatus from Elk Creek, Meade County, South

Dakota. BHI 4366. Two baculite specimens showing constrictions that appear to

indicate old apertures. Both x 1.

B. codyensis appear in the Middle Coniacian. Ornament consists predominantly

of lateral ribbing, although some variants of B. codyensis have distinct lateral

nodes rather than ribs—the B. ‘asper’ of previous authors. Lateral ribbing

remains the dominant mode of ornament in the baculitid succession of the US

Western Interior; in the Santonian, distinct ventral corrugations are initiated in

58 ANNALS OF THE SOUTH AFRICAN MUSEUM

B. thomi. Pseudobaculites nodosus is an exception, in having lateral ornament,

consisting in some specimens of two nodes connected by a rib—a mode of

ornament reminiscent of Eubaculites vagina. Pseudobaculites wyomingensis has

weak flank ornament and ventral undulations. Details of the ornament of the US

Western Interior baculites are admirably illustrated in Scott & Cobban (1965)

and Gill & Cobban (1973). The lineage from B. codyensis to B. eliasi

underwent progressive, sometimes reversed change. Ribbed forms gave rise to

smooth ones and vice versa in this lineage, which ends in the Lower

Maastrichtian with B. eliasi, a smooth species. Later forms arose from

B. baculus, an immigrant from the Gulf Coast.

The baculitid succession in the Indo-Pacific region most probably also has

its origins in B. yokoyamai, as suggested by Matsumoto & Obata (1963: 102,

text-fig. 218) and by the Baculites faunas of Zululand and Madagascar. The first

ornamented forms occur in the Middle Coniacian of Zululand as Baculites

capensis. However, unlike the baculite faunas of the US Western Interior, the

dominant form of lateral ornament of the Coniacian, Santonian and Lower

Campanian B. capensis lineage consists of nodes, rather than ribs. Details of the

ornament are shown in Klinger & Kennedy (1997) and only the salient points

need to be discussed here. Lateral ornament in B. capensis varies from absent,

through dorsolateral crescentic ‘schencki’, rounded weak dorsolateral ‘boulei’,

prominent lateral conical or longitudinally elongated, typical capensis ornament,

to elongated and pinched, oblique ‘wmsinenensis’ type of ornament. The typical

capensis type of ornament only occurs up to the Middle Santonian in Zululand

and Pondoland, but continues into the Lower Campanian in Madagascar and

Hokkaido as B. menabensis and B. tanakae, with essentially ‘umsinenensis’

oblique lateral tubercles.

Strongly ribbed forms occur in the Lower Campanian of Pondoland as

B. sulcatus, replacing, but probably originating in B. capensis. The Middle

Campanian baculites of Zululand and Madagascar, represented by B. increscens

and B. vanhoepeni have prominent lateral rib-like ornament, closely resembling

the Middle Campanian B. obtusus-B. asperiformis lineage of the US Western

Interior; a remarkable example of convergence, as mentioned above. Ventral

corrugation does occur in some specimens of B. vanhoepeni, but, in contrast to

the baculitids of the US Western Interior, it is not a consistent feature and of no

taxonomic value. A smooth baculite, B. duharti occurs more or less at the same

level as the first occurrence of B. vanhoepeni. The whorl section clearly

indicates that it belongs to the predominantly ornamented lineage of

B. capensis—B. vanhoepeni, and is not a derivative of Coniacian-Lower

Campanian smooth B. bailyi. The B. capensis—B. vanhoepeni lineage ended in

the Upper Campanian in Zululand with B. nibelae Klinger & Kennedy, 1997,

and in the Upper Campanian and/or Lower Maastrichtian of Madagascar with

B. bassei Besairie, 1930, with trigonal whorl section and lateral ribs. It possibly

continued as B. lomaensis in the Maastrichtian of California.

As mentioned above, Baculites appear rare in Europe and are poorly known

from the Coniacian to Campanian. However, Coniacian B. brevicosta and

Coniacian-Santonian B. incurvatus are similar to the B. capensis lineage of the

Indo-Pacific, if not conspecific. Some of the German Campanian forms figured

by Muller & Wolleman (e.g. 1906, pl. 2 (fig. 2)) as B. incurvatus have oblique

THE AMMONITE FAMILY BACULITIDAE 59

lateral tubercles as in B. menabensis; Baculites sp. 1 (pars) of 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) resembles B. vanhoepeni. Baculites leopoliensis, an Upper

Campanian (to Lower Maastrichtian?) European baculite has distinct lateral

ribbing that splits over the venter.

Apart from Eubaculites, the remaining Upper Campanian and Maastrichtian

baculites of the different geographic regions are difficult to relate to the above

lineages. Ornament in Eubaculites is variable, but, as shown in Figure 20

includes consistently smooth, laterally ribbed and bituberculate forms. The

ventral keel may be corrugated or smooth in the same species (e.g. E. vagina)

as in some Baculites, and this feature again is of no taxonomic importance.

Ornament in the remaining baculitids ranges from smooth to ribbed, and no

distinct tuberculate forms occur. The circumperipheral ribbing of

Trachybaculites columna and T.? furcillatus is conspicuous. Baculites kegeli

Oliveira, 1957, and B. lechitides Brunnschweiler, 1966, probably also belong to

the genus Trachybaculites. The ornament of B. anceps was discussed

extensively by Howarth (1965); it includes smooth and laterally ribbed forms,

some with, and others without constrictions—the only species of Baculites in

which constricted individuals occur with some frequency. |

ABSOLUTE SIZE, DEGREE OF TAPER, DIMORPHISM

AND SHAPE OF THE APERTURE

Before discussing these four closely related aspects, some words of caution.

Some species have mature adult apertures that differ in no significant respects

from the transient aperture morphology indicated by the growth lines. In others

there are distinct modifications that mark the cessation of growth at maturity.

But formation of an aperture apparently does not always indicate maturity:

Birkelund (1965: 62, pl. 9 (fig. 3a—c)) described and figured a specimen of

B. obtusus from West Greenland, which is part of a phragmocone, but has a

distinct flare, indicating that ‘growth does not necessarily stop when a flared

aperture has been formed’ (Birkelund 1965: 62). Steinmann (1895: 90, pl. 6

(fig. 4b)) illustrated a specimen of Eubaculites carinatus (as B. vagina) with an

old aperture (stehengebliebenen Mundrandern). However, judging by the

scarcity of phragmocones with parts of old apertures preserved, this appears to

be a rather rare phenomenon in Baculitidae. We figure here (Fig. 42) US

Western Interior Baculites with what may be transient apertures marked by

constrictions.

ABSOLUTE SIZE

Disregarding the possible effects of dimorphism and ecophenotypic vari-

ation, the family Baculitidae shows a progressive increase in overall size,

reaching a maximum in the late Campanian to early Maastrichtian (Fig. 43). In

the Maastrichtian, several dwarfed stocks appear, e.g. Fresvillia, Trachy-

baculites, Baculites larsoni, etc. Pseudobaculites, from the Upper Coniacian of

the US Western Interior are small, with a phragmocone whorl height of up to

ANNALS OF THE SOUTH AFRICAN MUSEUM

No @\

\ \

i O Lechites

\/_ Sciponoceras @

ae ©. Baculites

©} Eubaculites

] oo

70 @

MAXIMUM RECORDED WHORL HEIGHT

50 @® 0@ lo

& @)

4Q @ }

® @

2 @ @

ae OL” ©

30 @ @ @

o 8

20 @

@® © =e Oe

@ @ ® @

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L M U LM U L M

: SAN. CAMP. MAAST.

TIME SCALE NOT TO SCALE

Fig. 43. Diagram illustrating progressive increase in overall size in Baculitidae.

Time axis not to scale. Whorl height in millimetres. Data from various sources.

Genus Lechites. 1. L. comanchensis. 2. L. gaudini. 3. L. campichei.

4. L. vraconensis. 5. L. moreti. 6. L. antanimangaensis. Genus Sciponoceras.

1. S. roto. 2. S. baculoide. 3. S. kossmati. 4. S. orientale. 5. S. santacrucense.

6. S. intermedium. 7. S. gracile. Genus Baculites. 1. B. albertensis.

2. B. ambatryensis. 3. B. anceps. 4. B. subanceps. 5. B. androtsyensis.

6. B. ankilizatensis. 7. B. antsirasiraensis. 8. B. aquilaensis. 9. B. argentinicus.

11. B. asperiformis. 12. B. baculus. 13. B. bailyi. 14. B. bassei.

15. B. “besairier’. 16. B.- borealis. 17. B. boulet. 18. B. calamuse

19. B. capensis. 20. B. chicoensis. 21. B. claviformis. 22. B. coagmentatus.

23. B.codyensis. 24. B. robinsoni. 25. B. corrugatus. 26. B. reesidei.

27. B. compressus. 28. B. crickmayi. 29. B. cuneatus. 30. B. delvallei.

31. B. duharti. 32. B. eliasi. 33. B. falcatus. 34. B. fuchsi. 35. B. gilberti.

36. B. grandis. 37. B. gregoryensis. 38. B. haresi. 39. B. huenickeni.

40. B. increscens. 41. B. incurvatus. 42. B. inornatus. (continued on facing page)

THE AMMONITE FAMILY BACULITIDAE 61

46 mm; the Upper Campanian-Lower Maastrichtian P. natosini is a giant with a

whorl height of up to 240 mm on the body chamber.

True giants seem to occur in separate lineages of the family, but not

necessarily at the ends of these. In Sciponoceras, S. gracile is a true giant in

comparison with other contemporary Cenomanian, or younger, Turonian

members of the genus, or Baculites.

In Baculites, early Turonian and Coniacian species are all relatively small in

comparison with the majority of later, especially Campanian, species.

In the US Western Interior, a lineage can be followed from B. yokoyamai in

the Lower Turonian to Lower Coniacian, through B. codyensis in the Middle

Coniacian to Middle Santonian, then through a series of species in the Cam-

panian, starting with B. obtusus, reaching maximum size in B. rugosus in the

Upper Campanian, with whorl heights of 110 mm. Thereafter, there is a

decrease in size, ending with B. eliasi in the Lower Maastrichtian with whorl

heights of up to 57 mm.

In South Africa, a similar progressive increase in size can be traced in two

separate lineages. The first is predominantly nodose, starting with the group of

B. capensis in the Middle Coniacian to Middle? Santonian, probably through

B. sulcatus and B. increscens in the Lower Campanian, reaching maximum size

in the Middle Campanian in B. vanhoepeni and B. duharti, and followed by a

decrease in size in B. nibelae. The other, smooth lineage started with B. bailyi

in the Middle Coniacian, possibly ending in B. aff. rectus in the Campanian.

Both lineages probably had their origins in B. yokoyamai.

In the Campanian-Maastrichtian of the Gulf Coast region of the USA,

B. claviformis may reach gigantic proportions (Stephenson 1941, pl. 1) with

lengths in excess of 170 cm. (Neal Larson informed us that Stephenson’s

figured specimen, is, in fact, composite, consisting of eight to ten individuals.

The largest baculite they have seen is a B. obtusus with an aperture, and is

113.5 cm long, missing 20 cm of phragmocone).

Baculites rex, from the Maastrichtian of California is another giant with

whorl heights greater than 60 mm. :

Even larger are giant Eubaculites latecarinatus and E. carinatus with

estimated total lengths of two metres, known from the Maastrichtian of

Zululand and Neuquén Basin of Argentina, respectively.

Fig. 43 (continued). 43. B. jenseni. 44. B. kirki. 45. B. knorrianus. 46. B. kotanii.

47. B. latelobatus. 48. B. lechitides. 49. B. leopoliensis. 50. B. libyensis.

51. B. mclearni. 52. B. lomaensis. 53. B. malagasyensis.. 54. B. mamillatus.

55. B. mariasensis. 56. B. meeki. 57. B. menabensis. 58. B. natosini. 59. B. nugs-

suaquensis. 60. B. obtusus. 61. B. occidentalis. 62. B. ovatus. 63. B. perplexus.

64. B. princeps. 65. B. pseudobaculus. 70. B. pseudovatus. 71. B. rectangulatus.

72. B. rectus. 73. B. reduncus. 74. B. reesidei. 75. B. regina. 76. B. rex.

77. B. rioturbioensis. 78. B. roedereri. 79. B. rugosus. 80. B. schencki. 81. B. Scotti.

82. B. sparsinodosus. 83. B. subcircularis. 84. B. sweetgrassensis. 85. B. subtilis.

86. B. tanakae. 87. B. teicherti. 88. B. thomi. 89. B. trifidilobatus. 90. B. uedae.

91. B. undatus. 92. B. undulatus. 93. B. ventroplanus. 94. B. vertebralis.

95. B. yokoyamai. Genus Eubaculites. 1. E. vagina. 2. E. latecarinatus. 3. E. carinatus.

4. E. labyrinthicus. 5. E. simplex. Pseudobaculites natosini (Robinson, 1945) with

maximum whorl height of 260 mm (see Cobban & Kennedy 1994a) is not included in the

diagram.

62 ANNALS OF THE SOUTH AFRICAN MUSEUM

SIZE DISTRIBUTION IN EUBACULITES CARINATUS

ZULULAND (Klinger & Kennedy 1993: 219-220)

fa ARGENTINA (Riccardi 1974: 394)

7 || CHILE (Hiinicken & Covacevich 1975: 151)

Number of specimens

Maximum whorl height

Fig. 44. Histogram showing distribution in size of Eubaculites carinatus (Morton, 1834)

from Zululand (South Africa), Neuquén (Argentina), and Quiriquina (Chile).

Whorl height in mm.

Matsumoto (1959: 141) mentioned size differences in B. rex from different

localities in California, but could not decide whether these were due to simple

‘variation in size or . . . ecological difference between immature and mature

shells’ (i.e. ontogenetic separation). Matsumoto & Obata (1963: 67) mentioned

a similar difference in size between B. chicoensis from California and Hokkaido

respectively; those from the latter region are distinctly smaller than those from

California.

We (Klinger & Kennedy 1993: 235) noticed a similar difference in size in

Eubaculites carinatus. This is one of the few baculitid species with a near

world-wide distribution (see p. 6), and it is possible to compare the size distri-

bution from different geographic localities (Fig. 44). Giants with whorl heights

of 80 to 90 mm, and estimated lengths of 2 metres are known from the Neuquén

Basin (Argentina) and Zululand. In contrast, all the specimens recorded from

North America and Europe are much smaller. This may, however, be due to the

fact that the species is less common in these areas than in Argentina and in

Zululand, to their occurrence in facies where small fossils predominate (e.g.

phosphatic beds in the Prairie Bluff Chalk in the U.S. Gulf Coast Region, and,

in part due to collecting failure. Eubaculites carinatus is, however, very

common in southern Chile (see especially Htinicken & Covacevich 1975;

Klinger & Kennedy 1993, fig. 37). None of these Chilean specimens is as large

as the Neuquén and Zululand giants, and the whole known population seems to

consist of individuals which are smaller than those from Zululand and

Argentina.

A similar situation may occur in B. capensis. This species is well known

from Zululand, Pondoland, California and Hokkaido, but is apparently rare in

Madagascar. All the recorded specimens from California and Hokkaido are

small compared to the average size recorded from Zululand (Fig. 45).

THE AMMONITE FAMILY BACULITIDAE 63

Furthermore, most specimens from Pondoland appear to be smaller than those

from Zululand.

For the present, we are unable to interpret these data. We can only note that

there appear to be distinct differences in maximum size within the same

baculitid species from different geographic areas and/or depositional

environments. We suspect it may somehow be connected with different nutrient

supplies.

(eo)

Size distribution in Baculites capensis

[_]1. Zululand

[fj 2. Pondoland

[:]3. California

40 ZA 4. Hokkaido

NUMBER OF SPECIMENS

MAXIMUM WHORL HEIGHT IN MM

Fig. 45. Histogram showing distribution in size of Baculites capensis Woods,

1906, from Zululand and Pondoland (South Africa), California (USA), and

Hokkaido (Japan).

DEGREE OF TAPER

The taper index (Matsumoto & Obata 1963: 4) is the difference in whorl

height at two successive points on a shell divided by the distance measured

between them x 100. A taper index of less than 5 is considered low, between 5

and 10 moderate, and above 10 high, according to these authors. Converted into

the apical angle of the baculitid shell, these three categories of taper index

correspond to less than 3 degrees, 3-6 degrees and more than 6 degrees.

As Birkelund (1965: 44) has already noted, the taper index changes between

different ontogenetic stages; there is a considerable difference between juvenile

and adult stages, as well as individual variation. In addition, the degree of taper

differs between large and small populations of the same species as well as

between macro- and microconchs. Unless the taper index is very high, as in e.g.

64 ANNALS OF THE SOUTH AFRICAN MUSEUM

Pseudobaculites or very low as in e.g. B. thomi, we have found this feature to

be of very limited taxonomic significance. In addition, because of the generally

fragmentary nature of baculitids in South Africa, it is very difficult to determine

satisfactorily in the material we have studied.

DIMORPHISM AND SHAPE OF THE APERTURE

Dimorphism in the Baculitidae, especially in the genus Baculites has only

recently been demonstrated and/or suggested for some taxa, but as a whole,

remains more or less unexplored. In the major descriptions of the Baculitidae

from the US Western Interior (Reeside 1927a, 1927b; Cobban 1951 onwards),

California (Matsumoto 1959), Hokkaido (Matsumoto & Obata 1963), Honshu

(Obata & Matsumoto 1963), and Greenland (Birkelund 1965), dimorphism is

hardly mentioned—but in due fairness to these authors, dimorphism per se in

Cretaceous ammonites has only recently been recognized and investigated.

Matsumoto & Obata (1963: 52) did, however, note disparate sizes of adult

B. tanakae as did Obata & Matsumoto (1963: 90) in B. regina.

According to Cooper & Kennedy (1977), the earliest representative of the

family, the genus Lechites is distinctly dimorphic, as demonstrated by Lechites

gaudini from the Upper Albian Cambridge Greensand of England. Dimorphism

is manifested by differences in size, as well as shape of the aperture. A size-

frequency histogram (Cooper & Kennedy 1977, fig. 3) shows two size

groupings. Macro- and microconchs appear to have different apertures:

microconchs have an oblique down-turned, collared aperture, whereas

macroconchs have an expanded, trumpet-shaped aperture (Fig. 46). Recent

observations by Kennedy (1996 in Gale et al. 1996: 577-578) show that in

microconchs of L. (L.) gaudini the apterture is oblique to the long axis of the

shell, and preceded by weakening ribs. Macroconch apertures appear to have a

final sector of delicate, wiry ribs.

According to Wright & Kennedy (1981: 112) dimorphism is wide-spread in

Sciponoceras. In contrast to Lechites, numerous specimens of Sciponoceras with

apertures are known.

Fig. 46. Apertures of microconch and macro-

conch of Lechites gaudini (Pictet & Campiche,

1861). After Cooper & Kennedy (1977, fig. 7).

THE AMMONITE FAMILY BACULITIDAE 65

These show that some apertures are simple, others have a long ventral

rostrum, broad folds and a high collar, others have lappets, and some are

recurved (Figs 47-50). It appears that some of these apertural features are of

sexual as well as specific significance, but there is still disagreement on details.

Two examples illustrate the point:

1. Sciponoceras gracile (Shumard, 1860) (Fig. 46). Kennedy (1988: 108)

described the species as highly dimorphic. Micrcconchs develop a hooded

aperture at whorl heights of between 7.3 and 11.7 mm, ribs strengthen on the

venter towards the aperture, a dorsal constriction develops and the aperture

itself may be distinctly flared (see e.g. Cobban & Scott 1972, pl. 17

(figs 23-24)). Macroconchs are much larger, up to four times the size of

microconchs with whorl heights of up to 41.4 mm; complete apertures of

macroconchs have not yet been found. A specimen figured by Kennedy (1988,

pl. 20 (figs 17-20)) shows part of a macroconch aperture without a hood.

2. Sciponoceras baculoides (Mantell, 1822). Marcinowski (1980: 253)

described four body chambers of S. baculoides from the same bed; all are of the

same size, but with two different types of aperture. Three have well-developed

latero-ventral lappets (Fig. 50B), with the apertural constriction most

pronounced on the venter. The fourth specimen has a simple aperture with the

margin more or less parallel to the last rib, and no lappets. Marcinowski’s

(1980: 253) interpretation is that this is a special type of dimorphism; the micro-

and macro-conchs are indistinguishable in shell size and ornamentation, but

differ in the shape of the aperture. Marcinowski’s interpretation has still to be

tested on other species; certainly none of the other figured specimens of

S. baculoides with apertures show this type of dimorphism (see e.g. Mantell

1822, pl. 23 (figs 6-7); Sowerby 1822, pl. 592 (figs 2-3); D’Orbigny, 1842,

pl. 138 (figs 6, 8, 9); Crick 1896, figs a-e; Noetling 1885, pl. 8 (fig. 7);

Matsumoto & Obata 1963, pl. 2 (figs 1, 3); Juignet & Kennedy 1976, pl. 1

(figs 3-4, 6), pl. 2 (fig. 1) (Fig. 47); Kennedy & Juignet 1983, figs ila—b, d-e,

13a-—b; Thomel 1992, pl. 10 (fig. 2), pl. 11 (figs 1, 3); Wright & Kennedy

1995); instead they clearly show size-related dimorphism (Fig. 49).

Fig. 47. Macro- and microconch of Sciponoceras gracile

hima 1860). After Kennedy (1988, pl. 20 (figs 5, 14)).

66 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 48. Aperture of Sciponoceras cucullatum

Collignon, 1964, based on cast of holotype.

Some examples of Sciponoceras with well-preserved apertures include: |

Sciponoceras sp.—Marcinowski (1980: 254, pl. 3 (fig. 16)), from the Middle

Cenomanian of Dagestan with a pronounced pair of ventrolateral lappets |

(Fig. SOA).

Sciponoceras cucullatum Collignon (1964: 38, pl. 320 (fig. 1458)) from the |

Lower Cenomanian of Madagascar, with a distinctly recurved aperture |

(Fig. 48). |

Sciponoceras kossmati (?non Nowak, 1908). Matsumoto & Obata (1963, pl. 3 |

(fig. 2), pl. 4 (fig. 1)) from the Middle Cenomanian of Hokkaido, with a distinct

collar along the apertural margin.

Fig. 49. Aperture of Sciponoceras baculoide (Mantell, 1822). After Juignet &

Kennedy (1976, pl. 1 (fig. 3)).

THE AMMONITE FAMILY BACULITIDAE 67

Fig. 50. A. Aperture of Sciponoceras sp. with

distinct lappets. After Marcinowski (1980, pl. 3

(fig. 16)). B. Sciponoceras baculoide (Mantell,

1822), alleged microconch with lappets. After

Marcinowski (1980, pl. 3 (fig. 17)).

Sciponoceras orientale Matsumoto & Obata (1963, pl. 7 (fig. 3)) from the

Lower and Middle Turonian of Hokkaido. It has a hooded aperture with trigonal

ventral rostrum and acute apex.

Sciponoceras intermedium Matsumoto & Obata (1963, pl. 8 (fig. 1), pl. 11

(fig. 6)) from the Upper Turonian of Hokkaido—the aperture is oblique, facing

dorsally, the ventral rostrum is gently curved, the dorsal rostrum very short.

Sciponoceras matsumotoi Inoma (1980, text-fig. 4c, pl. 21 (figs 14, 16)) from

the Cenomanian of Hokkaido—the aperture is oblique and the ventral rostrum

slightly curved.

To summarize, apertures in microconch Sciponoceras are quite diverse, but

generally form a slight hood, or may even recurve, some with prominent

ventrolateral lappets, others without. Size and development of the dorsal

rostrum varies. In some, possibly macroconchs the aperture is simple and

oblique. With the possible exception of the case discussed by Marcinowski

(1980) dimorphism is also manifested by marked differences in size.

Apertures are quite commonly preserved in Baculites. Figured examples

include:

B. anceps Lamarck. Howarth (1965, pl. 5 (fig. 4)); Kennedy (1986c, pl. 20

(fig. 2)).

B. androtsyensis Collignon (1970, pl. 607 (fig. 2272)).

B. aquilaensis Reeside (1927a, pl. 8 (figs 1, 4, 7, 12); Reeside (1927Db, pl. 2

(figs 1, 4, 7, 12)).

B. asperiformis Meek. Cobban (19625, pl. 106 (figs 14-16)).

B. capensis Woods. Matsumoto & Obata (1963, pl. 19 (fig. 2)); Klinger &

Kennedy (1997, figs 46, 48A, 51-52).

B. codyensis Reeside (1927a, pl. 2 (figs 1-3) as B. asper); Kennedy & Cobban

(1991a, pl. 15 (figs 13-15)).

. mariasensis Cobban—Kennedy & Cobban (1991a, pl. 14 (figs 6-9)).

inornatus Meek—Obata & Matsumoto (1963, pl. 24 (fig. 6)).

obtusus Meek—Birkelund (1965, pl. 9 (fig. 3), pl. 13 (fig. 2)).

. occidentalis Meek—Matsumoto (1959, pl. 35 (fig. 2)).

. ovatus Say—Cobban (1974, pl. 2 (figs 13-14)).

. rectangulatus Collignon (1970, pl. 611 (fig. 2281)).

by by by by ty

68 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 51. Apertures of macro- and microconchs of Baculites capensis Woods, 1906.

Both x 0.9.

B. subanceps Haughton—Matsumoto (1959, pl. 35 (fig. 1) as B. aff. anceps);

Klinger & Kennedy (1997, fig. 130g).

. sulcatus Baily—Klinger & Kennedy (1997, figs 67h-j, 75a-c).

tanakae Matsumoto & Obata—Matsumoto & Obata (1963, pl. 16 (fig. 4)).

. thomi Reeside—Cobban & Kennedy (1991a, pl. 1 (figs 7-10)).

. undulatus d’Orbigny—Matsumoto & Obata (1963, pl. 11 (figs 2-3)).

. vanhoepeni Venzo—Klinger & Kennedy (1997, fig. 92d, g—h).

. ventroplanus Collignon—(1969, pl. 520 (fig. 2050).

. yokoyamai Tokunaga & Shimizu—Matsumoto & Obata (1963, pl. 11 (fig. 1),

pl. 14 (fig. 4); Renz (1982, pl. 34 (fig. 4)) (as B. inornatus); Cobban 1990, pl. 9

(figs 21-22).

In our South African material (Klinger & Kennedy 1997), apertures are

preserved in B. bailyi, B. capensis, B. sulcatus, B. vanhoepeni, B. nibelae and

B. aff. rectus.

The known apertures in all Baculites have a short dorsal rostrum, a longer

ventral rostrum, and lateral sinuses, but the relative proportions and orientation

vary considerably. In the earliest, Turonian, Baculites, e.g. B. undulatus and

B. yokoyamai the ventral rostrum is short, the dorsal one poorly developed, and

the aperture may face slightly upward, very much as in some Sciponoceras. In

later Baculites the ventral and dorsal rostra are more prominent. In some, the

ventral rostrum is very long and lingoid to spoon-shaped, with a distinct lateral

sinus (e.g. B. sulcatus in Klinger & Kennedy 1997, fig. 67h-j; B. thomi in

by Oy by by by by by

THE AMMONITE FAMILY BACULITIDAE 69

Cobban & Kennedy 1991a, pl. 1 (figs 7-10). In some Baculites, the aperture is

distinctly flared, e.g. B. obtusus in Birkelund (1965, pl. 13 (fig. 2a-c));

B. asperiformis Meek in Cobban (1962), pl. 106 (figs 14-16)) and some

B. vanhoepeni, or associated with a slight constriction, e.g. B. rex Anderson in

Matsumoto (1959, pl. 40 (fig. la—-c)) or B. subanceps Haughton (as B. aff.

anceps in Matsumoto 1959: 140). According to Birkelund (1965: 62), there

seems to be some variation in the aperture of B. obtusus; some specimens, e.g.

Birkelund (1965, pl. 10 (fig. la—c)) have less-inflated apertures; others, as

mentioned above (p. 59), retain traces of an early flared aperture on the

phragmocone.

In the South African material, the disparate sizes at which apertures are

formed in B. capensis (Fig. 51), B. sulcatus and B. vanhoepeni (Fig. 53) is

striking. In small specimens the apertures are usually slightly flared, whereas in

larger specimens this is less apparent, but this feature does not seem to be

consistent. |

Where large samples are available, e.g. in B. capensis (Fig. 52) and

B. vanhoepeni (Fig. 53), there seems to be a gradation from large to small

specimens with apertures, with no distinct, separate double peaks, indicating

micro- and macroconchs. This shows that there is a considerable degree of

overlap in size at which micro- and macroconchs form apertures. However, the

difference in size between the largest macroconch and the smallest microconch

is impressive.

Size distribution in Baculites capensis

@ indicates distribution of specimens with preserved

apertures.

NUMBER OF SPECIMENS

—_

us 10 15 20 25) . 30

MAXIMUM WHORL HEIGHT IN MM

Fig. 52. Histogram illustrating size distribution in Baculites capensis Woods, 1906,

to illustrate dimorphism.

70 ANNALS OF THE SOUTH AFRICAN MUSEUM

To summarize, the aperture in Baculites is relatively simple, parallel to the

long axis of the shell, with a short dorsal and a long ventral rostrum and a

lateral sinus of variable depth. In some it may be distinctly flared, but this is not

a very common feature. In general, dimorphism in Baculites seems to be only

size related. There is a large range of overlap in size between micro- and

macroconchs, and distinct separation into two size groupings is not clear in

some species, e.g. B. capensis and B. vanhoepeni.

Boehmoceras is thus far only known by two Upper Santonian species,

B. arculus (= B. loescheri Riedel, 1931) and B. krekeleri. These two species

co-occur in Austria, and Summesberger (1979) suggested that the two ‘species’

are in fact a dimorphic pair—B. arculus the microconch, and B. krekeleri the

macroconch. In the Gulf Coast Region of the USA, Boehmoceras is only

represented by B. arculus. Kennedy & Cobban (1991b: 183) regard specimens

of B. arculus with whorl heights of 15-17 mm at the base of the body chamber

as macroconchs, and small specimens with whorl heights of about 10 mm as

microconchs.

The aperture of Eubaculites is not as well-known as that of Baculites, but

seems to follow the same pattern, except that the whorl section does not expand.

Figured examples include:

E. vagina (Forbes)—Klinger & Kennedy (1993, fig. 12a—c) (Fig. 54A).

E. latecarinatus (Brunnschweiler)—Klinger & Kennedy (1993, fig. 46)

(Fig. 54B).

2E. simplex (Kossmat)—Klinger & Kennedy (1993, fig. 53b).

10 Size distribution in Baculites vanhoepeni

Oiindicates distribution of specimens with preserved

apertures.

NUMBER OF SPECIMENS

20 30 40 50

MAXIMUM WHORL HEIGHT IN MM

Fig. 53. Histogram illustrating size distribution in Baculites vanhoepeni

Venzo, 1936, to illustrate dimorphism.

THE AMMONITE FAMILY BACULITIDAE 71

The pattern of distribution of maximum whorl height in E. carinatus

(Fig. 44) from Neuquén, Zululand and Quiriquina, already mentioned above, is

interesting. The Zululand and Neuquén specimens respectively show a bimodal

distribution, suggesting size-related dimorphism, but the peaks do not fully

coincide. The Quiriquina specimens all occur within the size range of the

Zululand microconchs. We admit that these data are far from complete, but they

do suggest that differences in size occur between populations from different

localities, as suggested above, in addition to sexual differences.

The apertures of the remaining Maastrichtian baculitid lineages, e.g.

Fresvillia, are unknown but judging by the general pattern in the rest of the

Baculitidae, we assume that these genera are also dimorphic.

SUMMARY

1. Geographic distribution

Early representatives of the family, Lechites and Sciponoceras had a wide

north-south distribution, but appear absent or very rare in West Africa.

Baculites appears to have separated into distinct endemic lineages from the

Coniacian onwards; that of the US Western Interior is most conspicuous, and

persisted until the early Maastrichtian. In the Maastrichtian, Eubaculites

dominated the Gondwanan regions and Baculites was scarce. In contrast, in the

Northern Hemisphere, Eubaculites was a rarity and Baculites dominant.

Eubaculites shows distinct latitudinally restricted distribution; it is absent from

high northern and southern latitudes and equatorial regions, as well as from

West Africa, North Africa and the Middle East.

2. Coiling

Lechites and Sciponoceras are straight, except at the aperture. Some

Coniacian-Santonian Baculites may have curved body chambers, but this

feature is not consistent. Others are either completely straight or the whole shell

may be curved. Except in endogastric Boehmoceras and exogastric Crio-

baculites where the whole shell is a coiled criocone, curvature appears of little

taxonomic value.

3. Whorl section

Lechites and Sciponoceras have circular or elliptical whorl sections. Early,

Turonian Baculites have elliptical whorl sections; in later forms it varies

between species, but seems to be rather consistent within the same species.

Ventral keels first appear in the Santonian, but are not common, and only

dominate in the Maastrichtian. Near the Campanian—Maastrichtian boundary a

number of baculitid lineages acquire a cuneiform or trigonal whorl section. The

predominantly Maastrichtian genus Eubaculites has a distinct pyriform whorl

section with tabulate venter or trigonal whorl section with narrow venter. Late,

Maastrichtian aberrant baculitids have circular whorl sections.

4. Suture line

Early Baculites have a simple suture with quadrate saddles and lobes.

Pseudobaculites has a complex suture compared to contemporary Baculites. A

progressive increase in complexity of suture line took place in all Baculites

72 ANNALS OF THE SOUTH AFRICAN MUSEUM

lineages from the Coniacian onwards. In the Campanian, several lineages with

very complex sutures appeared suddenly. In the Maastrichtian, several small

lineages with simplified suture occur.

5. Ornament

Ornament in Lechites and Sciponoceras is simple: circumperipheral ribbing

or smooth with constrictions is a constant feature in the latter. In contrast,

ornament in Baculites is extremely variable. Some ornamented species have rare

smooth variants, and some smooth species have rare irregularly ornamented

forms. Also, smooth species may occur in otherwise ornamented lineages.

Lateral tubercles first occur in the Upper Turonian, and become dominant in the

Coniacian and Santonian, but persist to the Lower Campanian in the Indo-

Pacific. Prominent lateral ribbing is conspicuous in the Middle Campanian in

the US Western Interior and the Indo-Pacific.

6. Size, dimorphism and apertures

Early Baculites are small, but show a gradual increase in size, reaching

maximum size in the Upper Campanian—Lower Maastrichtian. True giants also

occur in some Eubaculites in the Maastrichtian. The family as a whole is

dimorphic. In Lechites apertures are different in macro- and microconchs.

Those of Sciponoceras are more varied, and of specific as well as of sexual

significance. Early Baculites have slightly curved apertures with short ventral

and dorsal rostra; later forms are straight, with longer rostra and may be

expanded. Eubaculites has a short dorsal and long ventral rostrum that is not

flared.

GENERAL COMMENTS ON BACULITIDAE

APTYCHI

Aptychi of Baculitidae are rarities. These are calcitic objects and would be

expected to have a high preservation potential. The best known occurrences are

in European chalk facies where original aragonitic shells are lost, but even here,

aptychi are absent from levels of abundant baculitids, as in the Middle

Cenomanian of Southern England. There is little doubt that development of

calcitic aptychi was limited to certain species only, as in the Scaphitaceae, and

perhaps only in some individuals, and that most species had unmineralized jaw

apparatus.

Thus far only two known occurrences of aptychi in Sciponoceras are

known: the first by Fritsch (1895: 78, text-fig. 63c) from Bohemia; the second

by Breitkreutz et al. (1991, figs 6-7) from the Upper Cenomanian

Neocardioceras juddii Zone of Ostwestfalendamm, near Bielefeld in the

Minster Basin in Germany. This specimen is shown as Figure 56 and

Fig. 54 (see facing page). Apertures in Eubaculites. A. E. vagina (Forbes, 1846).

BMNH C51146 from the Valudavur Formation of southern India. B. E. latecarinatus

(Brunnschweiler, 1966). SAS-H60H/1 from bed H, locality 133, Zululand, St Lucia

Formation, Maastrichtian I. Both x 1.

THE AMMONITE FAMILY BACULITIDAE

ANNALS OF THE SOUTH AFRICAN MUSEUM

F G

. 55. A. Rugaptychus flexus (Moberg, 1885). Syntype, Lund University Collection,

the original of Moberg (1885, pl. 1 (fig. 19)) from the Campanian of K6pinge,

Sweden. B-H. Rugaptychus insignis (Hébert, 1856) (= R. rugosus (Sharpe, 1857)).

B-C. Lund University Collections, the original of Moberg (1885, pl. 1 (figs 16-17)).

D-E. Natuurhistorisch Museum, Maastricht Collections no. 6309, from the lower third

of the Upper Campanian Zeven Wegen Chalk of Haccourt, Belgium. F. Syntype of

R. rugosus Norwich Castle Museum collections no. 3519 from the Upper Campanian

Chalk of Norwich, the original of Sharpe (1857, pl. 24 (fig. 31)). G. Lund University

Collections, the original of Moberg (18 (1885, pl. 1 (fig. 15)) from the Campanian of

Kopinge, Sweden. H. Lund University Collections, the original of Moberg (1885,

pl. 1 (fig. 14)), also from K6pinge. All x 1.

THE AMMONITE FAMILY BACULITIDAE 13

photographs were kindly supplied by Dr Bichner (Bielefeld) via Mr R.

Metzdorf. The Sciponoceras is ill-preserved but compares to the early

subspecies anterius Wright & Kennedy, 1984, of S. bohemicum (Fritsch &

Schlénbach, 1872). The specimen is 105 mm long, with a maximum preserved

whorl height of 7 mm. The adapertural 60 mm of the composite mould is body

chamber, lacking the adapertural part, the aptychus lies 23 mm from the last

septum, and is exposed as a convex mould of the inner, concave surface. There

are traces of what appears to be the original calcite material of the jaw,

suggesting it to be an aptychus rather than anaptychus. The partial exposure of

the specimen means that it is not clear if two plates are represented, or a single

fused plate. Ornament is of relatively coarse ridges and grooves that appear to

parallel the outer and lateral margins; the structure is 5 mm in maximum length,

the two halves 3.5 mm in maximum width. The specimen agrees with material

from the Upper Cenomanian Metoicoceras geslinianum Zone of southern

England described by Wright & Kennedy (1981: 118, pl. 31 (fig. 19)) as being

possibly associated with Scaphites, although Sciponoceras occurs in the same

interval and the latter assignation now seems more likely.

The form genus Rugaptychus Trauth belongs to the genus Baculites (Trauth

1927: 245; Arkell 1957b: L440). Records of Baculites with Rugaptychus are

nearly all from Campanian Boreal Chalk facies in Europe (Fig. 55).

Trauth (1927: 245) recognized the following species in Rugaptychus:

R. rugosus (Sharpe, 1853). Upper Campanian (Sharpe 1853: 57, pl. 24

(figs 8a—b, 9); Binckhorst 1861: 33 (said to be Maastrichtian); Lundgren 1874:

70, et seq., pl. 3 (fig. 14); Moberg 1885: 41, pl. 1 (figs 14-18), pl. 6 (fig. 26);

Blackmore 1896: 532-3, pl. 16 (fig. 16); De Grossouvre 1908: 39, pl. 10

(figs 7-13)).

R. rugosus insignis (Hébert, 1856). Upper Campanian (Hébert 1856: 367,

pl. 28 (fig. 6).

R. knorrianus Trauth, 1927. Upper Campanian (Schliiter 1876: 147, pl. 39

(fig. 16)).

R. flexus (Moberg, 1885). Upper Campanian (Schliiter 1876: 144, pl. 40

(fig. 8); Moberg 1885: 43, pl. 1 (fig. 19), pl. 6 (fig. 25)).

Rugaptychus form 1 Sharpe. Upper Campanian (Sharpe 1853: 58, pl. 26

(fig. 10a—b)). :

Rugaptychus form 2. Maastrichtian (Ravn 1902: 259). This specimen was not

figured and the identification is questionable.

The only possible older species of Rugaptychus is the specimen figured by

Fritsch (1893: 80, fig. 63c—-e) from the Coniacian of Bohemia.

To these may also be added: B. leopoliensis with aptychus in Nowak (1908:

339, pl. 14 (fig. 11)) and B. vertebralis Giers non Lamarck with aptychus in

Giers (1964: 256). Giers (1964: 256, text-fig. 2) also figured some problematic

structures in B. vertebralis, which he tentatively interpreted as jaw apparati

Fig. 56 (see overleaf). Sciponoceras bohemicum anterius? Wright & Kennedy, 1984.

Specimen with aptychus from the Upper Cenomanian Neocardioceras juddi zone of

Ostwestfalendamm near Bielefeld in the Miinster Basin, Germany, the original specimen

figured by Breitkreutz et al. (1991: 42, text-fig. 6). A x 4; B x 1.5.

ANNALS OF THE SOUTH AFRICAN MUSEUM

SNS

ASS

ig. 56

THE AMMONITE FAMILY BACULITIDAE

Fig. 57. Ammonitella of Baculites (small) and Scaphites (large) associated with

juvenile Baculites codyensis Reeside, 1927a. USNM 507252, from USGS Mesozoic

locality 21425, Santonian, Marias River Shale on east bank of Marias River,

18.5 km (11 miles) south-west of Shelby in the W% NE% SE% sec. 14, T. 31 N.,

R. 4 W., Toole County, Montana. x 2.

yyy

yf,

YG, Ui;

Y dy

ANNALS OF THE SOUTH AFRICAN MUSEUM

THE AMMONITE FAMILY BACULITIDAE 79

(Mundwerkzeuge). One of us (WJK) has examined the specimen and concluded

that the structures do not belong to the baculite. Picard (1929: 436) recorded an

aptychus in the same layer as B. vertebralis. Kennedy (1986c: 192, pl. 16

(figs 1-22)) figured numerous specimens of R. rugosus from the Campanian of

Folx-les-Caves, Belgium (previously figured by De Grossouvre 1908, pl. 10

(figs 7-13)), and later (Kennedy 1993: 114, pl. 7 (figs 18-25)) from Mons,

Belgium. Kennedy & Christensen (1997, fig. 3la-h) figured some aptychi from

southern Sweden, including some of Moberg’s original specimens.

Given the large number of Baculitidae, especially Baculites and Eubaculites,

often well-preserved in concretions, it is very surprising that neither aptychi nor

anaptychi associated with these genera have been recorded from the US Western

Interior, West Greenland, the Gulf Coast and Atlantic Seaboard regions of the

USA, Hokkaido, Madagascar, Zululand, Australia, India, Argentina and Chile.

MASS OCCURRENCES

Baculitidae, especially Baculites often occur in great monospecific

concentrations in single concretions, on single bedding planes and in limited

intervals such as individual concretions or layers of concretions, e.g.B. ‘ovatus’

(Reeside 19275, pl. 7 (figs 1-2)); B. asperiformis (Cobban 1962b: 708);

B. mclearni (Cobban 1962b: 712); B. perplexus (Cobban 1962b: 714; Gill &

Cobban 1966, pl. 11 (fig. 3)); B. cuneatus (Cobban 1962a: 128); B. jenseni

(Cobban 1962a: 130); B. perplexus (Kennedy & Cobban 1976, pl. 8 (fig. 2));

B. yokoyamai (Cobban & Hook 1983: 7); Sciponoceras gracile (Kennedy 1988:

109, text-fig. 38; herein Fig. 60); Eubaculites carinatus (Klinger & Kennedy

1993, fig. 37); and Baculites bailyi, B. capensis and B. duharti (Klinger &

Kennedy 1997).

In the US Western Interior, concretion layers filled with Baculites extend

over many hundreds of square kilometres. In southern England, Sciponoceras

baculoides occurs in profusion in a sequence of three limestone/marl couplets

(Kennedy 1969) in the Middle Cenomanian part of the Lower Chalk over an

area of more than 20000 square kilometres. Gale (1989, 1995) has shown these

couplets to represent an 18 000-23 000 Milankowitch cyclicity; for

72 000-92 000 years, Sciponoceras was the dominant organism preserved as

body fossils, but is very rare or absent in the Lower Cenomanian Chalk below.

It is equally abundant in contemporary phosphatized remanié faunas in

condensed units resting on hard grounds in Dorset (Kennedy 1970), Haute

Normandie (Juignet & Kennedy 1976) and Sarthe (Kennedy & Juignet 1983).

Similar brief flood abundances on single bedding planes can be recognized in

southern France with the Lower Cenomanian Sciponoceras sp. at Dieulefit

(Dréme) or Sciponoceras bohemicum anterius in the Upper Cenomanian

Neocardioceras juddii Zone in Devon, England.

Fig. 58 (see facing page). USNM 507253, from the Niobrara Shale, Santonian,

with mass occurrence of juvenile Baculites codyensis Reeside, 1927a, and rare

ammonite jaws, from USGS Mesozoic locality D6619, Seminoe Dam south-west

quadrangle in NW% SW% sec. 23, T. 25 S., R. 85 W., Carbon County,

Wyoming. x 5.

80 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 59. U.S. Geological Survey Collections, Denver, from the Niobrara Shale, Santonian

with mass occurrence of juvenile Baculites codyensis Reeside, 1927a, and rare ammonite

jaws, from USGS Mesozoic locality D6619, Seminoe Dam south-west quadrangle in NW%

SW% sec. 23, T. 25 S., R. 85 W., Carbon County, Wyoming. x 5.

THE AMMONITE FAMILY BACULITIDAE

Fig. 60. Mass occurrence of Sciponoceras gracile (Shumard, 1860). J. P. Conlin Collection

no. 7490, from 1.5 to 1.8 miles south-east of Britton, Ellis County, Texas. x 1.

82 ANNALS OF THE SOUTH AFRICAN MUSEUM

Mass occurrences are generally made up of whole or fragmentary adult or

subadult individuals and may sometimes show size-sorting. In the US Western

Interior, there are also rare occurrences with associated ammonitella and

juveniles (Figs 57-59), or laminae with abundant juveniles may occur in

concretions dominated by adults. Much rarer are mass occurrences of juveniles

only, as have been figured from Zululand (Klinger & Kennedy 1997) of

Baculites bailyi.

In most of these occurrences, including our Zululand material, these

Baculites show subparallel alignment, suggesting post-mortem drifting and

current alignment. However, the preservation and attachment to the rest of the

shell of the very delicate ammonitellas in some of these assemblages seems to

contradict current alignment of the shells. The very fine silty lithology of these

concretions suggests a low energy sedimentary environment. A detailed analysis

of the accompanying invertebrate faunas might throw more light on the

sedimentary environment of these mass occurrences of baculitids. The mass

occurrences suggest that, for part of their life-cycles at least, baculites were

gregarious.

AMMONITELLA AND EARLY ONTOGENY

In contrast to normally coiled ammonites where the ammonitella is partially

covered and protected by the succeeding planispiral whorls, the ammonitella of

the baculitids is very vulnerable, being totally exposed and attached to the rest

of the straight shell by a slender shaft. Notwithstanding these unfavourable

conditions, ammonitellas, often still attached to the early shaft have been

recorded on several occasions in Baculites—see e.g. Bandel et al. (1982, text-

figs 1c, 2d); Birkelund (1979, text-fig. 3a-c, g-h); 1981, text-fig. 4a—b?, c;

1993, text-fig. 6a—b, d-i)); Brown (1891: 159-160; 1892: 136-141, pl. 9

(figs 1-11)) (B. compressus); Cobban (1962b: 708-709); Landman (1982:

1235-1241, text-figs 1-2; 1987: 158, fig. 25c-d, p. 160, fig. 26); (1994);

Landman & Bandel (1985, figs 16-26); Reeside (1927b, pl. 7 (figs 6-8))

(B. ovatus), Reyment (1958: 7, fig. la—d) (B. yokoyamai? as B. ‘ovatus’ Say?)

and Smith (1901: 39-49). Our own material of B. bailyi (Klinger & Kennedy

1997) from the Coniacian also shows several specimens with the ammonitella

still attached to the early shaft. In some cases assemblages are dominated by, or

consist exclusively of such juveniles, suggesting age segregation of adults and

juveniles.

Unfortunately, it is not possible to dissect our Zululand specimens to

determine the early sutural ontogeny, nor is the preservation good enough to

show the early micro-ornament. For detailed discussions on these features the

reader is referred to Brown (1892), Smith (1901), Landman (1982) and Bandel

et al. (1982).

Dr W. A. Cobban (U.S. Geological Survey, Denver) has kindly allowed us

to illustrate a series of well-preserved Baculites codyensis Reeside, 1927a, from

the Santonian Colorado Shale (now Marias River Shale) of Toole County,

Montana (Figs 61-65). Specimens include ammonitella (Fig. 62A, B) and

ammonitella with juvenile phragmocone and, perhaps, body chamber

(Fig. 61A-B). Ammonitella with shell preserved (Fig. 63A-B) show the lack of

THE AMMONITE FAMILY BACULITIDAE

Fig. 61. Baculites codyensis Reeside, 1927a, USNM 507254, from

USGS Mesozoic locality 21425, the Santonian Marias River Shale on the east

bank of the Marias River, 18.15 km (11 miles) south-west of Shelby, in the

W'% NE'% SE% sec. 14, T. 31 N., R. 4 W., Toole County, Montana.

A x 15; B x 100.

84 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 62. Baculites codyensis Reeside, 1927a. A. USNM 207255. B. USNM 507256.

Both from USGS Mesozoic locality 21425, the Santonian Marias River Shale on the east

bank of the Marias River, 18.15 km (11 miles) south-west of Shelby, in the W'Z NE%

SE% sec. 14, T. 31 N., R. 4 W., Toole County, Montana. A x 90; B x 120.

THE AMMONITE FAMILY BACULITIDAE

Fig. 63. Baculites codyensis Reeside, 1927a, USNM 507257, from USGS Mesozoic

locality 21425, the Santonian Marias River Shale on the east bank of the Marias River,

18.15 km (11 miles) south-west of Shelby, in the W144 NE% SE% sec. 14, T. 31 N.,

R. 4 W., Toole County, Montana. A x 40; B x 90.

86 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 64. Baculites codyensis Reeside, 1927a, USNM 507258, from USGS Mesozoic

locality 21425, the Santonian Marias River Shale on the east bank of the Marias River,

18.15 km (11 miles) south-west of Shelby, in the W124 NE% SE% sec. 14, T. 31 N.,

R. 4 W., Toole County, Montana. A x 22; B x 110.

THE AMMONITE FAMILY BACULITIDAE 87

Fig. 65. Baculites codyensis Reeside, 1927a, USNM 507259, from USGS Mesozoic

locality 21425, the Santonian Marias River Shale on the east bank of the Marias River,

18.15 km (11 miles) south-west of Shelby, in the WZ NE% SE% sec. 14, T. 31 N.,

R. 4 W., Toole County, Montana. A x 14; B x 110.

88 ANNALS OF THE SOUTH AFRICAN MUSEUM

growth lines and microtuberculation as described by Bandel et al. (1982),

Landman (1982) and Landman & Bandel (1985), whereas internal moulds show

details of early suture ontogeny (Figs 61B, 62A, 64B, 65B) and the nepionic

constriction (Figs 61A, 64A-B).

MUSCLE SCARS

Records of muscle scars in Baculitidae are rare. In a pioneering study nearly

a century ago, Crick (1898) described and figured muscle scars in Baculites

ovatus (1898: 77, pl. 17 (figs 1-4)) and Eubaculites vagina (1898: 78, pl. 17

(fig. 1)). Kennedy & Cobban (1976, pl. 2 (fig. la—b)) figured paired dorsal

muscle scars in Baculites codyensis. Klinger & Kennedy (1980: 299, figs 3d,

4b, 5b) recorded a slight, median dorsal ridge in Eubaculites latecarinatus and

in E. vagina. They suggested that this structure might be associated with the

muscular attachment of the animal to the shell. Henderson (1984), in contrast,

suggested that subepithelial muscle attachment in ammonites was along the line

of contact of the septum and shell wall, and that the ‘muscle scars’ of previous

workers perhaps represented the ‘imprint of internal organs’, although they

could be viewed as ‘additional sites of muscle attachment, separated from those

Fig. 66. Copy of part of Crick’s (1898) plate 17. Figs 1-4 are labelled as Baculites ovatus

Say, Fig. 5 as Baculites vagina Forbes, and Fig. 8 as Hamites maximus J. Sowerby.

THE AMMONITE FAMILY BACULITIDAE 89

Fig. 67. Baculites sp. A-C. U.S. Geological Survey collections, Denver, locality D2140,

from the Pierre Shale, Campanian, 12.2-21.3 m above the top of the Groat Sandstone at

Mesozoic locality D2140, 25.6 km north of Belle Fourche in SW'4 SE% sec. 22, T. 11 N.,

R. 2 E., Butte County, South Dakota. The specimen has been coated with ammonium

chloride and shows the narrow groove on the internal mould that corresponds to the ridge on

the inside of the adapical end of the body chamber that defines the adapertural edge of the

bilobed dorsal muscle scar. The groove adapertural to the final septum corresponds to the

position of the postseptal prismatic zone of Henderson (1984). x 1.

of the subepithelial sheath’ (1984: 480). Henderson illustrated and described

pre- and post-septal prismatic zones, the latter in his view the site of muscle

attachment (1984, text-fig. 9).

Through the courtesy of Dr W. A. Cobban (U.S. Geological Survey,

Denver), we have been able to examine a series of rare and exceptionally

preserved Baculites from the US Western Interior that show detail of muscle

scars and related features with great clarity. Only a tiny percentage of generally

adult Baculites show muscle scars; development is variable even in individuals

from a single concretion. The post-septal prismatic zone of Henderson forms a

distinct groove, adapertural to the septa (Fig. 67). There is a well-developed

adaperturally biconvex groove on the mould (Figs 67-71), corresponding to a

ridge of shell material on the inside of the body chamber, that follows the

general line of the U/I saddle on the dorsum, and runs back towards the trace of

the post-septal prismatic zone. This bilobed trace is presumed to represent the

fused dorsal muscle scars described in other ammonites. Much rarer, and

90 ANNALS OF THE SOUTH AFRICAN MUSEUM

generally fainter, is an adaperturally convex groove that defines a single, ventral

muscle scar (Figs 69-70). Rare specimens show two sets of successive scars in

the adapical part of the body chamber, and one specimen (Figs 70-71) has a

series of successive scars on the walls of the last few chambers of the

phragmocone and the adapical part of the body chamber. These traces

presumably mark the adapertural boundary of the muscle scar associated with

temporary attachment following the pause in forward movement of the body that

coincided with the secretion of the last few septa.

Some internal moulds show delicate longitudinal ridges on both dorsum and

venter of moulds of phragmocone and body chamber, corresponding to ridges of

shell material on the inside of the shell. These may occur on specimens with the

muscle scar present (Fig. 69) or where they are not developed (Fig. 70). On the

dorsum there is a faint median groove, plus a lateral groove, corresponding to

lobe I and the outer flank of saddle U/I (Figs 69C, 70A).. A pair of closely

spaced median grooves may be present on the venter (Fig. 70), and there may

also be faint parallel striations.

None of the specimens studied showed any trace of an ‘annulus’ as

described by Crick (1898), and we accordingly re-examined his material. The

original of Crick’s Baculites ovatus are examples of one of the feebly

ornamented Campanian-—Maastrichtian Baculites from the Pierre Shale; the

locality is given as ‘Fox Hills beds (Upper Cretaceous) of Horse Head Creek,

South Dakota’. The original of Crick’s pl. 17 (figs 1-3) is BMNH C54150. It is

illustrated here as Figure 73D-H, and Crick’s original drawings are reproduced

as Figure 73A-C. It will be seen that the reported dorsal and ventral muscle

scars shown by Crick are indeed present. The original of his pl. 17 (fig. 4), is

shown here as Figure 74A and 74E; his original drawing is reproduced as

Figure 74B. We were unable to detect the annulus he described and illustrated.

We also examined the specimen of Eubaculites vagina (Forbes, 1846) described

and illustrated by Crick (1898, pl. 17 (fig. 5)), BMNH C73570) but were unable

to convince ourselves that a bilobed groove delineating a dorsal muscle scar was

indeed present (Fig. 74C, D, F).

DRAG BANDS AND RELATED FEATURES

Rare Baculites from the US Western Interior show longitudinal markings on

internal moulds of the chambers of the phragmocone (Fig. 75). Similar

structures have been discussed most recently by Zaborski (1986) and Hewitt et

al. (1991), and variously interpreted. In our view, they record the forward

movement of the tissues between successive pauses associated with septum

formation. The markings correspond to slight irregularities on the mould, and

variation in surface texture. The siphonal region of one specimen (Fig. 75B)

Fig. 68 (see facing page). Baculites haresi Reeside, 1927b, USNM 507260, from USGS

locality D1587, from the Gammon Member of the Pierre Shale, sec. 3, T. 10 N., R. 4 E.,

Butte County, South Dakota. A-B. Flanks (uncoated). C-D. Dorsum (coated in C,

uncoated in D). The bilobed dorsal muscle scar accompanied by a groove (C) with traces of

the original shell material (D). x 1.

THE AMMONITE FAMILY BACULITIDAE

Fig. 68

92 ANNALS OF THE SOUTH AFRICAN MUSEUM

B Cc D

Fig. 69. Baculites codyensis Reeside, 1927a. U.S. Geological Survey Collections, Denver;

from the Marias River Shale, Santonian, USGS MesoZoic locality 21425, near Shelby,

Montana. A and C are uncoated; B and D are coated with ammonium chloride. The venter

(A, B) shows a single incomplete adaperturally convex groove marking the adapertural edge

of the ventral muscle scar. The dorsum (C, D) shows the bilobed trace of the adapertural

edge of the dorsal muscle scar. In D, a median and a pair of lateral grooves (corresponding

to ridges on the inside of the shell) are present on both phragmocone and body chamber.

Their function is uncertain. x 2.

Fig. 70 (see facing page). Baculites codyensis Reeside, 1927a. A-C. USNM 507261.

B. USNM 507262. From the Colorado Shale, Santonian, USGS Mesozoic locality 21425,

18.15 km (11 miles) south-west Shelby, in W'Z NE% SE% sec. 14, T. 31 N., R. 4 W.,

Toole County, Montana. A, E, show the dorsum; G, the venter of specimens showing well-

developed grooves on internal moulds that correspond to ridges on the inside of the shell.

THE AMMONITE FAMILY BACULITIDAE

. 70

94 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 71. Baculites haresi Reeside, 1927b, USNM 507263, from the Gammon Member of

the Pierre Shale, Campanian, USGS Mesozoic locality D1587, sec. 3, T. 10 N., R. 4E.,

Butte County, South Dakota. Dorsum (A) shows traces of four successive positions of the

bilobed dorsal muscle scar. Two are overlain by the final two septa of the phragmocone, and

two are visible at the adapical end of the body chamber. Flank (B) shows final dorsal muscle

scar at adapical end of body chamber clearly. Venter (C) shows traces of four successive

ventral muscle scars. x 2.

shows well-developed longitudinal lines, in part picked out by traces of shell, in

the siphonal region. These features were described by a number of previous

workers (see review in Kennedy & Cobban 1976), and interpreted as traces of

the ligaments supporting the siphuncle. The width of the structure figured here

suggests that it may be the trace of the adapertural migration of the ventral

muscle.

MICROSCULPTURES

Checa (1995) recognized eight morphological types of ‘microsculpture’ on

the outer surface of a range of well-preserved Jurassic and Cretaceous

ammonites, and interpreted them as having been produced by compressive

stresses acting on a free, uncalcified periostracum, and consequently preserved

by mineralization of the wrinkled surface. Structures of this type are quite

common in US Western Interior Baculites, both where the outer shell layer is

THE AMMONITE FAMILY BACULITIDAE 95

Way

Fig. 72. Baculites haresi Reeside,.1927b, USNM 507263,

from the Gammon Member of the Pierre Shale, USGS

Mesozoic locality D1587, sec. 3, T. 10 N., R. 4 E., Butte

County, South Dakota. Details of dorsum of specimen shown in

Fig. 71A, with positions of four successive dorsal muscle scars

indicated. x 6.

preserved, and on partially exfoliated surfaces within the nacreous layer

(Figs 76-78, 79A-C, E). Some of the most striking examples extend through on

to the internal mould (Fig. 76D-F).

ANNALS OF THE SOUTH AFRICAN MUSEUM

HO basis

‘I

135 toe

THE AMMONITE FAMILY BACULITIDAE S|

We are uncertain as to the origin of these structures, the best developed of

which may disrupt ornament and appear to record a pathological condition

(Fig. 76D-F, 77-78) rather than the exceptional preservation of normal

structures produced in an unmineralized periostracum.

FEATHER STRUCTURE

Arkell (1957a) used the term “feather structure’ for the low-relief adapically

diverging chevrons described from a range of generally oxycone or very

compressed feebly ornamented ammonites of Jurassic and Cretaceous age.

Wepfer (1920) referred to these structures as ‘Streifenbiischel’ and noted

their presence in what he termed Baculites compressus from the US Western

Interior. Feather structure is in fact relatively common in US Western Interior

Baculites (Fig. 79D). Well-preserved specimens show this structure to be

present on the outer surface of the outer shell layer (on rare occasions where

this is preserved), but is typically seen on exfoliated surfaces within the thick

nacreous layer, and is also present on the internal mould. Wepfer (1920)

thought this structure could possibly be the impression of the nidamental gland.

Kessler (1923) thought the structure to be associated with muscle

attachment, as did Seilacher (1988) who interpreted it as reflecting the

successive sites of attachment of the retractor muscle to the wall of the body

chamber. In an alternative interpretation Checa (1995: 885) regarded feather

structure as a variant of what he described as ‘adoral convergent wrinkles’

produced in unmineralized periostracum. He dismissed the muscle attachment

hypothesis on the basis that the feather structure was external rather than

internal. The present material shows it to be both (for further discussion of

feather structure see Enay & Dominjon 1964, 1967, and references therein).

PARASITISM

Distortion and irregular section of shell has been interpreted as the result of

parasitism by some workers (Keupp 1984, 1995; Keupp & Dietze 1987;

Hengsbach 1990, 1991, and reviews therein). Through the courtesy of Dr

W. A. Cobban and Messrs P. and N. Larson (Black Hills Institute of Geological

Research, Hill City, South Dakota) we have seen Baculites showing two

conditions that may be the result of parasitism. The most spectacular example

(Fig. 80) is a specimen of B. eliasi Cobban, 1958, from the Maastrichtian Pierre

Shale of Garfield County, Montana.

A partially exfoliated fragment from the adapical end of the body chamber

bears a large, asymmetric blister-like swelling on the venter (Fig. 80).

A possible interpretation of this specimen is that the mantle at the aperture of

the shell was either parasitized or infected, producing a tumour-like growth, but

Fig. 73 (see facing page). Baculites sp., BMNH C54150, from the ‘Fox Hills beds (Upper

Cretaceous) of Horse Head Creek, South Dakota’, the original of Crick (1898, pl. 17

(figs 1-3)). Crick’s figures are reproduced as A-C; photographs of the corresponding views

as D-F. A-F x 1; G, H x 1.5.

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 74

THE AMMONITE FAMILY BACULITIDAE 99

A B

Fig. 75. Drag bands in Baculites sp. A. USNM 507264. B. USNM 507265. From the

Pierre Shale, Campanian, 12.2-21.3 m above the top of the Groat Sandstone at USGS

Mesozoic locality D2140, 25.6 km north of Belle Fourche, in SW'%4 SE% sec. 22, T. 11 N.,

R. 2 E., Butte County, South Dakota. A shows drag markings associated with movement of

mantle on dorsolateral surface of mould. B shows drag markings on the venter associated

with the ventral muscle. A x 3; B x 2.

Fig. 74 (see facing page). A, B, E. Baculites sp. BMNHC54150, from the ‘Fox Hills beds

(Upper Cretaceous) of Horse Head Creek, South Dakota’, the original of Crick (1898, pl. 17

(fig. 4)). Crick’s original figure is reproduced as B, photographs of the corresponding view

as A and E. C, D, F. Eubaculites vagina (Forbes, 1846). BMNH C73570, Valudavur

Group, Maastrichtian, Pondicherry, South India, the original of Crick (1898, pl. 17 (fig. 5)).

Crick’s original figure is reproduced as D, photographs of the corresponding views as C

and F. A-D x 1; E, F x2.

100 ANNALS OF THE SOUTH AFRICAN MUSEUM

E F

Fig. 76. Baculites codyensis Reeside, 1927a. From Marias River Shale, Santonian, USGS

Merorore locality 21425, east bank of Marias River, 18.15 km (11 miles) south-west of

Shelby in W% NE% SE% Secu 47a SINE: R. 4 W., Toole County, Montana.

A-C. USNM 506266, well-preserved individual with shell present, showing normal ornament

and growth lines. D-F. USNM 507267, example with well-developed wrinkles affecting

whole of shell surface, and also present on internal mould in a more subdued form. All x 1.

Fig. 77 (see facing page). Baculites codyensis Reeside, 1927a. USNM 507267, Marias River

Shale, Santonian, USGS Mesozoic locality 21425, east bank of Marias River, 18.5 km

(11 miles) south-west of Shelby in W% NE% SE% sec. 14, T. 31 N., R. 4 W., Toole

County, Montana. Detail of specimen shown in Fig. 76D-F, showing well-developed

wrinkles. x 4.

THE AMMONITE FAMILY BACULITIDAE 101

102 ANNALS OF THE SOUTH AFRICAN MUSEUM

not affecting normal secretory processes, as a result of which the shell was

secreted around the growth, beyond which the mantle returned to its normal

shape, and a simple tubular shell, without growth irregularities, was secreted.

Keupp (1984, fig. 4; 1995, fig. 2) illustrated and described a similar blister-like

growth anomaly in a Jurassic (Kimmeridgian) Orthosphinctes.

The second distinctive growth anomaly that may be the result of parasitism

is shown by specimens of Baculites scotti Cobban, 1958, and Baculites sp. from

the Campanian Pierre Shale of South Dakota. In the case of the latter, 3 per cent

of a total of 300 specimens showed the abnormality. Gill & Cobban (1966: A27,

40, pl. 10 (figs 1-9)) had previously described and illustrated these structures on

moulds of Baculites gregoryensis Cobban, 1951, from the Pierre Shale of

Wyoming, and interpreted them as marking the attachment areas of some

unknown organism.

The structures are restricted to the body chambers and on internal moulds

take the form of crudely elliptical pits 0.3-0.8 mm across and approximately

0.1 mm deep (Figs 81, 82). They may affect the whole of the body chamber as

preserved (Fig. 82) part of the length only (Fig. 81A) or be restricted to parts of

the flanks, dorsum or venter only. Where the Baculites retain traces of shell,

this may be differentially preserved within the pits. In other cases, where traces

of shell extend over the surface of the mould, the pits appear to have been

originally hollow, rather than solid pillars of shell material. That they appear to

have been hollow suggests that they record the presence of some foreign body

between the shell interior and mantle surface that irritated the mantle cells and

led to the secretion of shell material that sealed off the source of irritation and

hence secretory stimulus. These structures do not extend through the shell, and

are thus not a response to external borings through the shell that stimulated

mantle activity to repair damage.

PREDATION

There is an extensive literature on shell damage to ammonites that has been

interpreted as a result of predation, in some cases with subsequent and

successful shell repair. Previous workers have attributed damage to predatory

Fig. 78 (see facing page). Baculites codyensis Reeside, 1927a. Marias River Shale,

Santonian, USGS Mesozoic locality 21425, east bank of Marias River, 18.15 km (11 miles)

south-west of Shelby in W% NE% SE% sec. 14, T. 31 N., R. 4 W., Toole County,

Montana. A-C, F, G-I. Internal moulds of body chambers showing ribbing interrupted by

wrinkles of the type shown in Figs 76-77. A-C. USNM 507268. F. USNM 507269.

G-I. USNM 507270. D-E and G show the dorsal muscle scar, whereas D also shows

longitudinal dorsal grooves, corresponding to ridges on the inside of the shell. All x 1.

Fig. 79 (see overleaf). A-C. Microsculpture wrinkles in Baculites texanus Kennedy &

Cobban, 1999. From the Pierre Shale, Campanian, USGS Mesozoic locality D1411, west-

facing bluff, 1.6 km (1 mile) north-north-east of Oral in NW% NW4% sec. 26, T. 7 S.,

R. 8 E., Fall River County, South Dakota. A-B. USNM507271. C. USNM

507272. D. Feather structure in Baculites scotti, USNM507273, horizon and locality as for

A-C. E. Microsculpture wrinkles in Baculites rugosus Cobban, 1962a, USNM 507274,

Pierre Shale, Campanian, Exiteloceras jenneyi Zone, SE% NE% sec. 17, T. 7S., R. 8 E.,

Fall River County, South Dakota. All x 1.

THE AMMONITE FAMILY BACULITIDAE 103

Fig. 78

104

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 79

THE AMMONITE FAMILY BACULITIDAE 105

A B C

Fig. 80. Baculites eliasi Cobban, 1958. BHI 4362, Pierre Shale, Maastrichtian,

Garfield County, Montana. Blister-like protuberance on venter of partially exfoliated

specimen is interpreted as the result of parasitism, corresponding to the

“Volumensanomalie’ of Keupp (1984, 1994). x 1.

decapods, fish and mosasaurs (see reviews in Kennedy & Cobban 1976; Keupp

1984, etc.). See also Seilacher (1998) for a different interpretation of alleged

mosasaur predation on Placenticeras. The only previous record of predation in

Baculites is by Kauffman (in Kennedy & Cobban 1976: 18) of mosasaur tooth

marks. We have not ourselves seen evidence of such predation, but through the

Fig. 81 (see overleaf). A, C. Baculites scotti Cobban, 1958. USNM 507273, from the Pierre

Shale, Campanian, USGS Mesozoic locality D1411, west-facing bluff 1 mile north-east of

Oral in NW% NW% sec. 26, T. 7S., R. 7 E., Fall River County, South Dakota. Specimen

is uncoated in A and coated with ammonium chloride in C. The specimen has lost most of its

Shell, but traces survive in the pits on the surface of the mould as shown in A. Detailed

inspection suggests that these pits correspond to hollow blisters on the inside of the body

chamber, interpreted as a result of infestation by an unknown parasitic organism.

B. Baculites sp., USNM 507275, from the Pierre Shale, Campanian, 12.2-21.3 m above the

top of the Groat Sandstone at USGS Mesozoic locality D2140, 25.6 km north of Belle

Fourche in SW% SE% sec. 22, T. 11 N., R. 2 E., Butte County, South Dakota. Specimen is

an uncoated internal mould, and shows drag markings associated with the ventral muscle scar

on the phragmocone, and intense pitting, regarded as a response to the same type of

parasitism as shown in A and C. A-B x 1; C x 4.5.

ANNALS OF THE SOUTH AFRICAN MUSEUM

106

Fig. 81

THE AMMONITE FAMILY BACULITIDAE 107

A C

Fig. 82. Baculites sp., USNM 507275, from the Pierre Shale, Campanian, 12.2-21.3 m

above top of Groat Sandstone at USGS Mesozoic locality D2140, 25.6 km north of Belle

Fourche in SW% SE% sec. 22, T. 11 N., R. 2 E., Butte County, Montana. A. Dorsum.

B. Flank. C. Venter. All of an internal mould coated with ammonium chloride, showing

dense pitting, corresponding to blisters on the inside of the shell of the body chamber,

interpreted as a response to infestation by an unknown parasite. x 1.

courtesy of Messrs P. and N. Larson (Black Hills Institute of Geological

Research, Hill City, South Dakota), we have been able to study a number of

spectacular examples of predation on Baculites. The most frequent evidence of

predation is traumatic damage to the body chamber in which the end of the

shell, part or all of the dorsum, venter or flank have been broken-off, followed

by subsequent repair, irregular growth and, eventually a return to normal shell

development (Fig. 83).

In some cases shells retain dents and depressions, as though significant

amounts of tissue had been bitten out and subsequently regenerated. Such

damage affects individuals with whorl height of up to 50 mm, and may possibly

be the result of attacks by pycnodont fish.

A second striking type of healed shell damage that appears to record

predation takes the form of a ‘V’-shaped break to the shell on both flanks

(Fig. 84), tapering away from the adapertural end. The outline of the inferred

ANNALS OF THE SOUTH AFRICAN MUSEUM

108

THE AMMONITE FAMILY BACULITIDAE 109

break is irregular, with one side of the ‘V’ feebly curved towards the apex, the

apex rounded, and the other side of the “V’ with a minor embayment at the

apical end. The healed area may bulge out, or be irregularly impressed and

folded, presumably recording the morphology of the healed tissue that bulged

out of the shell locally. If it is assumed that Baculites lived with the dorso-

ventral plane vertical, damage was clearly produced by a lateral closing

movement of an approximately mirror image pair of structures. Three

possibilities come to mind—a fish that rotated the jaws and head through

90 degrees before striking, a large cephalopod that captured the Baculites and

rotated it before biting, or a decapod that simply snapped at the shell, the

direction of movement of the closing claw being normal to the dorso-ventral

plane of the ammonite. We believe the last to be the most likely culprit on the

basis of the clear asymmetry of the damage, and the minor differences in outline

and size of the opposing flanks of the specimens. Large decapods are known

from the Pierre Shale and equivalent rocks in the Western Interior, and are

described by Whitfield (1907), Feldman et al. (1977), Kammer & Raff (1978),

and others.

A third indicator of possible predation is shown by BHI 4377 (Fig. 85), a

specimen of Baculites texanus Kennedy & Cobban, 1999, from the Campanian

Pierre Shale, Baculites scotti Zone of Lyman County, South Dakota. The

specimen is a phragmocone 53.5 mm long, with a maximum preserved whorl

height of 17 mm. The adapical end of the fragment shows a prominent healed

“V’-shaped area of damage on both flanks, of the type attributed above to

decapod attack. The fragment also shows five prominent circular to elliptical

holes in one flank, and a further hole in the venter. These are borings, and

penetrate through the full thickness of the nacreous layers. The borings are

subcircular, elliptical, tear-shaped or bilobate, with maximum external

diameters of 1.8-3.5 mm. In some cases, a distinct groove leads to the boring,

or a pair of borings may be partially superimposed. There are also suggestions

of incomplete borings with associated, sometimes linking grooves, mere

depressions in the outer surface of the nacre. The borings taper slightly with a

countersunk appearance. These holes look like octopus borings. Closely

comparable borings of undoubted octopus origin are discussed by Bromley

(1993), and others (e.g. Saunders et al. 1987; Saunders et al. 1991) all of whom

provide references to earlier accounts. The present examples are drilled into the

phragmocone, and, in the absence of evidence of damage to the body chamber

Fig. 83 (see facing page). A-D. Baculites gregoryensis Cobban, 1951. BHI collections,

from the Pierre Shale, Campanian, Lyman County, South Dakota. Specimen retains partially

exfoliated shell, and is a section of the phragmocone. The individual appears to have suffered

traumatic damage to the venter and one flank, interpreted as the result of a bite by a

vertebrate, possibly a pycnodont fish. The swollen portion records regeneration and shell

repair by the damaged mantle tissues. E-G. Baculites reesidei Elias, 1933. BHI collections,

from the Pierre Shale, Campanian, Meade County, South Dakota. Specimen retains partially

exfoliated shell and is part phragmocone, part body chamber. The individual has suffered

massive damage in life, with removal of the venter and most of the flanks as a result of a bite

by a vertebrate, possibly a pycnodont fish, followed by extensive regeneration of the shell.

Ils:

110 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 84. Baculites showing healed asymmetric V-shaped bites to both flanks, interpreted as a

result of a decapod attack. A-D. Baculites sp., BHI 4358, Baculites compressus or

B. cuneatus zone, Campanian, Meade County, South Dakota. Specimen is an internal mould

of part of the body chamber. E-H. Baculites gregoryensis Cobban, 1951, BHI 4370, Pierre

Shale, Campanian, Lyman County, South Dakota. Specimen is a phragmocone, with partially

exfoliated shell. x 1.

(which is not preserved), it cannot be shown that this represents attack on a

living animal. Professor R. Bromley (Copenhagen, telefax 07.02.96) has

examined this specimen, and does not think that the holes were made by octopi;

rather he suggests them to be due to diagenetic processes. He has seen similar

holes in aragonite developed by chemical diagenesis around pyrite grains.

111

THE AMMONITE FAMILY BACULITIDAE

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112 ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNOTATED LIST OF SPECIES REFERRED TO THE

FAMILY BACULITIDAE

The most important references for each species are given, but we do not

claim these to be complete synonymies.

The symbol * preceding an entry indicates species figured photographically

herein. For photographic illustrations of most of the type specimens of Baculites

from Madagascar and South Africa, and Eubaculites in general, readers are

referred to Klinger & Kennedy (1993) and Klinger & Kennedy (1997).

The following abbreviations are used to indicate the repositories of material

referred to in the text and appendix:

ANSP Academy of Natural Sciences, Philadelphia

ASUE _ Ain Shams University, Egypt

BEG Bureau of Economic Geology at Austin, Texas

BGS British Geological Survey, Keyworth, Nottingham

BHI Black Hills Institute of Geological Research, Hill City, South Dakota

BMNH_ Natural History Museum, London

CAS California Academy of Sciences (type collection), San Francisco

CBC Commonwealth Bureau of Mineral Resources, Geology and Geophysics, Canberra

CPC Facultad de Ciencias Exactas, Fisicas y Naturales, Universidad Nacional de

Cordoba, Argentina

CTB Collection Klauman

DSGUB Department of Geology, University of Bologna

GD Institut des Sciences de la Terre de l’université Dijon (ex Collignon collection)

GDUA Department Geology, University of Adelaide

GK Department of Geology, Kyushu University, Fukuoka

GLKU Geological Laboratory, Kagawa University, Tamatsu

GMUA_ Geology Museum, University of Alberta

GMUK_ Geology Museum, University of Kansas

GPIB Institut fir Geologie der Friedrich-Wilhelms Universitat, Bonn

GSC Geological Survey, Canada

GSI Geological Survey of India

GSNZ Geological Survey, New Zealand

GT Geological Institute, University of Tokyo

HTW __— Henryk Teisseyre Geological Museum, Wroctaw

HU Hebrew University, Department Geology, Jerusalem

IGG Institute Geology-Geophysics, Bucarest

IGP Instituto di Geologia, Padova

IRSNB Institut Royal des Sciences Naturelles, Brussels

LSJU Stanford University, California

KU Kansas University

MGL Museé Géologique, Lausanne

MLP Museo de Ciencias Naturales de La Plata

MNHP Museum Histoire Naturelle, Paris

MMH Mineralogical Museum, University Copenhagen (Museum Mineralogicum

Hafniense)

NHMW Naturhistorisches Museum, Vienna

NMB National Museum, Bloemfontein (presently in the South African Museum)

OUM Oxford University Museum

SAM South African Museum, Cape Town

SAS Geological Survey of South Africa, Pretoria

SMC Sedgewick Museum, Cambridge

SP Collections Sorbonne, now Université Pierre et Marie Curie, Paris

TMM

UAB

UCLA

UCS

USGS

UPST

USNM

YPB

THE AMMONITE FAMILY BACULITIDAE

Texas Memorial Museum, Austin

Geology Department, Universitat Autonoma, Barcelona

University California, type collection

Geology Department, University Chile, Santiago

United States Geological Survey

Université Paul Sabatier, Toulouse

United States National Museum, Washington

Peabody Museum, Yale

Genus Baculites Lamarck, 1822

Baculites acuminatus Glazunova, 1960

113

Glazunova 1960: 162, pl. 36 (figs 6-7), text-fig. 26.

Type. ?

Occurrence. Asiatic Russia.

Baculites alavaensis Santamaria Zabala, 1996

Santamaria Zabala 1996: 14, pl. 3 (figs 7-9). Klinger & Kichler 1998:

287-295, figs 1, 2a-f, 3-4, 51-1.

Type. Holotype is the original of Santamaria Zabala (1996, pl. 3 (fig. 8))

from the Campanian of Ullibarri-Jauregui, northern Spain. UAB-1979.

Occurrence. Lower Campanian, Alava Province, northern Spain.

* Baculites albertensis Warren, 1930 Fig. 117D-E

(= B. codyensis Reeside, 1927a, fide Kennedy & Cobban 1991a: 72).

Warren 1930: 64, pl. 5 (figs 5-8, 10, 13).

Type. Lectotype here designated is the original of Warren (1930, pl. 5

(figs 7-8)), from the Coniacian Smoky River Shale, Little Smoky River,

Peace River District, Alberta, Canada. GMUA Ct 458.

Occurrence. Coniacian, Canada.

Baculites alonsoi Santamaria Zabala, 1996

Santamaria Zabala 1996: 13, pl. 3 (figs 3-6).

Type. Holotype is the original of Santamaria Zabala (1996, pl. 3 oo 6))

from the Campanian of Ullibarri-Jauregui, northern Spain. UAB-1069

Occurrence. Lower Campanian, Alava Province, northern Spain.

Baculites ambatryensis Collignon, 1971

Collignon 1971: 15, pl. 645 (fig. 2392). Klinger & Kennedy 1997: 128,

fig. 62a. Fatmi & Kennedy 1999: 657, figs 13.1-13.18, 13.23-13.28, 16.1,

16.4, 16.5.

Type. Holotype by monotypy is the original of Collignon (1971, pl. 645

(fig. 2392)) from the Lower Maastrichtian of gisement 400, Céte d’ Ambatry

(Betioky), Madagascar. GD 12392.

Occurrence. Lower Maastrichtian, Madagascar; Upper Maastrichtian,

Baluchistan and India.

114 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 86. Baculites anceps Lamarck, 1822. A-B. Musée d’Histoire Naturelle, Genéve,

unregistered specimen, ex Pictet Collection. C-D. NHMW 7482. Both from the Upper

Maastrichtian Calcaire a Baculites of Valognes, Manche, France. x 1.

Baculites ambiguus Eichwald, 1868

Eichwald 1868: 1176, pl. 37 (fig. 3a-c). = ?B. claviformis Stephenson,

1941 (fide Cobban 1974: 8).

* Baculites anceps Lamarck, 1822 Figs 86-89

Lamarck 1822: 648. Schliiter 1876, non p. 145, pl. 40 (figs 2, 6) only. non

Johnson 1903: 132, pl. 11 (fig. 30a-c). non Miller & Wolleman 1906: 5.

Lopuski 1911: 122, pl. 4 (fig. 4). non Spengler 1923: 55, pl. 4 (fig. 8a, b)

(as B. cf. anceps). ?Basse 1931: 21, pl. 2 (fig. 5) ?Haas 1943: 13,

figs 15-19. ?Mikhailov 1951: 44, pl. 2 (figs 11-12), text-fig. 13. non Hagg

1954: 55. Naidin & Shimanskij 1959: 183, text-fig. 17-20. non Young

1963: 42, pl. 2 (figs 18, 20-22) (as B. cfr. anceps). non Giers 1964: 257,

text-fig. 3. Howarth 1965: 363, pl. 4 (fig. 4), pl. 5 (figs 4-5), pl. 6

(figs 1-5), text-figs 2-3, 5-12. non Atabekian & Khakimov 1976: 94, pl. 2

THE AMMONITE FAMILY BACULITIDAE 1

Cc D

Fig. 87: Baculites anceps Lamarck, 1822. SMC F2822 from the Upper Maastrichtian of

Maastricht, The Netherlands. x 1.

(figs 3-4), pl. 11 (figs 8-10). Matsumoto & Obata 1963: 59. Naidin 1974:

163, pl. 53 (fig. 1), 2non pl. 53 (figs 2-4), text-figs 22.3-22.5, 23a (as

B. anceps anceps hamarck) [sic]. ?Martinez 1982: 169, pl. 30 (figs 1-2)

116

ANNALS OF THE SOUTH AFRICAN MUSEUM

(= ?E. simplex). Balan 1982: 209, pl. 19 (figs 5-7). Kennedy 1986a: 58,

pl. 11 (figs 12-14), pl. 12 (figs 7-11), text-figs 3e-h, 7a-c. Kennedy 1986c:

189, pl. 20 (fig. 2), pl. 28 (figs 3, 11-13, 19-23), text-fig. 11c-d. Kennedy

1986d, fig. 10n—o. Ward & Kennedy 1993: 52, fig. 45.2.

Type. Neotype, designated by Howarth (1965: 365, pl. 5 (fig. 5)) from

‘Normandy’, BMNH C32573.

Occurrence. Upper Maastrichtian, Denmark (as B. valognensis by

Birkelund 1979: 53), Cotentin Peninsula, France, Belgium, Poland,

Commonwealth Independent States, and possibly Spain. At Zumaya (Spain),

B. anceps is recorded from the Lower Maastrichtian (Ward et al. 1991:

1183, fig. 2; Ward & Kennedy 1993: 52); herein also possibly from the

Lower Maastrichtian of Angola.

Fig. 88. Baculites anceps Lamarck, 1822. Muséum Nationale d’Histoire Naturelle

Collections, Paris, ex D’Orbigny collections no. 7204, from the Upper

Maastrichtian of Néhou, Manche, France.

All x 1.

THE AMMONITE FAMILY BACULITIDAE 117

Fig. 89. Baculites anceps Lamarck, 1822. From Piconville, Manche, France.

Lund University Collections. The original of Baculites schliiteri n. sp. of

Moberg (1885, pl. 4 (fig. 13)). x 1.

Baculites anceps var. angustisellata Lopuski, 1911

Lopuski 1911: 124, 139. (= B. anceps s.s. fide Kennedy 1986c: 190).

Type. Lopuski only figured the suture; lectotype designation must await

study of the original material.

Occurrence. Maastrichtian, Poland.

Baculites anceps var. latisellata Lopuski, 1911

Lopuski 1911: 124, 139.

Type. Lopuski only figured the suture; lectotype designation must await

study of the original material.

Occurrence. Maastrichtian, Poland.

Baculites anceps var. leopoliensis Nowak, 1908

See B. leopoliensis.

Nowak 1908: 328, pl. 14 (figs 1-5), text-figs 1-5 (p. 329), text-figs 5-10

(p. 331).

Baculites anceps var. obtusa Meek, 1876

See B. obtusus.

Meek 1876: 406, text-figs 57-60.

118 ANNALS OF THE SOUTH AFRICAN MUSEUM

E F G H

Fig. 90. Baculites anceps pacificus Matsumoto, 1959. A-I. CAS-31245.01, all

from Alameda County, Arroyo del Valle, from creek south of Aqueduct tunnel,

SE% of NW% of sec. 13, T. 3 S., R. 2 E., California.

All x 1.

* Baculites anceps pacificus Matsumoto & Obata, 1963 Fig. 90

(=.B. subanceps)

Matsumoto & Obata 1963: 59, pl. 20 (fig. 3), text-figs 145-146, 156.

Baculites anceps var. sublaevis Griepenkerl, 1889

(= nom. nud. fide Howarth 1965: 366; non anceps, nom. dub. fide Kennedy

1986c: 191).

Baculites anceps var. valognensis Bohm, 1891

See B. valognensis.

Baculites androtsyensis Collignon, 1970

(= B. increscens)

Collignon 1970: 5, pl. 608 (figs 2270-2272). Klinger & Kennedy 1997,

figs 83, 85c.

THE AMMONITE FAMILY BACULITIDAE 119

Type. Holotype by original designation is the original of Collignon (1970,

pl. 608 (fig. 2270)), from the Middle Campanian at Gisement 329, Coupe

Ampolypoly-Antsirasira-Behamotra (Belo sur Tsiribihina), Madagascar.

GD 12270.

Occurrence. Middle Campanian, Madagascar.

* Baculites angustus Moberg, 1885 Fig. 91E-G

Moberg 1885: 39, pl. 4 (fig. 10). Hagg 1954: 55. Kennedy & Christensen

1997: 110, fig. 27, pl. 13 (figs 5-8), text-fig. A-E. Nomen dubium fide

Kennedy & Christensen (1997: 110).

Baculites ankilizatensis Collignon, 1970

Collignon 1970: 13, pl. 612 (figs 2282-2284). Klinger & Kennedy 1997,

figs 85b, 89.

Type. Holotype by original designation is the original of Collignon (1970,

pl. 612 (fig. 2282)), from the Middle Campanian, of Gisement 153 Coupe

d’ Ankilizato (Belo sur Tsiribihina), Madagascar, GD 12282.

Occurrence. Middle Campanian, Madagascar.

?Baculites annulatus Conrad, 1856

Conrad 1856: 265.

Baculites antsirasiraensis Collignon, 1969

(= B. menabensis)

Collignon 1969: 18, pl. 519 (figs 2040-2041). Klinger & Kennedy 1997,

figs 59a—c, 62b.

Type. Holotype by original designation is the original of Collignon (1969,

pl. 519 (fig. 2040)) from the Lower Campanian of Gisement 304, Coupe

Ampolypoly-Antsirasira-Behamotra (Belo sur Tsiribihina), GD 12040.

Occurrence. Lower Campanian, Madagascar.

* Baculites aquilaensis Reeside, 1927b Figs 92-96, 139-140

Reeside 1927b: 12, pl. 6 (figs 11-13), pl. 8 (figs 1-14). Landes 1940: 167.

Young 1963: 41, pl. 1 (figs 1-4, 9) (as B. sp. cfr B. aquilaensis). Cobban &

Kennedy 1992a: 82, pl. 5 (figs 1-6, 10-15), pl. 6 (figs 3-4), pl. 7

(figs 3-4), text-fig. 4d (as Baculites sp. group of aquilaensis). Kennedy

1993: 110, pl. 4 (figs 22-24). Kennedy & Christensen, 1997: 110, fig.

26f-g, h, j-k (as B. cf. aquilaensis. Larson et al. 1997: 20.

Type. Holotype is the original of Reeside (1927), pl. 8 (figs 7-11)), from

the Lower Campanian, sandstone in the Steele Shale, Sheridan County,

Wyoming, USNM 73298.

Occurrence. Lower Campanian, US Western Interior (Wyoming, Montana,

Utah, South Dakota, Colorado, New Mexico), Alberta, Canada;

questionably Middle Campanian, Texas; Campanian, Belgium and perhaps

Sweden.

120 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 91. Baculites sp. A-B. The original of Baculites vertebralis Lamarck of Moberg

(1885, pl. 4 (fig. 9)) from the Campanian of Balsberg, Sweden. C-D. Baculites sp., the

pagal of Baculites vertebralis Lamarck of Moberg (1885, pl. 4 (fig. 8)), from a loose block

at Ahus, Sweden. E-G. Baculites angustus Moberg, 1885, the original of Moberg (1885,

pl. 4 (fig. 10)), from the Campanian of Képinge, Sweden. H. Baculites suecicus Moberg,

1885, the original of Moberg (1885, pl. 4 (fig. 1)), from the Lower Campanian of

Kaseberga, Sweden. All from Lund University Collections. All x 1.

THE AMMONITE FAMILY BACULITIDAE 121

A B C

Fig. 92. Baculites aquilaensis Reeside, 1927b. USNM 73298 from sandstone

in Steele Shale, 900 ft below top, 2 miles north-west of Slack, Sheridan

County, Wyoming. x 1.

Fig. 93 (see overleaf). A-D. Baculites aquilaensis Reeside, 1927b. Collections of the

U.S. Geological Survey, Denver, from USGS Mesozoic locality 23639 lower Gammon Shale

Member of Pierre Shale along west-facing scarp at head of Owl Creek in SE% sec. 12,

T.9S., R. 61 E., and N% sec. 13, T. 9 S., R. 16 E., Carter County, Montana. Lower

Campanian Zone of Scaphites hippocrepis sensu lato.

122 ANNALS OF THE SOUTH AFRICAN MUSEUM

THE AMMONITE FAMILY BACULITIDAE 123

A B

Fig. 94. A-C. Baculites aquilaensis Reeside, 1927b. Collections of the

U.S. Geological Survey, Denver, from USGS Mesozoic locality 23639

lower Gammon Shale Member of Pierre Shale along west-facing scarp at head

of Owl Creek in SE% sec. 12, T. 9 S., R. 61 E., and N% sec. 13, T. 9 S.,

R. 16 E., Carter County, Montana. Lower Campanian Zone of Scaphites

hippocrepis sensu lato.

Fig. 95 (see overleaf). Baculites aquilaensis Reeside, 1927b. Collections of the

U.S. Geological Survey, Denver, from USGS Mesozoic locality 23639 lower Gammon

Shale Member of Pierre Shale along west-facing scarp at head of Owl Creek in SE%

sec. 12, T. 9 S., R. 61 E., and N% sec. 13, T. 9 S., R. 16 E., Carter County,

Montana. Lower Campanian Zone of Scaphites hippocrepis sensu lato. x 1.

Fig. 96 (see overleaf). Baculites sp. gr. of aquilaensis Reeside, 1927).

A. USNM 441411. B-C. USNM 441410. D-E. USNM 441407.

F-G. USNM 441406. All from the Trachyscaphites spiniger zone of the Ozan

Formation, 46 m above the base at USGS Mesozoic locality D10154, Fannin County,

Texas. All x 1.

124 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 95

THE AMMONITE FAMILY BACULITIDAE 125

126 ANNALS OF THE SOUTH AFRICAN MUSEUM

* Baculites aquilaensis var. obesus Reeside, 1927b Fig. 97A

Reeside 1927b: 12, pl. 10 (figs 1-8).

Type. Holotype is the original of Reeside (19275, pl. 10 (figs 1-6)),

from the Lower Campanian, sandstone in the Steele Shale, Wyoming,

USNM 73305.

Occurrence. Lower Campanian, US Western Interior (Wyoming,

Montana, New Mexico).

* Baculites aquilaensis var. separatus Reeside, 1927b Fig. 97E-F, 98

Reeside 1927b: 12, pl. 8 (figs 15-21), pl. 9 (figs 6-15), pl. 45

(figs 5-6). Landes 1940: 168.

Type. Holotype is the original of Reeside (1927), pl. 8 (figs 18-21)

from the Lower Campanian, sandstone in the Steele Shale, Wyoming,

USNM 73302.

Occurrence. Lower Campanian, US Western Interior (Montana,

Wyoming, New Mexico), Alberta, Canada.

Baculites sp. (nov.?) cf. Aquilaensis (non Reeside) Collignon

Collignon 1970: 81, pl. 639 (fig. 2358). Klinger & Kennedy 1997,

fig. 12la-c.

Occurrence. Upper Campanian, Madagascar.

Baculites argentinicus Weaver, 1927

(?= Eubaculites simplex (Kossmat); nom. dub. Eubaculites fide

Riccardi 1974)

Weaver 1927: 429, fig. 2.

Type. Weaver based his description on the figured specimen from “The

Roca beds at locality no. 115 on the western side of Sierra de

Huantraico in the north-western part of Gobernacion del Neuquén,

Argentina’, but also mentioned another poorly preserved specimen

from Los Ramblones. The former is here designated lectotype.

Allegedly housed in the collections of University of Washington,

Seattle.

Occurrence. Upper Campanian or Maastrichtian, Neuquén Province,

Argentina.

Fig. 97 (see facing page). A. Baculites aquilaensis obesus Reeside, 1927b.

USNM 73305 from sandstone in Steele Shale, 1000 ft below top in SW% sec. 19,

T. 58 N., R. 87 W., Sheridan County, Wyoming. B-D. Baculites ovatus var. haresi

Reeside, 1927b. USNM 73296 from Elk Basin sandstone member of Telegraph Creek

Formation in sec. 2, T. 7 S., R. 23. E., Carbon County, Montana. E-F. Baculites

aquilaensis separatus Reeside, 1927b. USNM 73302 from sandstone in Steele Shale,

1040 ft below top, in sec. 32, T. 58 N., R. 87 W., Sheridan County, Wyoming.

H-I. Baculites thomi Reeside, 1927b. Specimen with complete aperture in the U.S.

Geological Survey Collections, Denver. From USGS Mesozoic locality 21419, 8 miles

west of Shelby, Montana, in the NE% sec. 31, T. 32 N., R. 3 W., Marias River

Shale, Kevin Member. x 1.

THE AMMONITE FAMILY BACULITIDAE 127

ANNALS OF THE SOUTH AFRICAN MUSEUM

128

Fig.

THE AMMONITE FAMILY BACULITIDAE 129

* Baculites asper Morton, 1830 Fig. 99H-K

Morton 1830: 291. Morton 1834: 43, pl. 1 (figs 12-13), pl. 13 (fig. 2).

non Roemer 1852: 36, pl. 2 (fig. 2). Johnson 1905: 26. non Reeside

1927a: 4, pl. 1 (figs 19-24), pl. 2 (figs 1-5). non Reeside 1927b: 13

(pars), pl. 11 (figs 5-16). non Picard 1929: 442, pl. 10 (fig. 8).

Reeside 1962: 116 (pars). non Scott & Cobban 1964, pl. 3 (fig. 5),

pl. 4 (fig. 3), pl. 7 (figs 5-6). non Kennedy 1977, text-fig. 17.5-17.6.

non Kauffman 1977: 268, pl. 28 (fig. 6).

Type. According to Reeside (1962: 117), only one of Morton’s

syntypes (Morton 1834, pl. 1 (figs 12-13)) survives in the Academy of

Natural Sciences, Philadelphia, ANSP 19878, allegedly from Prairie

Bluff, Alabama.

Occurrence. Upper Campanian, Selma Chalk, Cahawba, Lower

Maastrichtian. Prairie Bluff. All records of B. asper subsequent to

Morton’s original descriptions seem to be misidentified B. codyensis

(fide Kennedy & Cobban 1991a: 72).

Baculites asper var. larteti Blanckenhorn, 1905

See B. larteti

Blanckenhorn 1905: 111.

*Baculites asperiformis Meek, 1876 Figs 100-101

Meek 1876: 405, pl. 39 (fig. 10a, d). Landes 1940: 168. Cobban

19625: 708, pl. 106 (figs 1-16). Gill & Cobban 1973: 5, fig. 2). Larson

et al. 1997: 23.

Type. Lectotype by subsequent designation of Cobban (1962b: 708) is

the original of Meek (1876, pl. 39 (fig. 10a, d)), from the Middle

Campanian, allegedly from ‘near mouth of Judith River, Montana’ (see

Cobban 1962b: 711 for locality data), USNM 178a.

Occurrence. Middle Campanian, US Western Interior (New Mexico,

South Dakota, Wyoming, Kansas, Montana, Colorado), Alberta and

Saskatchewan, Canada.

Baculites asperoanceps Lasswitz, 1904

Lasswitz 1904: 16, pl. 3 (15) (fig. la-b). ?7Roemer 1852: 36, pl. 2

(fig. 2). non Taubenhaus 1920: 10, pl. 6 (fig. 3) (= B. palestinensis).

Adkins 1928: 206. non Collignon 1938: 89, pl. 6 (fig. 7) (as B. cf

aspero-anceps). Klinger et al. 1996: 100, pi. 1; Klinger & Kennedy,

1997, fig. 129.

Fig. 98 (see facing page). Baculites aquilaensis separatus Reeside, 1927b. Collections

of the U.S. Geological Survey, Denver, from USGS Mesozoic locality 23639 lower

Gammon Shale Member of Pierre Shale along west-facing scarp at head of Owl Creek

in SE% sec. 12, T. 9 S., R. 61 E., and N% sec. 13, T. 9S., R. 16 E., Carter County,

Montana. Lower Campanian Zone of Scaphites hippocrepis sensu lato. x 1.

130 ANNALS OF THE SOUTH AFRICAN MUSEUM

THE AMMONITE FAMILY BACULITIDAE 131

F G H

Fig. 100. A-B. Baculites asperiformis Meek, 1876. USNM 178 from the mouth of

Judith River, Montana; the specimen figured by Meek (1876, pl. 39 (fig. 10a, 10d)).

C-H. Baculites obtusus Meek, 1876. C-E. USNM 1934, from Deer Creek, North

Platte River, Wyoming; the specimen figured by Meek (1876, text-fig. 57).

F-H. USNM 1934 from the same locality, the specimen figured by Meek (1876,

text-figs 59-60). All x 1.

Type. Lectotype by subsequent designation of Klinger et al. (1996,

pl. 1) is the original of Lasswitz (1904, pl. 3 (15) (fig. la—b)) from

Austin, Texas. HTW 3045s (k), MGUWr.

Occurrence. Coniacian-?Santonian, Texas.

Fig. 99 (see facing page). A-C. Baculites meeki Elias, 1933. KU 1702A1, the

holotype, from KU locality 1702-1, Beecher Island Shale Member, Pierre Formation,

2 miles north-west of Beecher Island, Yuma County, Colorado. D. Baculites

pseudovatus var. A. Elias, 1933. KU 1662A1, the holotype, from KU locality 1662-1,

Salt Grass Shale Member (Baculites zone), Pierre Formation, sec. 2, T. 12 S.,

R. 42 W., Wallace County, Kansas. E-G. Baculites reesidei Elias, 1933.

USNM 73304, the holotype, from the Bearpaw Shale, sec. 34, T. 16 N., R. 28 E.,

Fergus County, Montana, the specimen figured by Reeside (1927b, pl. 9 (figs 1-5)).

H-K. Baculites asper Morton, 1830. ANSP 19878, according to data on original label

in Morton’s handwriting ‘one of cotypes . . . Prairie Bluff Alabama’.

All x 1.

132 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 101. Baculites asperiformis Meek, 1876. A-B. USNM 131015e.

C. USNM 131015f. D-F. USNM131015a. All from 40 ft below the top of the Sharon

Springs Member of the Pierre Shale at USGS Mesozoic locality D2951, 1 mile north-

east of McAllaster, Logan County, western Kansas. Copy of Cobban (1962), pl. 106

(figs 12-14, 1-3)). All x 1.

Baculites asperoides Meek & Hayden, 1861

Meek & Hayden 1861: 421. (= nom. nud. fide Cobban 19625: 708).

* Baculites baculus Meek, 1862 Fig. 102

Meek & Hayden 1861: 445. Meek 1876: 397, text-figs 51-52. Scott &

Cobban 1965: 3, map I-439. Gill & Cobban 1973: 10, fig. 3d, 7a.

Riccardi 1983, pl. 26 (figs 6-10). Kennedy 1993: 110, pl. 4 (figs 10,

20-21). Larson et al. 1997: 34.

Type. Holotype is from the ‘Fox Hill Sandstone near Glenrock,

Wyoming’ (Meek 1876, text-figs 51-52).

Fig. 102 (see facing page). Baculites baculus Meek, 1862. U.S. Geological Survey

Collections, Denver. From the Pierre Shale near Midland, South Dakota. x 1.

THE AMMONITE FAMILY BACULITIDAE

133

Fig. 102

134 ANNALS OF THE SOUTH AFRICAN MUSEUM

Occurrence. Maastrichtian, US Western Interior (Wyoming, Colorado,

South Dakota, Montana), Saskatchewan, Alberta, Canada, U.S. Gulf

Coast and Belgium.

Baculites bailyi Woods, 1906

Baily 1855: 457 (pars), pl. 11 (fig. 5a—b only, non 5c). ?Etheridge

1904: 90. Woods 1906: 341, pl. 44 (fig. 5). ?Van Hoepen 1921: 18,

pl. 3 (figs 9-10). non Spath 1921: 261. Spath 1922: 146. non Besairie

1930: 223, pl. 21 (figs 6-7). Glazunova 1955: 186, pl. 3 (fig. 7) (as B.

cf. bailyi). Matsumoto & Obata 1963: 35, pl. 20 (figs 1-2), pl. 21

(fig. 5), text-figs 88-89, 116-120, 140-142. Collignon 1969: 21,

pl. 520 (fig. 2051). Klinger & Kennedy 1977: 75, fig. Sd. Ward 1978:

1148, pl. 1 (figs 5-7), text-fig. 5. Olivero 1984: 57, pl. 1 (figs 1-5),

text-fig. la—b. Klinger 1985: 5, fig. 4e-h. Klinger & Kennedy 1997,

figs 13-23, 67k-q, 78a-b.

Type. Holotype by original designation of Woods (1906: 342) is the

original of Baily (1855, pl. 11 (fig. 5a—b)), from an unspecified horizon

in the Mzamba Formation at the Mzamba River Estuary,

BMNH C11372..

Occurrence. Upper Santonian—Lower Campanian, Pondoland,

Zululand, Natal offshore, Madagascar, James Ross Island, Antarctica,

British Columbia.

Baculites bassei Besairie, 1930

Besairie 1930: 222, pl. 22 (fig. 8, 8a), text-fig. 17. Klinger & Kennedy

1997, fig. 121d-e.

Type. Holotype, by monotypy? is the original of Besairie (1930, pl. 22

(fig. 8, 8a)) from the Upper Campanian or Lower Maastrichtian,

Maintirano, Madagascar. The repository of the type is not known to

us.

Occurrence. Upper Campanian or Lower Maastrichtian, Madagascar.

Baculites besairiei Collignon, 1931

(= B. yokoyamai Tokunaga & Shimizu)

Collignon 1931: 37, pl. 5 (figs 6-9), pl. 9 (fig. 16). Collignon 1965:

18, pl. 420 (figs 1745-1746).

Type. Lectotype, here designated is the original of Collignon (1931,

pl. 5 (fig. 6)) from the Upper Coniacian of Mahagaga, Madagascar,

GD unregistered.

Occurrence. Upper Coniacian, Madagascar.

Fig. 103 (see facing page). A-L, R-S. Baculites sp. NHMW 1890. xiii.209, labelled

Baculites faujasi Lam. var. bohem. Frit, Priesener Schichten, Priesen, Bohemia.

Probably Coniacian. T-W. Baculites sp., NHMW 1890.xiii.210, from the same locality

and horizon as A-L, R-S. (= B. brevicosta Schliiter). M-P. Sciponoceras bohemicum

(Fritsch, 1872). NHMW 1893.111.10. The original of Jahn (1895, pl. 8 (fig. 7)).

All x 1.

THE AMMONITE FAMILY BACULITIDAE 135

Fig. 103

136 ANNALS OF THE SOUTH AFRICAN MUSEUM

* Baculites borealis Warren, 1930 Fig. 117A-C

(= B. codyensis Reeside, 1927a, fide Kennedy & Cobban 1991a: 72)

Warren 1930: 65, pl. 5 (figs 3-4, 9, 14).

Type. Lectotype here designated is the original of Warren (1930, pl. 5

(figs 3-4), from the Smoky River Shale, Little Smoky River, Alberta,

Canada. GMUA Ct 465.

Occurrence. Middle Coniacian to Middle Santonian, Alberta, Canada.

* Baculites boulei Collignon, 1931

(= B. capensis)

Collignon 1931: 35, pl. 5 (fig. 2), pl. 9 (fig. 14). non Collignon 1938:

88, pl. 6 (fig. 6, 6a, 6b). Matsumoto 1959: 118, pl. 32 (fig. 7a-c),

pl. 33 (fig. 4a-c, 5a-b, 6a-d, 7a-b), text-figs 27a—b, 28-32.

Matsumoto & Obata 1963: 43, pl. 13 (figs 3, 5), pl. 15 (fig. 6), text-

figs 93, 152-155. non Forster 1975: 168, pl. 4 (figs 3, 9), text-fig. 37.

?non Haggart & Ward 1989: 226, fig. 3.7-3.10 (as Baculites cf.

boulei).

Type. Lectotype by the subsequent designation of Matsumoto (1959:

118) is the original of Collignon (1931, pl. 5 (fig. 2, 2a), pl. 9 (fig. 14))

from the Upper Coniacian of Mahagaga, Madagascar, GD

unregistered.

Occurrence. Upper Coniacian—Lower Santonian Madagascar,

Zululand, California, Hokkaido. (Neal Larson informed us that

Baculites cf. boulei occurs in the Turner Sand Member of the Carlile

Formation from South Dakota, Wyoming, and questionably New

Mexico).

* Baculites brevicosta Schliiter, 1876 ?Figs 103R-S, 128Q-S

Schliiter 1876: 141, pl. 39 (figs 9-10). non Moberg 1885: 37, pl. 4

(figs 5-6). Wegner 1905: 207. non Spath 1921: 260, pl. 24 (fig. 5) (as

B. cf. brevicosta). non Hagg 1930: 58. non Collignon 1931: 34, pl. 5

Fig. 104 (see facing page). A-C. Baculites schencki Matsumoto, 1959.

CAS-67731.01, paratype from Fresno County, Panoche Quadrangle, c. 1100 feet of

south-east corner of sec. 28, T. 14 S., R. 11 E., Panoche Hills, California.

D. Baculites aff. B. capensis Woods, 1906. CAS-31289.01; from Yolo County,

3 miles north-west of Rumsey on Lake Co. highway, on Cache Creek, a little above the

concrete bridge, California. The original figured by Anderson (1958: 192, pl. 48

(fig. 8, 8a)). E-G. Baculites buttensis Anderson, 1958. CAS-27835.01, the holotype,

from Butte County, from bluff on south-east side of Big Chico Creek at Frank

Mickey’s place, California. H-K. Baculites subcircularis Anderson, 1958.

CAS-28442.01, the holotype, from Contra Costa County, 4 miles south of Brentwood,

Marsh Creek, at the mouth of Briones Creek, c. 1250 feet south-west of the old John

Marsh House, from large concretion, S42 SW'% NW% sec. 35, T. 1 N., R. 2 E.

L-N. Baculites lomaensis Anderson, 1958. CAS-2361.03, a paratype, from Fresno

County, north-west of Oil City, 11 miles north of Coalinga, Joaquin Ridge, California,

the specimen figured by Anderson (1958, pl. 48 (fig. 6)). A-G, J-N x 1; H-I x 2.

THE AMMONITE FAMILY BACULITIDAE 137

Fig. 104

138

ANNALS OF THE SOUTH AFRICAN MUSEUM

(fig. 1), pl. 9 (fig. 13) (as B. cf. brevicosta). Kennedy 1984: 146,

pl. 33 (figs 23-25) (as B. cf. brevicosta). Kaplan & Kennedy 1994: 59,

pl. 40 (figs 15-19). ?Kennedy & Christensen 1997: 109.

Type. Lectotype by the subsequent designation of Kennedy (1984:

146), is the original of Schliiter (1876, pl. 39 (figs 9-10)) from the

Emscher Marls of Horst in Westphalia, Germany. The lectotype could

not be located by us in the collections in Bonn and has to be assumed

lost.

Occurrence. Coniacian of Germany and possibly France.

*Baculites buttensis Anderson, 1958 Fig. 104E-G

(= uninterpretable; probably B. capensis; see also Matsumoto 1959:

121-2)

Anderson 1958: 191, pl. 49 (fig. 6).

Type. Holotype (by monotypy?) is the original of Anderson (1958,

pl. 49 (fig. 6)) from the Upper Cretaceous of Chico Creek, California,

CAS type collection.

Occurrence. .?Santonian, California.

* Baculites calamus Morrow, 1935 Fig. 105A

Morrow 1935: 473, pl. 49 (fig. 8a, b). Cobban & Scott 1972: 49,

pl. 34 (fig. 7).

Type. Holotype is the original of Morrow (1935, pl. 49 (fig. 8a, b))

from the Upper Turonian, Bridge Creek Member of the Greenhorn

Formation, Hamilton County, Kansas. GMUK.

Occurrence. Upper Turonian, Kansas and Colorado.

Baculites capensis Woods, 1906

Woods 1906: 342, pl. 44 (figs 6-7). ?Boule et al. 1907: 65 (45), pl. 15

(fig. 3, 3a) (as B. vagina). Crick 1907: 240 (as Baculites sp.). Spath

1921: 257, pl. 24 (figs 6-7). Spath 1921: 259, pl. 24 (fig. 4, 4a) (as B.

cf. aspero-anceps); 1921: 260, pl. 24 (fig. 5, 5a) (as B. cf. brevicosta);

1921: 258 (as B. sp. aff. capensis); 1921: 260 (as B. cf. sulcatus).

Spath 1922: 146. ?Spath 1925: 31, pl. 1 (fig. 1) (as Baculites sp. ind.).

Collignon 1931: 22, pl. 3 (fig. 6) (as B. aff. capensis). Besairie 1930:

620. Venzo 1936: 116 (58). Venzo 1936: 116 (58), pl. 10 (6) (fig. 13)

(as B. capensis var. umsinenensis). ?Anderson 1958: 191, pl. 49

Fig. 105 (see facing page). A. Baculites calamus Morrow, 1935. From the Middle

Turonian Bridge Creek Member of the Greenhorn Formation, Hamilton County,

Kansas. B-R. Baculites yokoyamai Tokunaga & Shimizu, 1926. B-E. USNM 433862

from USGS Mesozoic locality 20611. F-G. USNM 433867 from USGS Mesozoic

locality 23438. H-I. USNM 433858 from the same locality as F-G.

J-L. USNM 433861, from the same locality. M-O. USNM 438857, from the same

locality. P-R. USNM 433863, from the same locality. All from the Lower Coniacian

Inoceramus erectus zone. All x 1.

THE AMMONITE FAMILY BACULITIDAE 139

140 ANNALS OF THE SOUTH AFRICAN MUSEUM

(fig. 6, 6a, 6b) (as B. buttensis). Anderson 1958: 192, pl. 48 (fig. 8, 8a) (as

B. aff. capensis). Matsumoto 1959: 121, pl. 33 (fig. la-d, 2a—c, 3a-b),

pl. 45 (fig. la-d, 2a-d, 3a-d, 4a-d), text-figs 33a—b, 34a—-b. Matsumoto &

Obata 1963: 47, pl. 14 (fig. 2), pl. 15 (figs 3-5), pl. 19 (fig. 2), text-

figs 95-96, 147-151. Collignon 1966: 6, pl. 457 (fig. 1862). Collignon

1966: 6, pl. 457 (figs 1863-1864) (as B. capensis var. tenuetuberculata).

Collignon 1966: 7, pl. 457 (fig. 1865) (as B. malagasyensis). non Kennedy

& Klinger 1973: 100, pl. 4 (figs 1-5), pl. 5 (fig. la—-d), pl. 6 (figs 4-5) (as

Baculites sp. group of B. capensis Woods). Klinger & Kennedy 1977: 71,

figs 2a-f, 3g. Haggart 1984: 233, fig. a-d. ?Cooper 1988: 210, fig. 1g-1.

Kennedy & Cobban 1991b: 182, figs 6: 4; 8: 1-8; 10: 7-10, 12-14; 12: 2,

5. ?Kennedy & Christensen 1993: 152, fig. 4a (as Baculites sp. group of

capensis). Klinger & Kennedy 1997: 53, figs 12m-o, 27-55.

Type. Lectotype by subsequent designation of Matsumoto & Obata (1963:

48) is the original of Woods (1906, pl. 44 (fig. 6a—b)) from the Santonian of

the Mzamba Formation, Mzamba River Estuary, Pondoland, SAM-48723.

Occurrence. Middle Coniacian to Middle Santonian, Pondoland, Zululand,

Madagascar, Santonian California and Hokkaido, Texas and Mississippi in

the USA, doubtfully Campanian of Angola and Sweden.

Baculites capensis var. tenuetuberculata Collignon, 1966

(= B. capensis s.s.)

Collignon 1966: 6, pl. 457 (figs 1863-1864), p. 22, pl. 463

(figs 1894-1895).

Type. Holotype, by original designation is the original of Collignon (1966,

pl. 457 (fig. 1863)) from the Lower Santonian of Gisement 270,

Coupe Ampolypoly-Antsirasira-Behamotra (Belo sur Tsiribihina),

Madagascar, GD 11865.

Occurrence. Lower Santonian, Zone of Texanites oliveti, Madagascar.

Baculites capensis var. umsinenensis Venzo, 1936

(= B. capensis Ss.s.)

Venzo 1936: 116 (58), pl. 10 (6) (fig. 13).

Type. Lectotype, here designated is the original of Venzo (1936, pl. 10

(fig. 13)) from the Upper Coniacian-or Lower Santonian, Mzinene River,

Zululand.

Occurrence. Upper Coniacian or Lower Santonian, Zululand.

Baculites carinatus Morton, 1834

(see Eubaculites)

Morton 1834: 44, pl. 13 (fig. 1).

Fig. 106 (see facing page). Baculites chicoensis Trask, 1856. CAS-61997.08 (Stanford

University collection) from Butte County, east bank of Chico Creek, 3.6 miles along country

road to Big Chico Creek from Humboldt Road at the ‘10 mile House’, California. The

specimen figured by Matsumoto (1959: 146, pl. 37 (fig. la-d), text-fig. 59a-d). x 1.

141

THE AMMONITE FAMILY BACULITIDAE

106

Fig.

142 ANNALS OF THE SOUTH AFRICAN MUSEUM

Baculites carinatus Binckhorst, 1861, non Morton, 1834

(= presumably an atypical B. vertebralis fide Kennedy 1986c: 192)

Binckhorst 1861: 43, pl. 5d (fig. 2).

Type. Holotype by monotypy is the original of Binckhorst (1861, pl. 5d

(fig. 2a—d).

Occurrence. Maastrichtian, Belgium.

Baculites cazadorianus Paulcke, 1907

(= ?B. anceps or Eubaculites simplex)

Paulcke 1907: 11 (177), pl. 16 (fig. 5), text-fig. 2. (as B. vagina var. nov.

cazadorianus).

Type. Lectotype here designated is the original of Paulcke (1907, pl. 16

(fig. Sa-c)) from the Upper Campanian? of Cerro Cazador, Patagonia,

Argentina. The type, originally housed in the Albert-Ludwigs Universitat,

Freiburg im Breisgau, Germany, appears to be lost (letter Dr U. Leppig

13.07.1992).

Occurrence. Upper Campanian?, Patagonia.

* Baculites chicoensis Trask, 1856 Figs 106, 107G-I, 108D-F

Trask 1856: 92, pl. 2 (fig. 2). Gabb 1864: 80, pl. 14 (fig. 27), pl. 17

(fig. 27, 27a). Usher 1952: 96, pl. 26 (figs 1-4), pl. 31 (fig. 18), text-fig. 3.

Matsumoto 1959: 145, pl. 36 (fig. 2), pl. 37 (fig. 1), text-figs 59a-d, 60a,

6la—b, 62a—-b, 63a-b. Matsumoto & Obata 1963: 66, pl. 21 (figs 2, 4), text-

figs 159, 163-164. Ward 1978: 1148, pl. 1 (figs 3-4, 8-9), text-fig. 5.

Matsumoto & Miyauchi 1984: 69. ?Grabovskaja 1984: 86 (as B. cf.

chicoensis). Haggart 1984: 233, fig. 7e-h. Haggart 1991, pl. 2 (figs 6-7).

Type. Trask’s original material is lost (see Taff et al. 1940: 1321,

Matsumoto 1959: 145). Taff et al. 1940 designated two ‘neosyntypes’.

Matsumoto (1959: 146) has selected a better specimen in the collections of

Stanford University, LSJU 8537 (Matsumoto 1959, pl. 36 (fig. 2a—d), text-

fig. 60a—b) from Chico Creek, and indicated that an application would be

made to the International Commission of Zoological Nomenclature to

designate the latter as the neotype, and to invalidate the designation of Taff

et al. 1940.

Occurrence. Lower Campanian, California (see Ward 1978: 418), British

Columbia, Hokkaido, and tentatively southern Saghalin.

Fig. 107 (see facing page). A-F. Baculites occidentalis Meek, 1862.

A-B. CAS-66773.02 from Fresno County, Coalinga Quadrangle from c. 50 ft above

contact with massive sandstone, west of Los Gatos Creek, California. C-F. CAS-

66769.01 from Fresno County, Los Gatos Creek, from the SE% of sec. 4, T. 20S., R.

14 E., from the upper part of the Panoche Formation, California. G-I. Baculites

chicoensis Trask, 1856. CAS-61997.11; from Butte County, east bank of Chico Creek,

3.6 miles along country road to Big Chico Creek from Humboldt Road at the ‘10 mile

House’, California. All x 1.

143

THE AMMONITE FAMILY BACULITIDAE

107

Fig.

ANNALS OF THE SOUTH AFRICAN MUSEUM

144

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l H A a q

THE AMMONITE FAMILY BACULITIDAE 145

Baculites chicoensis Waring, 1915, non Trask, 1856

(= uninterpretable)

Waring 1915, map, fig. 6. See also Matsumoto 1959: 130.

Occurrence. Upper Campanian, Baha California, Mexico.

Baculites chicoensis yezoensis Matsumoto & Miyauchi, 1984

Matsumoto & Miyauchi 1984: 70, pl. 25 (figs 1-5), text-fig. 11b-c.

Fig. 109. Baculites claviformis Stephenson, 1941. A. USNM 44940.

B. USNM 44943. C-D. USNM 449431. All from the Upper Campanian,

Nostoceras hyatti zone fauna of the Coon Creek Tongue of the Ripley

Formation, Coon Creek, McNairy County, Tennessee.

All x 1.

146

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 110. Baculites clinolobatus Elias, 1933. USNM

507276, from the Pierre Shale near Wasta, South Dakota.

pe. Holotype is the original of Matsumoto & Miyauchi 1984, pl. 25

(fig. 1)), from the lower Campanian Fukiyose Bed of Kiyohama II of

northern Hokkaido, GK H5975.

Occurrence. Lower Campanian, northern Hokkaido.

THE AMMONITE FAMILY BACULITIDAE 147

Fig. 111. Baculites clinolobatus Elias, 1933. USNM 507276, from the Pierre

Shale near Wasta, South Dakota. x 1.

148 ANNALS OF THE SOUTH AFRICAN MUSEUM

* Baculites claviformis Stephenson, 1941 Fig. 109

Stephenson 1941: 403, pl. 1, pl. 77 (figs 6-8), pl. 78 (figs 1-6). Cobban

1974: 5, pl. 3 (figs 7-8, 12-14). Cobban & Kennedy 1991b: E4, pl. 3

(figs 1-9). Cobban & Kennedy 1994c: B8, pl. 8 (figs 1-8), pl. 11, text-

fig. 3.

Type. Holotype is the original of Stephenson (1941, pl. 77 (figs 6-8)) from

the Upper Campanian, Nacatoch Sand Kaufman County, Texas,

USNM 77241.

Occurrence. Upper Campanian-?Maastrichtian, Texas, New Jersey,

Arkansas, Alabama, Mississippi, Tennessee, and possibly Colorado.

Fig. 112. Baculites sp. (smooth). A-C. USNM 131014a. D. USNM 131014b. Both

from the Pierre Shale, USGS Mesozoic locality D2102 near Red Bird, Wyoming.

Copy of Cobban (1962), pl. 108 (figs 1-4)). Both x 1.

* Baculites clinolobatus Elias, 1933 Figs 110-111

Elias 1933: 310, pl. 30 (figs 1-2), pl. 34 (figs 1, 2a—-b, 3). Scott & Cobban

1965: 3, map I-439. Gill & Cobban 1973: 11, fig. 7c. Larson et al. 1997:

36.

Type. Lectotype, here designated is the original of Elias (1933, pl. 30

(fig: 1)) from Beecher Island, Yuma County, Colorado. GMUK 59709.

Occurrence. Maastrichtian, US Western Interior (Wyoming, Colorado,

South Dakota, Kansas and Nebraska).

Baculites coagmentatus Collignon, 1970

(= ?B. increscens)

Collignon 1970: 7, pl. 609 (figs 2275-2276). Klinger & Kennedy 1997: 136,

fig. 88.

THE AMMONITE FAMILY BACULITIDAE 149

Type. Holotype is the original of Collignon (1970, pl. 609 (fig. 2275)) from

the Middle Campanian of Gisement 177, Coupe d’Ankilizato (Belo sur

Tsiribihina), Madagascar, GD 12275.

Occurrence. Middle Campanian, Zone of Pachydiscus grossouvrei, top of

subzone of Pachydiscus bassae, Madagascar.

* Baculites cobbani Khakimov, 1976 Fig. 112

Cobban 1962b: 714, pl. 108 (figs 1-4), text-fig. li-j (as Baculites smooth

species). Atabekian & Khakimov 1976: 98, pl. 11 (figs 2-7). Kennedy 1993:

111, pl. 5 (figs 1-12), pl. 6 (figs 5-10), text-fig. 6 (as Baculites sp.

(smooth). Larson et al. (1997), however, think that the Central Asian

material is different from that of the US Western Interior; Dr W. A. Cobban

(pers. comm.) agrees that the sutures are different.

Type. According to Khakimov (1976: 99, in Atabekian & Khakimovy), the

holotype is number 45/1168; however, none of their figured specimens

bears this number.

Occurrence. Lower Campanian, Central Asia; 7Middle Campanian, U.S.

Western Interior; ?Campanian, Belgium.

* Baculites codyensis Reeside, 1927 Figs 113-117

Reeside 1927a: 4, pl. 2 (figs 6-19). Reeside 19275: 13, pl. 10 (figs 9-12),

pl. 11 (figs 5-16) (as B. asper). Reeside 1927a: 4 ,pl. 1 (figs 19-24), pl. 2

(figs 1-5) (as B. asper). Warren 1930: 64, pl. 5 (figs 5-8), 10, 13) (as

B. albertensis). Warren 1930: 65, pl. 5 (figs 3-4, 9, 14) (as B. borealis).

Warren 1930: 65, pl. 5 (figs 1-2, 11-12) (as B. trifidilobatus). Cobban

1955: 204, pl. 2 (fig. 2). Scott & Cobban 1964, pl. 5 (fig. 4), pl. 7

(figs 3-4). Scott & Cobban 1964, pl. 3 (fig. 5), pl. 5 (fig. 3), pl. 7 (figs 5-6)

(as B. asper). Birkelund 1965: 52, pl. 6 (figs 3-5), pl. 7 (fig. 1), text-

figs 42-45. Cobban 1976: 4, pl. 2 (figs 6-19). Kennedy & Cobban 1976,

pl. 2 (fig. 1). Kennedy 1977, text-fig. 17.3-17.4. Kennedy 1977, text-

fig. 17.5-17.6 (as B. asper). Kauffman 1977: 268, pl. 28 (fig. 7). Scott er

al. 1986, figs 12d, 13f. Kennedy & Cobban 1991a: 72, pl. 15 (figs 1-30),

pl. 16 (figs 1-13), pl. 17 (figs 1-8), text-fig. 25f.

Type. Holotype is the original of Reeside (1927a, pl. 2 (figs 6-10)) from the

Coniacian Cody Shale, Park County, Wyoming. YPB 6408.

Occurrence. Middle Coniacian to Middle Santonian, predominantly in the

western half of the US Western Interior, from north-western Alberta in the

north to central New Mexico in the south, Middle Santonian, West

Greenland.

Baculites collignoni Klinger & Kennedy, 1997

B. leopoliensis Collignon non Nowak in Collignon (1970: 10, pl. 610

(figs 2277-2278)) (= ?B. increscens)

Klinger & Kennedy 1997, fig. 86.

ANNALS OF THE SOUTH AFRICAN MUSEUM

150

Fig. 113

THE AMMONITE FAMILY BACULITIDAE 151

Type. Holotype is the original of Collignon (1970, pl. 610 (fig. 2277)) from

the Middle Campanian of Gisement 156, Coupe d’Ankilizato (Belo sur

Tsiribihina), Madagascar, GD 12277.

Occurrence. Middle Campanian, Madagascar.

Baculites columna Morton, 1834

= Trachybaculites columna (Morton, 1834).

Baculites complex Say

Matsumoto 1959: 112. (= ?lapsus B. compressus).

*Baculites compressus Say, 1821 Fig. 118

Say, 1821: 41. Morton 1834: 43, pl. 9 (fig. 1). Meek 1876: 400, pl. 20

(fig. 3a—c), text-figs 55-56. Brown 1892, pl. 9 (figs 1-11). Johnson 1905:

26. Dowling 1917: 31, pl. 30 (fig. 1). non Reeside 1927b: 10, pl. 9

Fig. 114. Baculites codyensis Reeside, 1927a.

A-C. USNM 433895. D-F. USNM 433896, both from the Coniacian, USGS

Mesozoic locality 21097, of the Cody Shale, shore of Ray Lake, Fremont

County, Wyoming. Both x 1.

Fig. 113 (see facing page). Baculites codyensis Reeside, 1927a. A. USNM 433886 from the

Coniacian, USGS Mesozoic locality D3797, of the Cody Shale, Oregon Basin, Park County,

Wyoming. B-D. USNM 433984 from the Coniacian, USGS Mesozoic locality 21097, of the

Cody Shale, shore of Ray Lake, Fremont County, Wyoming. E-F. USNM 433890 from the

Coniacian, USGS Mesozoic locality D1655, of the Cody Shale, Hot Springs County,

Wyoming. All x 1.

ANNALS OF THE SOUTH AFRICAN MUSEUM

152

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wo WOg

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q V

153

THE AMMONITE FAMILY BACULITIDAE

"T x Wog

"euRUOW ‘AJUNOD IOORIH ‘a[eYS JOATY Seley 9y) JO ‘OpEd Ajl[esdo] DIOZOSeW] SOS jo ueeluoD

ay) WOT OO6EEP WNSN “A-d ‘“surwohm ‘AjunoD yeg ‘opeys ApoD ay) Jo “yet Ait[eoo] ofozosayy

sosn

UBTORIUOTD 94} WOl} 868ECh WNSM O-V

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154 ANNALS OF THE SOUTH AFRICAN MUSEUM

F G H

Fig. 117. Baculites codyensis Reeside, 1927a. A-C. GMUA Ct465, a specimen

labelled as ‘Baculites borealis, topotype used as cotype’ from the Middle

Coniacian to Middle Santonian of the Smoky River Shale, Little Smoky River,

Alberta, Canada. D-E. GMUA Ct459, labelled ‘topotype, Baculites albertensis

Warren’ from the same locality. F-H. GMUACt448, data as for D-E.

I-K. GMUA Ct 464, specimen labelled ‘Baculites trifidilobatus topotype used as

cotype’. Locality as for A-C.

All x 1.

(figs 1-5) (= B. compressus robinsoni, W. A. Cobban pers. comm.).

Warren 1931, pl. 2 (figs 1-2). Elias 1933: 299. Elias 1933: 300, pl. 28

(fig. 4), pl. 32 (figs 3a-b, 4a-c, 5a—b). Landes 1940: 172. Gill & Cobban

1973: 10, fig. 2n. Nelson 1975, pl. 65 (figs 3-4). Cobban et al. 1992: A7,

pl. 2 (figs 1-12), pl. 3 (fig. 1) (with complete synonymy). Larson et al.

1997: 29.

Type. According to Elias (1933: 294) and Landes (1940: 172) the types are

lost. Neotype designation is desirable.

Occurrence. Upper Campanian, US Western Interior.

THE AMMONITE FAMILY BACULITIDAE 155

Baculites compressus var. corrugatus Elias, 1933

(= B. corrugatus Elias, 1933)

Elias 1933: 303, pl. 28 (fig. 3), pl. 30 (fig. 3), pl. 32 (fig. la-c).

Baculites compressus var. ornatus Robinson, 1945

(= B. compressus robinsoni Cobban, 1962a)

Robinson 1945: 51, pl. 1 (figs 1-4). Cobban 1962a: 128.

Baculites compressus var. reesidei Elias, 1933

(= B. reesidei Elias)

Elias 1933: 302, pl. 28 (fig. 1), pl. 31 (fig. 3), pl. 32 (fig. 2a-c), pl. 33

(figs la-c, 2a-c).

* Baculites compressus robinsoni Cobban, 1962a Fig. 119

Robinson 1945: 51, pl. 1 (figs 1-4) (as B. compressus var. ornatus).

Cobban 1962a: 128 nom. nov. Larson et al. 1997: 30: (as var. robinsoni).

Jerzykiewicz 1996: 99 (fig. 2).

Type. Holotype is the original of Robinson (1945, pl. 1 (figs 1-2)), from the

Bearpaw Formation of southern Saskatchewan, Canada, GSC 9070.

Occurrence. Upper Campanian, Canada (Alberta), US Western Interior

(Montana, South Dakota).

B | Cc

Fig. 118. Baculites compressus compressus Say, 1821. USNM 507277, from

USGS Mesozoic locality 23349, Pierre Shale, near Wasta, South Dakota.

x1.

156 ANNALS OF THE SOUTH AFRICAN MUSEUM

A B Cc

Fig. 119. Baculites compressus robinsoni Cobban, 1962a. USNM 507278, from

USGS Mesozoic locality D4133, Bearpaw Shale, near Lavina, Montana. x 1.

* Baculites corrugatus Elias, 1933 Figs 120-121

Elias 1933: 303, pl. 28 (fig. 3), pl. 30 (fig. 3), pl. 32 (fig. 1) (as B. com-

pressus var. corrugatus). Cobban 1962a: 134. Larson et al. 1997: 28.

Type. Holotype is the original of Elias (1933, pl. 28 (fig. 3)), from the

Campanian Lake Creek Shale Member, Pierre Shale, Wallace County,

Kansas. GMUK 50164.

Occurrence. Upper Campanian, US Western Interior.

Baculites crickmayi Williams, 1930

Williams 1930: 3, pl. 1 (fig. 1), pl. 2 (figs 1-3). Landes 1940: 173;

Kennedy & Cobban 1993a: 95, figs 10.10, 10.13-10.15, 10.17,

10.20-10.27; 12.5. Larson et al. 1997: 27.

Type. Lectotype here designated is the original of Williams (1930, pl. 1,

pl. 2 (figs 1-3)) from the Upper Campanian, Bearpaw Formation of Alberta.

No repository data were given.

THE AMMONITE FAMILY BACULITIDAE 157

Fig. 120. Baculites corrugatus Elias, 1933. OUM KT7445, from the Pierre Shale,

Didymoceras cheyennense zone, Upper Campanian, Indian Creek in SW% sec. 22,

R. 12 E., T. 3 S, Pennington County, South Dakota. x 1.

Occurrence. Upper Campanian, Bearpaw Formation, Alberta, Canada,

Campanian, US Western Interior (Arkansas, Montana, Wyoming and

Colorado).

* Baculites cuneatus Cobban, 1962a Fig. 122

Meek 1876, text-figs 55-56 (as B. compressus). Warren 1930, pl. 2

(figs 1-2) (as B. compressus). Cobban 1962a: 127, pl. 25 (figs 1-8),

text-fig. 1b. Scott & Cobban 1965: 2, map I-439. Gill & Cobban 1973: 10,

fig. 20. Riccardi 1983, pl. 25 (figs 1-3). Larson et al. 1997: 31.

158 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 121. Baculites corrugatus Elias, 1933. KU 1566A1, the holotype, from upper

part of Lake Creek Shale Member, Pierre Formation, NW SW'% sec. 5, T. 15S.,

R. 41 W., Wallace County, Kansas. x 1.

Type. Holotype is the original of Cobban (1962a, pl. 25 (figs 6-8)) from the

Upper Campanian, near Hardin, Montana, USNM 108966.

Occurrence. Upper Campanian, US Western Interior (Montana,

South Dakota, Colorado, Kansas, Wyoming), Canada (Alberta,

Saskatchewan).

Baculites delvallei Riccardi, 1980

Riccardi 1980: 324, pl. 1 (figs 1-2), text-fig. 1.

THE AMMONITE FAMILY BACULITIDAE 159

Fig. 122. Baculites cuneatus Cobban, 1962a. A, C-D. USNM 108966, the holotype.

B. USNM 108967a, paratype. Both from the Upper Campanian Baculites cuneatus zone of

the Bearpaw Shale, from USGS Mesozoic locality 21363 near Hardin, Montana. Copy of

Cobban (1962a, pl. 25 (figs 4, 6-8)). Both x 1.

Type. Holotype by monotypy is the original of Riccardi (1980, pl. 1

(figs 1-2), text-fig. 1) from the Upper Santonian or Lower Campanian of

Seymour (Vicecomodoro Marambio) Island, Antarctica. MLP 12248.

160 ANNALS OF THE SOUTH AFRICAN MUSEUM

Occurrence. Upper Santonian-—Lower Campanian, Seymour Island,

Antarctica.

Baculites dissimilis Desmarest, 1817

Desmarest 1817: 49, pl. 2 (figs 4-6) (= B. vertebralis fide Kennedy 1986c:

187).

Baculites distans De Grossouvre, 1901

De Grossouvre 1901, table 35 (in zone of Pachydiscus neubergicus) (=

nom. nud. fide Kennedy 1986b: 112).

Baculites duharti Hinicken, 1975

Hiinicken (in Hinicken 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. Hiinicken et al. 1980: 224, pl. 1

(figs 1-2), pl. 2 (figs 1-6), text-figs 3-9. Klinger & Kennedy 1997: 178,

figs 80c-d, 117e, 121-127.

Type. Holotype is the original of Htinicken (1975, pl. 1 (figs 1-4), text-

figs 2c-d, 4) from the Middle-Upper Campanian Cerro Matero Formation,

at Rio Sur, at the confluence with the Rio Condor, Chile. UCS D-1127.

Occurrence. Middle-Upper Campanian, southern Chile, Middle

Campanian, Zululand.

* Baculites eliasi Cobban, 1958 Fig. 123

Elias 1933: 306, pl. 28 (fig. 2), pl. 33 (fig. 5a-b) (as B. pseudovatus Elias

var. A.). Cobban 1958: 663, pl. 91 (figs 1-11), text-figs 1f-g, i-j. Scott &

Cobban 1965: 3, map I-439. Gill & Cobban 1973: 10, fig. 2r. Kennedy &

Cobban 1976, pl. 6 (fig. 3). Riccardi 1983, pl. 25 (figs 4-6). Larson et al.

1997: 33.

Type. Holotype is the original of Cobban (1958, pl. 91 (figs 9-11), text-

fig. 11)) from the Lower Maastrichtian Bearpaw Shale, near Fort Peck,

Montana, USNM 108969.

Occurrence. Lower Maastrichtian, US Western Interior (Montana,

Wyoming, Colorado, South Dakota, Kansas and Nebraska) and Alberta and

Saskatchewan, Canada.

Baculites embaensis Balan, 1982

Balan 1982: 210, pl. 19 (fig. 4).

Type. Holotype is the original of Balan (1982, pl. 19 (fig. 4)) from the

Upper Turonian of Kazakstan. PIN 3766/18.

Occurrence. Upper Turonian, Kazakstan.

Fig. 123 (see facing page). Baculites eliasi Cobban, 1958. A-B, F. USNM 108972,

paratype from the Bearpaw Shale, 1.7 miles north of Oswego, Montana.

C-E. USNM 108969, the holotype from the Bearpaw Shale, near Fort Peck, Montana. Copy

of Cobban (1958, pl. 91 (figs 4-6, 9-11)). Both x 1.

161

THE AMMONITE FAMILY BACULITIDAE

Fig. 123

162 ANNALS OF THE SOUTH AFRICAN MUSEUM

Baculites fairbanksi Anderson, 1902

(= uninterpretable fide Matsumoto 1959: 135, Howarth 1965: 370)

Anderson, 1902: 92, pl. 7 (figs 152-153), pl. 10 (fig. 194). Anderson 1958:

190, pl. 48 (fig. 4), pl. 49 (fig. 4).

Type. Anderson did not designate a type in his original (1902): 92)

description. Anderson (1958: 190) referred to the largest specimen as the

‘holotype’—this may be taken as lectotype designation. The lectotype was

recorded from Silverado Canyon, Orange County and dated as Turonian in

the plate description. UCLA collections.

Occurrence. The lectotype was dated as Turonian, another specimen (1958,

pl. 49 (fig. 4, 4a) as Coniacian, in the text (1958: 190) it is recorded as

being associated with Metaplacenticeras pacificum, which would date it as

late Campanian.

Baculites fairbanksi germanicum Lommerzheim, 1976

(probably a Sciponoceras)

Lommerzheim 1976: 222, pl. 1 (fig. la—-c), text-fig. 5.

Type. Holotype is the original of Lommerzheim (1976, pl. 1 (fig. 1b), text-

fig. 5b, d) from the Turonian of the Becker Brickworks, Miilheim-Broich,

Westphalia, Germany, Collection Klauman, CTB 1.

Occurrence. Turonian, northern Germany.

Baculites falcatus Collignon, 1969

(= B. menabensis)

Collignon 1969: 20, pl. 520 (figs 2045-2047). Klinger & Kennedy 1997: 94,

figs 58a-c, 61b.

Type. Holotype is the original of Collignon (1969, pl. 520 (fig. 2045)) from

the Lower Campanian, Gisement 303, Coupe Ampolypoly-Antsirasira-

Behamotra (Belo sur Tsiribihina), Madagascar, GD 12045.

Occurrence. Lower Campanian, Madagascar.

Baculites faujasii Lamarck, 1822

(= B. vertebralis Lamarck, 1801, fide Schliiter 1876: 143; Kennedy 1986a:

57)

Lamarck, 1822: 647. Favre 1869: 29.

Baculites fuchsi Redtenbacher, 1873

Redtenbacher 1873: 134, pl. 30 (fig. 15). Summesberger 1979: 113, pl. 1

(figs 2-4), text-figs 2-3. Summesberger 1979: 115, pl. 1 (figs 5-7), text-

fig. 4 (as B. cf. fuchsi). Immel et al. 1982: 28, pl. 11 (fig. 8). Immel 1987:

129. Kennedy & Christensen 1991: 217, pl. 7 (fig. la—b) (as B. cf. fuchsi).

Santamaria Zabala 1991: 269, pl. 18 (fig. 9), 1992: 239.

Type. Holotype by monotypy is the original of Redtenbacher (1873, pl. 30

(fig.15)) from the Santonian of the Gosau Beds at Tiefengraben, Austria,

NHMW 1865/1/138.

163

THE AMMONITE FAMILY BACULITIDAE

‘| x “BuTUOAM ‘(e[suRIPENd epseomeN) AJUNOD UOJsaM “"MA ~O “A “'N 8p LL “LI “90S YMN NAS UI ‘ESLTIC Artfeoo] stozosayy

qTeYS Wold oY} WO “YCTLLM WNO “OL8I “AW Sipunss sajnovg

SOSN ‘aUu0Z SIpUDIS SajIINIDg URIYOLISee, JOMOT

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p71 “3d

164 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 125. Baculites grandis Meek, 1876. OUM KT7201, from the Pierre Shale, Lower

Maastrichtian Baculites grandis zone, USGS Mesozoic locality D11783, in SE% sec. 17,

T. 48 N., R. 64 W., Weston County (Newcastle Quadrangle), Wyoming. x 1.

THE AMMONITE FAMILY BACULITIDAE 165

Occurrence. Lower to Upper Santonian, Austria and Spain.

* Baculites furcillatus Blanckenhorn?, 1905 Fig. 194A-C

(= ?Trachybaculites)

Taubenhaus 1920: 10, pl. 6 (fig. 2). (According to Picard (1929: 441),

possibly only a more strongly ribbed form of B. palestinensis.)

Type. Lectotype here designated is the original of Taubenhaus (1920, pl. 6

(fig. 2)) from West of Es-Salt, Palestine, Upper Campanian. Blanckenhorn’s

material was originally deposited in Breslau (Wroctaw) in the care of Frech

(see Taubenhaus 1920: 2). The material appears lost, and neotype

designation may be desirabie.

Occurrence. Upper Campanian, Israel.

Baculites gallischi Fritsch, 1897

(= indeterminable fide Diener 1925: 60)

Fritsch 1897: 40.

Baculites gigantea Desmarest, 1817

(= Diplomcceras cylindraceum Defrance) (fide Schliiter 1876: 148 footnote;

also Kennedy 1986c: 181)

Baculites gigas Sinzow, 1888

(= 2B. claviformis fide Cobban 1974: 8)

Sinzow 1888: 126.

* Baculites gilberti Cobban, 1962b Fig. 126

Cobban 1962b: 716, pl. 108 (figs 5-13), text-fig. 1d-f. Gill & Cobban 1973:

8, fig. 3a. Larson et al. 1997: 24.

Type. Holotype is the original of Cobban (1962), pl. 108 (figs 11-13)) from

the Middle Campanian Pierre Shale, just below Hygiene Member, north of

Boulder, Colorado, USNM 108911.

Occurrence. Middle Campanian, US Western Interior (Colorado, Wyoming,

South Dakota).

* Baculites grandis Hall & Meek, 1854 Figs 124-125

Hall & Meek 1854: 402, pl. 6 (fig. 10), pl. 7 (figs 1-2), pl. 8 (figs 1-2).

Meek 1876: 398, pl. 33 (fig. la—c), text-figs 53-54. Wade 1926: 182, pl. 60

(figs 8, 12). Elias 1933: 307, pl. 31 (figs la—b, 2a—-b), pl. 34 (figs 4, Sa-c).

Scott & Cobban 1965: 3, map I-439. Gill & Cobban 1973: 10, fig. 7b.

Larson et al. 1997: 35.

Type. ?

Occurrence. Maastrichtian, US Western Interior (Colorado, Wyoming,

Montana, Nebraska, South Dakota).

* Baculites gregoryensis Cobban, 1951 Fig. 127

Cobban 1951: 820, pl. 118 (figs 1-5), text-figs 8-13. Scott & Cobban 1965:

2, map I-439. Gill & Cobban 1973: 7, figs 21, 3b. Larson et al. 1997: 25.

166 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 126. Baculites gilberti Cobban, 1962b. A-C. USNM 108912b, paratype.

D-E. USNM 108912a, paratype. F-H. USNM 108911, the holotype. I. USNM 108912c,

paratype. All from the Pierre Shale just below the Hygiene Member at USGS Mesozoic

locality D236, north of Boulder, Colorado. Copy of Cobban (19625, pl. 108 (figs 5-13)).

All x 1.

THE AMMONITE FAMILY BACULITIDAE 167

A B C

Fig. 127. Baculites gregoryensis Cobban, 1951. USNM 507279.

From the Pierre Shale near Fort Thompson, South Dakota. x 1.

Type. Holotype is the original of Cobban (1951, pl. 118 (fig. 1), text-

figs 12-13), from the Middle Campanian Gregory Member of the Pierre

Shale, west of Oacama, Lyman County, South Dakota, USNM 106987.

Occurrence. Middle Campanian, US Western Interior (Montana, Wyoming,

South Dakota, Colorado, New Mexico).

* Baculites haresi Reeside, 1927 Fig. 97B-D

Reeside 1927b: 10, pl. 6 (figs 5-10), pl. 7 (figs 9-10) (as B. ovatus var.

haresi). Elias 1933: 298, pl. 35 (figs 3a—b, 4a—b) (as B. ovatus var. haresi).

Landes 1940: 170. ?Birkelund 1965: 56, pl. 7 (fig. 2), text-fig. 46 (as B. cf.

haresi). Cobban & Kennedy 1992b: 449, figs 6.5-6.8, 6.11-6.13,

6.16-6.24, 7.1. Kennedy & Cobban 1993e: 844, figs 14.18-14.35, 14.37,

15.2, 16.1-16.6. Larson et al. 1997: 20.

Type. Lectotype by subsequent designation of Cobban & Kennedy (1992b:

449) is the original of Reeside (1927), pl. 6 (figs 5-6)), from the Lower

Campanian, Elk Basin Sandstone Member of the Telegraph Creek

Formation, Carbon County, Montana, USNM 73296.

Occurrence. Widely distributed in the Lower Campanian, US Western

Interior (Montana, Wyoming, Utah, New Mexico, Kansas, South Dakota);

168 ANNALS OF THE SOUTH AFRICAN MUSEUM

THE AMMONITE FAMILY BACULITIDAE 169

Gober Chalk of Texas, U.S. Atlantic Seaboard (New Jersey),and possibly

West Greenland.

Baculites hochstetteri Liebus, 1902

Liebus 1902: 119 [7], pl. 6 (figs 4-6). Nowak 1908: 331, fig. 11. non

?Naidin 1974: 164, pl. 53 (figs 6-7), text-figs 22.6-7, 23.v (as B. anceps

hochstetteri).

Type. Lectotype by subsequent designation of Matsumoto & Obata (1963:

63) is the original of Liebus (1902, pl. 6 (figs 5a, 6) text-fig. 2b) from the

Senonian Friedekar Baculite Marls of the Silesian Carpathians. The types

could not be located in Munich and are presumed destroyed during WW II

(letter of Prof. H. Immel 25.5.1992).

Occurrence. Senonian, Galicia.

Baculites huenickeni Stinnesbeck, 1986

Stinnesbeck 1986: 201, pl. 8 (figs 7-8), pl. 9 (figs 1-2), text-fig. 24b-c.

Type. Holotype is the original of Stinnesbeck (1986, pl. 8 (fig. 7)), from the

Maastrichtian of La Gloria, San Vicente near Talcahuano, Chile,

GPIB VO/96.

Occurrence. Maastrichtian, southern Chile.

Baculites increscens Collignon, 1970

Collignon, 1970: 3, pl. 607 (figs 2266-2268), p. 5, pl. 608 (fig. 2269).

Klinger & Kennedy 1997: 124, figs 79a-l, n—-o, 80a—b, 82.

Type. Holotype is the original of Collignon (1970, pl. 607 (fig. 2266)) from

the Middle Campanian, Gisement 177, Coupe d’Ankilizato (Belo sur

Tsiribihina), Madagascar, GD12266.

Occurrence. Middle Campanian, Madagascar, Zululand, ?Israel.

* Baculites incurvatus Dujardin, 1837 Fig. 128A-P

Dujardin 1837: 232, pl. 17 (fig. 13a-d). D’Orbigny 1842: 564, pl. 139

(figs 8-10). D’Orbigny 1842: 565 (as B. tuberculata). Geinitz 1843: 9, pl. 1

(fig. 5). Fritsch & Schlénbach 1872: 51, pl. 13 (fig. 21). Schliiter 1876:

142, pl. 39 (figs 6-7), pl. 40 (fig. 3). non Moberg 1885: 36, pl. 4 (figs 2-4).

non Holzapfel 1887: 64, pl. 4 (figs 5-6), pl. 5 (fig. 10). Wegner 1905: 206.

Fig. 128 (see facing page). A-P. Baculites incurvatus Dujardin, 1837.

A-D. MNHPR1052, paralectotype; locality as for K-L. E-G. MNHP R1044b, D’Orbigny

collection no. 7208, the original of D’Orbigny (1842, pl. 139 (figs 8-10)) from Tours.

H-J. OUM KZ16643, Upper Coniacian Gauthiericeras margae zone, 30-50 cm above the

Calcaire Durs de la Ribochére of St Paterne-Racan (Indre-et-Loire). K-L. Lectotype,

MNHP R1025a; the locality is simply given as Touraine. M. Paralectotype, MNHP

R1025ci; locality as for K-L. N-P. MNHPR1044a, D’Orbigny collection no. 7208, the

original of D’Orbigny (1842, pl. 139 (figs 8-10)), from ‘Tours’. Q-S. Baculites cf.

B. brevicosta Schliter, 1876. OUMKZ17023, from the same locality and horizon as H-J.

All x 1.

170 ANNALS OF THE SOUTH AFRICAN MUSEUM

THE AMMONITE FAMILY BACULITIDAE 171

non Miller & Wolleman 1906: 4, pl. 2 (figs 2-5). ?Scupin 1913: 102, pl. 3

(figs 1, 8). non Hagg 1930: 58. non Hagg 1935: 58. non Collignon 1938:

88, pl. 6 (figs 4-5). Wright 1957: L218, fig. 245.3. Immel e¢ al. 1982: 27,

pl. 11 (figs 5-7). Kennedy 1984: 143, pl. 32 (figs 12, 15-19), pl. 33

(figs 1-22), text-figs 41, 42f-m. Immel 1987: 128. ?Alizade et al. 1988:

378, pl. 27 (figs 2-5) (as B. incervatus [sic.]). Thomel 1988: 18. Kennedy

& Christensen 1991: 217, pl. 1 (fig. 2a—b) (as Baculites cf. incurvatus).

Santamaria Zabala 1991: 267, pl. 18 (fig. 8). VaSi¢ek 1993: 182, pl. 3

(fig. 4), text-figs 10-11. Santamaria Zabala 1992: 238, pl. 2 (fig. 8). Klinger

& Kennedy 1997, fig. 56.

Type. Lectotype by the subsequent designation of Immel et al. (1982: 27), is

the original of Dujardin (1837, pl. 17 (fig. 13a)), imprecisely located as

‘Craie Tufau’, MNHP R1025a.

Occurrence. Middle Coniacian to Santonian France, Germany, Spain,

Czechoslovakia, Austria, ?Sweden, ?Azerbaijan.

Baculites incurvatus Moberg, 1885 non Dujardin, 1837

Kennedy & Christensen 1997: 108, fig. 20c-—e.

Occurrence. Upper Middle Santonian of Eriksdal, Sweden.

* Baculites inornatus Meek, 1862 Fig. 108G-I, 129

Meek, 1862: 316. Anderson & Hanna 1935: 24, pl. 8 (figs 1-2). Anderson

1958: 190, pl. 48 (figs 2-3). Matsumoto 1959: 155, pl. 38 (fig. la—c), pl. 43

(fig. 5a—c), text-figs 72a—b, 73a—d, 74-79. Obata & Matsumoto 1963: 78,

pl. 22 (fig. 1), pl. 24 (fig. 6), pl. 26 (figs 4-6), text-figs 169-170, 187-190.

Ward 1978: 1151, pl. 1 (figs 1-2), text-fig. 5. non Renz 1982: 105, pl. 34

(figs 3-6), text-fig. 80. Matsumoto & Miyauchi 1984: 72, pl. 25 (fig. 6),

text-fig. 12. Morozumi 1985: 45. Haggart 1991, pl. 3 (fig. 3-4).

Type. Lectotype by the subsequent designation of Matsumoto (1959: 155), is

one of Meek’s unfigured syntypes, figured by Matsumoto (1959, pl. 43

(fig. 5a—c), text-fig. 72a—b) from Sucia Island, Washington, USNM 1259.

Occurrence. Middle to Upper Campanian, California, Baha California

(Mexico), British Columbia and Washington State, and Lower Campanian

of Honshu.

* Baculites jenseni Cobban, 1962a Figs 130-131

Cobban 1962a: 129, pl. 26 (figs 1-12), text-fig. la. Scott & Cobban 1965:

3, map I-439. Gill & Cobban 1973: 10, fig. 2q. Larson et al. 1997: 32.

Type. Holotype is the original of Cobban (1962a, pl. 26 (figs 1, 11-12))

from the Upper Campanian, upper part of the Bearpaw Formation, north-

east of Melstone, Montana, USNM 131117.

Fig. 129 (see facing page). Baculites inornatus Meek, 1862. A-C. CAS-66772.03; from

Fresno County, Coalinga Quadrangle, from bottom of Copper Canyon, in lens of boulder

conglomerate near bottom of Joaquin Ridge Sandstone. D-F. CAS-66772.02; from the

same locality as A-C. Both x 1.

172 ANNALS OF THE SOUTH AFRICAN MUSEUM

A B

Fig. 130. Baculites jenseni Cobban, 1962a. USNM 131118b, paratype,

from the Baculites jenseni zone of the Bearpaw Shale, USGS Mesozoic

locality D915, 12 miles north-east of Melstone, Montana.

Copy of Cobban (1962a, pl. 26 (figs 7-8)). x 1.

Occurrence. Upper Campanian, US Western Interior (Montana, South

Dakota, Colorado, Wyoming, Kansas), Alberta and Saskatchewan, Canada.

Baculites kegeli Oliveira, 1957

(= ?Trachybaculites)

Oliveira 1957: 22, pl. 2 (figs 6-7).

Type. Holotype by monotypy is the original of Oliveira (1957, pl. 2

(figs 6-7), text-fig. 1) from the Maastrichtian, Gramame Formation, Olinda

State, Brazil, Nr 4486, Cat. de Inv. da Diviséo de Geologia e mineralogia

do departamento Nacional da Producao Mineral, Rio de Janeiro.

Occurrence. Maastrichtian, Brazil.

Baculites kirki Matsumoto, 1959

Matsumoto 1959: 143, pl. 43 (figs la-c, 2a—-c, 3a-c), text-figs 53a-b,

54-57, 58a—b. Matsumoto & Obata 1963: 65, pl. 18 (fig. 2), text-fig. 114

THE AMMONITE FAMILY BACULITIDAE 173

Fig. 131. Baculites jenseni Cobban, 1962a. A-B. USNM 131119b, paratype.

C-F. USNM 131119a, paratype. G. USNM 131118a, paratype. H-J. USNM 131117, th

holotype. All from the upper part of the Bearpaw Shale, USGS Mesozoic locality D915,

12 miles north-east of Melstone, Montana. Copy of Cobban (1962a, pl. 26 (figs 1-6,

11-12)). x 1.

174

ANNALS OF THE SOUTH AFRICAN MUSEUM

(as B. cf. kirki). ?Riccardi & Aguirre Urreta 1988, pl. 3 (figs 4-8) (as B. cf.

kirki). Olivero 1992: 51, pl. 1 (figs 1-3) (as B. aff. kirki).

Type. Holotype is the original of Matsumoto (1959, pl. 43 (fig. la—c), text-

fig. 53a—b) from the Santonian of Hooten Gulch in the Redding Area,

California, UCLA UC 35693.

Occurrence. Santonian s.1., California, doubtfully Santa Cruz Province,

Argentina and James Ross Island, Antarctica.

*Baculites knorrianus Desmarest, 1817 Figs 132-133

Desmarest 1817: 48, pl. 1 (fig. 3). Favre 1869: 27, pl. 7 (figs 2-4). Schliiter

1876: 146 (ars), pl. 39 (figs 16-18, 20 non 19). Mikhailov 1951: 46, pl. 3

(fig. 15) (as B. anceps var. leopoliensis). Birkelund 1979: 53. Kennedy &

Summesberger 1987: 32, pl. 4 (figs 4-6), pl. 5 (figs 1-14), text-fig. 2 (cum.

synon.). Vasicek 1988: 77, pl. 1 (fig. 4), text-fig. 2. Birkelund 1993: 52,

pl. 13 (figs 12-14). Kennedy 1993: 109, pl. 5 (figs 13-22), pl. 6

(figs 11-13, 18-23), text-fig. 5a-c. Kennedy & Christensen 1997: 111,

fig. 29.

Type. Desmarest based this species on a figure in Knorr & Walch (1755,

pl. 12), which he reproduced as pl. 1 (fig. 3); the type is lost. Neotype

designated by Kennedy & Summesberger (1987: 33) is the original of

Kennedy & Summesberger (1987, pl. 5 (figs 5, 7-8) from the Lower

Maastrichtian of the Ukraine, NHMW 7459a.

Occurrence. Lower Maastrichtian Ukraine, Belgium and northern Germany;

in Denmark it spans the Lower-Upper Maastrichtian boundary; also Poland

and Czechoslovakia.

Baculites kotanii Matsumoto, Hashimoto & Furuichi, 1980

Matsumoto ef al. 1980: 408, figs 1-2. Matsumoto & Miyauchi 1984: 73,

pl 251g. 7):

Type. Holotype is the original of Matsumoto ef al. (1980, figs la—b, 2a))

from the Upper Campanian of Gesho, Shikoku, Japan, Geological

Laboratory, Kagawa University, Takamatsu, GLKU IZ 80001.

Occurrence. Upper Campanian (see also Bando & Hashimoto 1984),

Shikoku, Japan.

Baculites labyrinthicus Morton, 1834

(see Eubaculites)

Morton 1834: 44, pl. 13 (fig. 10).

Baculites latelobatus Collignon, 1931

(= B. yokoyamai Tokunaga & Shimizu, 1926 = B. besairiei Collignon,

1931)

Collignon 1931: 38, pl. 5 (figs 11, 1la, 12, 12a), pl. 9 (fig. 18).

Type. Lectotype here designated is the original of Collignon (1931, pl. 5

(fig. 11)) from the Upper Coniacian of Mahagaga, Madagascar, GD

unregistered.

Occurrence. Upper Coniacian, Madagascar.

THE AMMONITE FAMILY BACULITIDAE 175

Fig. 132. Baculites knorrianus Desmarest, 1817. A-C. NHMW 7459, the original of Favre

(1869, pl. 7 (fig. 2)). D-F. Neotype, NHMW 7459, the original of Favre (1869, pl. 7

(figs 3-4, 6)) from the Lower Maastrichtian of Nagoryany, Lvov, Ukraine. Both x 1.

176 ANNALS OF THE SOUTH AFRICAN MUSEUM

A B Cc

Fig. 133. Baculites knorrianus Desmarest, 1817. NHMW 7459c from the

Lower Maastrichtian of Nagoryany, Lvov, Ukraine. x 1.

Baculites larsoni Cobban & Kennedy, 1992

Cobban & Kennedy 1992c: 682, figs 1.5, 1.6, 1.10-1.20, 2, 3.1.

Type. Holotype is the original of Cobban & Kennedy (1992c, figs 1.14-

1.16, 3.3) from the Timber Lake Member of the Fox Hills Formation, Bear

Creek gravel pit, Dewey County, South Dakota, Maastrichtian.

Occurrence. Maastrichtian, South Dakota.

Baculites larteti Blanckenhorn, 1905

Blanckenhorn 1905: 111. Picard 1929: 443, pl. 10 (fig. 9) (as B. asper var.

larteti.

Type. ?

Occurrence. Maastrichtian, Palestine.

THE AMMONITE FAMILY BACULITIDAE 177

Fig. 134. Baculites leopoliensis Nowak, 1908. OUMKZ20219. Upper Campanian,

Nostoceras pozaryskii zone, Piotrawin, Poland.

aut

Baculites lechitides Brunnschweiler, 1966

(?= Trachybaculites)

Brunnschweiler 1966: 23, pl. 1 (figs 1-3), text-fig. 8. Henderson ef al.

1992: 162, fig. 23g-j (as Baculites? lechitides).

Type. Holotype is the original of Brunnschweiler (1966, pl. 1 (figs 1-3)),

from the Maastrichtian of the Marilla Anticline, Miria Marl, Western

Australia. CBC 2689.

Occurrence. Maastrichtian, Western Australia.

178

ANNALS OF THE SOUTH AFRICAN MUSEUM

F G H

Fig. 135. Baculites lomaensis Anderson, 1958. A-B. USNM 463147.

C. USNM 463148. D-E. USNM 463149. F-H. USNM 463158. All from the

Maastrichtian Prairie Bluff Chalk of Alabama. All x 1.

* Baculites leopoliensis Nowak, 1908 Fig. 134

Nowak 1908: 328, pl. 14 (figs 1-5), text-figs 1-5 (p. 329), text-figs 5-10

(p. 331) (as B. anceps var. Leopoliensis). Elias 1933: 310, pl. 34 (fig. 3) (as

B. anceps var. leopoliensis). non Mickailov 1951: 46, pl. 3 (fig. 15) (as

B. anceps var. leopoliensis). Glazunova 1955: 180, pl. 1 (figs 1-10). non

Tzankov 1964: 149, pl. 10 (fig. 2) (as B. anceps leopoliensis). non

Collignon 1970: 10, pl. 610 (figs 2277-2278). non Naidin & Shimanskij

1959: 164, pl. 53 (fig. 5) (as B. anceps leopoliensis). non Atabekian &

Khakimov 1976: 96, pl. 11 (figs 11-13) (as B. anceps leopoliensis). non

Tzankov 1982: 17, pl. 3 (figs 2-3) (as B. anceps leopoliensis). Kennedy

1986e: 1013, pl. 2 (figs 1-2, 11-12), pl. 3 (figs 22-24). Kennedy 1986b:

114, pl. 18 (figs 1, 4-12). Hancock & Kennedy 1993: 165, pl. 20

(figs 9-11, 14, 16-19). Klinger & Kichler 1998: fig. 2g.

Type. Lectotype by the subsequent designation of Kennedy (1986e: 1013) is

the original of Nowak (1908, pl. 14 (fig. 2)) from the Upper Campanian of

Lipniki, formerly Galicia, now in Ukraine.

Occurrence. Upper Campanian of Ukraine, Poland and France. (Previous

records from the Lower Maastrichtian, e.g. Kennedy 1986b: 115, are

reputed to be incorrect, fide Hancock & Kennedy 1993: 165.)

Fig. 136 (see facing page). A-B. Baculites mclearni Landes, 1940. A. GSC 9366a,

paratype, from the Pakowki Shale, 30 ft below top, NW'%4, sec. 20, tp. 1, range 4th Mer.,

east side of Bear Gulch, Alberta. B. GSC 9366a, the holotype from the same locality as A.

C-E. Baculites minerensis Landes, 1940. GSC 9365, the holotype, from the Lower Pakowki

Formation, at the mouth of a small coulée entering Milk River Valley from the south near its

junction with Deadhorse Coulée, NW sec. 11, tp. 2, range 11, W. 4th mer.

Aiole

THE AMMONITE FAMILY BACULITIDAE 179

Fig. 136

180 ANNALS OF THE SOUTH AFRICAN MUSEUM

Baculites libyensis Hassan, 1971

(= ?B. anceps)

Hassan 1971: 71, pl. 2 (fig. 12).

Type. Holotype (by monotypy?) is the original of Hassan (1971, pl. 2

(fig. 12)) from the Maastrichtian of Gebel Tarawan, Kharga Oasis, Egypt,

ASUE 210.

Occurrence. Maastrichtian, Egypt.

*Baculites lomaensis Anderson, 1958 Figs 104L-N, 135

Anderson 1958: 191, pl. 48 (figs 5, 5a, 6). Matsumoto 1959: 126, pl. 34

(figs la—c, 2a—c), text-figs 35-38, 39-41. ?7Dundo 1971, pl. 1 (fig. 7, 7a) (as

B. cf. lomaensis). Cobban & Kennedy 1995: 18, figs 13.1-13.3,

13.7-13.59, 14.8, 14.10, 15.9.

Type. Holotype is the original of Anderson (1958, pl. 48 (fig. 5)) from the

?Upper Campanian of Joaquin Ridge, north of Coalinga, upper part of

Panoche Formation, California, CAS type collection.

Occurrence. ? Upper Campanian, California, Maastrichtian Prairie Bluff

Chalk of Alabama and Mississippi; ?Russian Arctic.

Baculites maximus Hagenow, 1842

(= ?B. knorrianus fide Schliter 1876: 148 footnote)

Hagenow 1842: 567.

* Baculites mclearni Landes, 1940 Fig. 136A-B

Landes 1940: 165, pl. 7 (figs 1-3) . Cobban 1962b: 712, pl. 105 (fig. 15),

pl. 107 (figs 17-19), text-fig. 1g-h. Scott & Cobban 1965: 1, map I-439.

Gill & Cobban 1973: 5. Kauffman 1977: 267, pl. 27 (figs 19-21), pl. 30

(fig. 9). Cobban & Kennedy 1993: 75, figs 5.1-5.2, 6.1-6.14. Larson et al.

1997: 22.

Type. Holotype is the original of Landes (1940, pl. 7 (fig. 1)) from the

Middle Campanian of the Pakowski Formation, Montana Group, Alberta,

Canada, GSC 9366.

Occurrence. Middle Campanian, US Western Interior (Montana,

?Wyoming, Colorado, South Dakota), north-eastern Texas, and Alberta and

Saskatchewan, Canada.

Baculites malagasyensis Collignon, 1966

(= B. capensis)

Collignon 1966: 7, pl. 457 (fig. 1865).

Type. Holotype (by monotypy?) is the original of Collignon (1966,

pl. 457 (fig. 1865)) from the Lower Santonian of Gisement 272, coupe

Ampolypoly-Antsirasira-Behamotra (Belo sur Tsiribihina), Madagascar,

GD 11865.

Occurrence. Lower Santonian, Madagascar.

THE AMMONITE FAMILY BACULITIDAE 181

Fig. 137. Baculites mariasensis Cobban, 1951. A-C. USNM 433869 from the Mancos

Shale, USGS Mesozoic locality D7230, in Sevier County, Utah. D-F. USNM 433877 from

the Marais River Shale, USGS Mesozoic locality 21422, in Toole County, Montana.

G-I. USNM 433885 from the same locality. J-M. USNM 433876 from the same locality.

All from the Middle Coniacian Inoceramus deformis zone. All x 1.

182 ANNALS OF THE SOUTH AFRICAN MUSEUM

Baculites mamillatus Collignon, 1970

(= B. increscens)

Collignon, 1970: 7, pl. 609 (figs 2273-2274). Klinger & Kennedy 1997:

132, figs 84a-c, 85a.

Type. Holotype is the original of Collignon (1970, pl. 609 (fig. 2273)) from

the Middle Campanian of Gisement 181, Coupe d’Ankilizato (Belo sur

Tsiribihina), Madagascar, GD 12273.

Occurrence. Middle Campanian, Madagascar and Zululand.

* Baculites mariasensis Cobban, 1951 Fig. 137

Cobban 1951: 818, pl. 118 (figs 10-12), text-figs 4-7. Cobban 1955: 204,

pl. 2 (fig. 1). Cobban 1976: 124, pl. 2 (figs 5-6). Kauffman 1977: 270,

pl. 29 (fig. 10). 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.

Type. Holotype is the original of Cobban (1951, pl. 118 (fig. 12), text-

figs 5-6) from the Middle Coniacian Colorado Shale, south of Shelby, Toole

County, Montana, USNM 106985.

Occurrence. Middle Coniacian, US Western Interior (Montana, Wyoming,

Utah).

* Baculites meeki Elias, 1933 Fig. 99A-C

Meek 1876: 404, pl. 39 (fig. 10b-c, non 10a, d) (as B. asper). Elias 1933:

312, pl. 30 (fig. 4a-b). ?Birkelund 1965: 62, pl. 14 (figs 2-3), pl. 15

(fig. 1), text-figs 53-55 (as B. cf. meeki).

Type. Holotype is the original of Elias (1933, pl. 30 (fig. 4a—b)) from the

Maastrichtian, Beecher Island Shale Member, Pierre Shale, Beecher Island,

Yuma County, Colorado. GMUK 59670.

Occurrence. Maastrichtian, US Western Interior (Colorado, South Dakota),

possibly West Greenland.

Baculites menabensis Collignon, 1969

(= ?B. tanakae)

Collignon 1969: 15, pl. 518 (figs 2036-2037). ?Matsumoto & Miyauchi

1984: 74, pl. 25 (fig. 8) (as B. sp. aff. menabensis). Klinger & Kennedy

1997: 93, figs 57, 62c.

Type. Holotype is the original of Collignon (1969, pl. 518 (fig. 2036)), from

the Lower Campanian of Gisement 307, Coupe Ampolypoly-Antsirasira-

Behamotra (Belo sur Tsiribihina), Madagascar, GD 12036.

Occurrence. Lower Campanian Madagascar, possibly Hokkaido.

* Baculites minerensis Landes, 1940 Fig. 136C-E

Landes 1940: 166, pl. 6 (figs 7-8).

Type. Holotype is the original of Landes (1940, pl. 6 (figs 7-8)), from the

lower Pakowski Formation, GSC 9365.

Occurrence. Upper Campanian, Canada.

THE AMMONITE FAMILY BACULITIDAE 183

Baculites natosini Robinson, 1945

(= Pseudobaculites)

Robinson 1945: 52, pl. 1 (figs 5-6).

Baculites nibelae Klinger & Kennedy, 1997

Klinger & Kennedy 1997: 162, figs 116, 117a—d, f-i, 118-120.

Type. Holotype is SAM-PCZ9148 from locality 111, Nibela, Zululand,

St Lucia Formation, Campanian III.

Occurrence. Campanian, Zululand.

Baculites nitidus Glazunova, 1955

Glazunova 1955: 185, pl. 3 (figs 4-6).

Type. ?

Occurrence. Siberia.

Baculites nugssuaquensis Birkelund, 1965

Birkelund 1965: 48, pl. 4 (fig. 1), pl. 5 (figs 1-4), pl. 6 (figs 1-2), text-

figs 35-41.

Type. Holotype is the original of Birkelund (1965, pl. 4 (figs la-c), text-

fig. 35a-e) from the Lower Santonian of Tuperssuarta, West Greenland,

MMH 9755.

Occurrence. Lower Santonian, West Greenland.

Baculites oberholzeri Bohm, 1909

(= ?B. anceps)

Bohm 1909: 52, pl. 1 (fig. 9a-b).

Type. Holotype by monotypy is the original of Bohm (1909, pl. 1

(fig. 9a—b)) from the Senonian, Leiboden marls of Leibodenalp,

Switzerland, Geological Institute of ETH, Zurich.

Occurrence. Senonian, Switzerland.

Baculites obliquatus Sowerby, 1828

(= Sciponoceras baculoides)

*Baculites obtusus Meek, 1876 Fig. 100C-H, 138

Meek 1876: 406, text-figs 57-60. Landes 1940: 169. Cobban 19625: 706,

pl. 105 (figs 1-14). Birkelund 1965: 58, pl. 8 (fig. 1), pl. 9 (figs 1-3), pl. 10

(fig. 1), pl. 11 (figs 1-2), pl. 12 (figs 1-2), pl. 13 (figs 1-2), pl. 14 (fig. 1),

text-figs 47-52 (as B. obtusus s.1.). Scott & Cobban 1965: 1, map I-439.

Gill & Cobban 1973: 5, fig. 2i. Larson et al. 1997: 22.

Type. Meek did not designate a type. Landes (1940: 169) referred to Meek’s

(1876, figs 58-60) as the ‘holotype’. Cobban (1962b: 708) referred to the

Same specimen as the lectotype. It is from the Middle Campanian of Deer

Creek on the North Platte River, from the Fox Hills Group of the Upper

Missouri Cretaceous Series, USNM 1934a.

184

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 138. Baculites obtusus Meek, 1876. A-B. USNM 13101le.

C-D. USNM 131011c. E-G. USNM 131011b. H-J. USNM1310ila. All from the

Pierre Shale, USGS Mesozoic locality D1958, north-east of Canon City in SW%

sec. 10, T. 18 S., R. 70 W., Fremont County, Colorado. All x 1.

Occurrence. Middle Campanian US Western Interior (New Mexico,

Wyoming, South Dakota, Colorado, Montana), Alberta and Saskatchewan,

Canada and ?Lower Campanian of West Greenland.

Baculites occidentalis Meek, 1862

(= Eubaculites fide Klinger & Kennedy 1993: 190)

Meek 1862: 316.

Fig. 139 (see facing page). Baculites aquilaensis Reeside. Collections of the U.S.

Geological Survey, Denver, from USGS Mesozoic locality 23639 lower Gammon Shale

Member of Pierre Shale along west-facing scarp at head of Owl Creek in SE% sec. 12,

T.9S., R. 61 E., and N% sec. 13, T. 9 S., R. 16 E., Carter County, Montana. Lower

Campanian Zone of Scaphites hippocrepis sensu lato.

All x 1.

185

THE AMMONITE FAMILY BACULITIDAE

aS CX.

mene

Fig. 139

186 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 140. A-D. Baculites aquilaensis Reeside. Collections of the U.S. Geological

Survey, Denver, from USGS Mesozoic locality 23639 lower Gammon Shale

Member of Pierre Shale along west-facing scarp at head of Owl Creek in SE% sec. 12,

T. 9S., R. 61 E., and N% sec. 13, T. 9 S., R. 16 E., Carter County, Montana.

Lower Campanian Zone of Scaphites hippocrepis sensu lato.

lle

Baculites orientalis Matsumoto MS

(= nom. nud.) (= Sciponoceras orientale Matsumoto & Obata, 1963: 21)

Matsumoto 1942: 195, 197.

THE AMMONITE FAMILY BACULITIDAE 187

Baculites ornatus d’Orbigny, 1847

(= Eubaculites vagina)

D’Orbigny 1847, pl. 3 (figs 3-6).

Baculites ovatoides Hanna & Anderson, 1928

(= nom. nud. fide Anderson & Hanna 1935: 24).

Baculites ovatus Say, 1821

Say 1821: 41, pl. 5 (figs 5-6). Morton 1834: 42, pl. 1 (figs 6-8). Meek

1876: 394, pl. 20 (figs la—-b, 2a—-b, d). Johnson 1905: 26. non Dowling

1917, pl. 30 (fig. 2). ?7Taubenhaus 1920: 11, pl. 6 (fig. 1). Wade 1926: 181

(pars) non. pl. 60 (fig. 9). Reeside 1927b: 9 (pars). ?Picard 1929: 446, pl. 9

(fig. 11) (as B. vertebralis Lamarck (?) var. ovatus). non Elias 1933: 297,

pl. 33 (fig. 3a—c). Landes 1940: 171. non Reyment 1958: 7, pl. 1 (figs 1-2),

text-figs 1-2 (= B. yokoyamai). ?Bodylevski & Shulgina 1958, pl. 44

(fig. 5a—b). Reeside 1962: 113, pl. 68 (figs 1-4). Cobban 1974: 3, pl. 1

(figs 1-32), pl. 2 (figs 1-14), pl. 3 (figs 1-6, 9-11), text-fig. 4. ?7Luger &

Groéschke 1989: 398, pl. 49 (fig. 2), text-fig. 15i-o. ?Al-Harithi & Ibrahim

1992: 433, pl. 1 (fig. 3), pl. 2 (fig. 5). Kennedy & Cobban 1993c: 426,

figs 12.3-12.4, 14.5, 14.12, 14.15-14.17, 14.20, 14.22-14.42, 15.1-15.12,

15.14-15.24. Kennedy & Cobban 1994b: 1297, figs 5.1-5.2, 14.1-14.30,

14.33-14.35.

Type. Johnson (1905: 26) recorded the type in the collections of the

Academy of Natural Sciences, Philadelphia, but this is disputed—see

Cobban (1974: 4) for details.

Occurrence. Atlantic Highlands, New Jersey, where it is dated as late

Campanian and Arkansas; the US Western Interior, Middle East and North

African records are dubious.

Baculites ovatus var. baculus Meek, 1862

(= B. baculus)

Meek 1862: 445. Meek 1876: 397, text-figs 51-52.

Baculites ovatus var. haresi Reeside, 1927

(= B. haresi)

Reeside 1927b- 10, pl. 6 (figs 5-10), pl. 7 (figs 9-10).

Baculites palestinensis Picard, 1929

(= ?B. subanceps fide Howarth 1965: 370)

Picard 1929: 438, pl. 10 (figs 1-7), text-figs 1-2.

Type. Lectotype here designated, is the original of Picard (1929, pl. 10

(fig. la—b)) from the Maastrichtian? of Palestine, collections Department of

Geology, Hebrew University of Jerusalem, Israel (cf. Ferber 1985: 66) nr

HU 5278.

Occurrence. Maastrichtian?, Palestine.

188 ANNALS OF THE SOUTH AFRICAN MUSEUM

Baculites paradoxus Pervinquiére, 1907

(= probably not Baculites; Polyptychoceratinae)

Pervinquiére 1907: 94, pl. 4 (figs 10-11), text-fig. 24.

Type. The specimens may be lost, and lectotype designation is deferred.

Occurrence. Maastrichtian, Tunisia.

* Baculites perplexus Cobban, 1962b Figs 141-142

Cobban 1962b: 714, pl. 107 (figs 1-16), text-fig. la—c. Scott & Cobban

1965: 2, map I-439. Gill & Cobban 1973: 7, fig. 2k. Kennedy & Cobban

1976, pl. 8 (fig. 2), pl. 10 (fig. 8). Larson et al. 1997: 24.

Type. Holotype is the original of Cobban (19625, pl. 107 (figs 12-14)) from

the Middle Campanian upper part of the Steele Shale near Glenrock,

Wyoming, USNM 108914.

Occurrence. Middle Campanian, US Western Interior (Wyoming, Montana,

Colorado, South Dakota, Nebraska, New Mexico, Utah).

Baculites princeps Matsumoto & Obata, 1963

Matsumoto & Obata 1963: 55, pl. 13 (figs 1-2), pl. 15 (figs 1-2), text-

figs 134-139. Obata & Matsumoto 1963: 80, pl. 22 (fig. 2), pl. 25 (fig. 2),

pl. 27 (fig. 10), text-figs 171, 197-199. (as B. cf. princeps).

Type. Holotype is the original of Matsumoto & Obata (1963, pl. 13 (fig. 1),

text-fig. 134, 138), from the Santonian, Kotodo-no-sawa, Haboro Valley,

Teshio Province, Hokkaido, GK H4454.

Occurrence. Santonian, Hokkaido, ?Honshu.

Baculites pseudobaculus Matsumoto & Obata, 1963

Matsumoto & Obata 1963: 68, pl. 14 (fig. 3), text-figs 160-162, 166, 167.

Type. Holotype is the original of Matsumoto & Obata (1963, pl. 14 (fig. 3),

text-figs 162a—b, 166) from the Lower Santonian, Sankebetsu, a branch of

the Chikubetsu, Teshio Province, Hokkaido, GK H4803.

Occurrence. Lower Santonian, Hokkaido.

* Baculites pseudovatus Elias, 1933 Figs 99D, 143

Elias 1933: 304, pl. 29 (figs la—b, 2) only, non pl. 28 (fig.2), pl. 33

(fig. 4a—b, non fig. 5a—b)). Landes 1940: 174. Larson et al. 1997: 27.

Type. Holotype is the original of Elias (1933, pl. 29 (fig. la—b)) from the

upper Weskan Shale Member, Pierre Formation, Wallace County, Kansas,

GMUK 59668.

Fig. 141 (see facing page). Baculites perplexus Cobban, 1962b. A. Holotype

USNM 108914, from the Lower Campanian Pierre Shale near Glenrock, Wyoming, USGS

Mesozoic locality D255. B. Mass occurrence from the Campanian Pierre Shale near Red

Bird, Wyoming, USGS Mesozoic locality D1871. After Gill & Cobban (1966).

Both x 1.

189

THE AMMONITE FAMILY BACULITIDAE

Fig. 141

ANNALS OF THE SOUTH AFRICAN MUSEUM

190

Fig. 142

THE AMMONITE FAMILY BACULITIDAE 191

Occurrence. Upper Campanian, US Western Interior (Kansas, Colorado,

South Dakota, Montana, Nebraska, New Mexico).

Baculites rectangulatus Collignon, 1970

(= ?B. increscens)

Collignon 1970: 12, pl. 611 (figs 2279-2281). Klinger & Kennedy 1997:

135, fig. 87.

Type. Holotype is the original of Collignon (1970, pl. 611 (fig. 2279)) from

the Middle Campanian of Gisement 157 Coupe d’ Ankilizato (Belo sur

Tsiribihina), Madagascar, GD 12279.

Occurrence. Middle Campanian, Madagascar.

Baculites rectus Marshall, 1926

Marshall 1926: 154, pl. 19 (fig. 1), pl. 32 (figs 9-10). Spath 1953: 19, pl. 7

(fig. 2a—c) (as B. aff. rectus). Henderson 1970: 23, pl. 3 (figs 2-3), text-

fig. 6. ?Collignon 1977: 10 (as Baculites sp. indet.). Olivero 1984: 64, pl. 1

(figs 6-9), text-figs lc, 2. ?7Klinger & Kennedy 1997: 47, figs 25-26 (as

Baculites sp. aff. B. rectus).

Type. Lectotype by the subsequent designation of Henderson (1970: 23) is

the original of Marshall (1926, pl. 32 (figs 9-10)) from the ?Campanian of

Bull Point, New Zealand, GSNZ CE722.

Occurrence. ?Campanian—Maastrichtian, New Zealand, James Ross Island,

Antarctica, and possibly Graham Land, subsurface deposits, Durban, and

New Caledonia.

* Baculites reduncus Cobban, 1977 Figs 144-145

Cobban 1977: 462, figs 2-6. Kennedy & Cobban 1993d: 144, pl. 7

(figs 28-32). Larson et al. 1997: 25.

Type. Holotype is the original of Cobban (1977, fig. 2f-g) from the Middle

Campanian, lower part of the Rock River Formation near Rock River,

Wyoming, USNM 108909.

Occurrence. Middle Campanian, US Western Interior (Wyoming, Colorado,

South Dakota), and Gulf Coast (Arkansas).

* Baculites reesidei Elias, 1933 Figs 99E-G, 146

Reeside 1927b: 10 (pars), pl. 9 (figs 1-5) (as B. compressus). Elias 1933:

302, pl. 28 (fig. 1), pl. 31 (fig. 3), pl. 32 (fig. 2a—-c), pl. 33 (figs 1-2), (as

B. compressus var. reesidei). Gill & Cobban 1973: 10, fig. 2p, 3c. Riccardi

1983, pl. 26 (figs 3-5). Larson et al. 1997: 32.

Type. Holotype is the original of Reeside (1927b, pl. 9 (figs 1-5)), said to be

from the Campanian Eagle Sandstone in Fergus County, Montana, but from

the Bearpaw Shale, probably from central Montana, USNM 73304.

Fig. 142 (see facing page). Baculites perplexus Cobban, 1962b. A-C. OUM KT7590.

D-F. OUM KT7588. Both from the Mitten Member of the Pierre Shale, Middle Campanian

B. perplexus zone in NE% sec. 21, T. 46 N., R. 64 W., Weston County, Wyoming.

Both x 1.

192 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 143. Baculites pseudovatus Elias, 1933. USNM 507280, from USGS

Mesozoic locality D13537, Pierre Shale near Newcastle, Wyoming. x1.

Occurrence. Upper Campanian, US Western Interior (Kansas, South

Dakota, Montana, Wyoming, North Dakota, Colorado) and Canada.

Baculites regina Obata & Matsumoto, 1963

Obata & Matsumoto 1963: 85, pl. 22 (figs 3-6), pl. 23 (figs 1-2), pl. 24

(figs 1-5), pl. 25 (figs 3-5), pl. 27 (figs 1, 6-7, 9), text-figs 191-196,

200-214. Matsumoto & Morozumi 1980: 24.

Type. Holotype is the original of Obata & Matsumoto (1963, pl. 24 (fig. 1))

from the Campanian? Azenotani Shale Member at Kuratani, Shinke-mua,

Sennan-gun, Osaka Prefecture, Honshu, GT I-575.

Occurrence. Campanian, Honshu.

*Baculites rex Anderson, 1958 Figs 108A-C, 147

Anderson 1958: 191, pl. 49 (fig. 2). Matsumoto 1959: 136, pl. 31

(fig. 5a-d), pl. 34 (fig. 5), pl. 39 (fig. la-c, 2a-c, 3), pl. 40 (fig. la-c),

text-figs 45a—b, 46a-c, 47, 48a—-c, 49a—b, 50-52. Matsumoto & Obata 1963:

Fig. 144 (see facing page). Baculites reduncus Cobban, 1977. A, G-H. USNM 240423,

paratype. B. USNM 240245, paratype. C. USNM 240246, paratype. D. USNM 240421,

paratype. E-F. USNM 240422, paratype. All from the Upper Campanian Rock River

Formation near Rock River, Wyoming. x1.

THE AMMONITE FAMILY BACULITIDAE

Fig. 144

193

194

ANNALS OF THE SOUTH AFRICAN MUSEUM

64, pl. 19 (fig. 3), text-fig. 158, 165. ?7Dundo 1971, pl. 1 (fig. 5, 5b) (as

Baculites cf. rex). Ward 1978: 1152, pl. 2 (figs 7, 10, text-fig. 2. Haggart

1991, pl. 4 (figs 4-5).

Type. Holotype is the original of Anderson (1958, pl. 49 (fig. 2)) from the

Upper Campanian of Crow Creek, West San Joaquin Valley, California,

CAS type collection.

Occurrence. Upper Campanian—?Lower Maastrichtian, California, Baha

California (Mexico), British Columbia and possibly Arctic Russia.

A B Cc D E

Fig. 145. Baculites reduncus Cobban, 1977. A-B. USNM 411300.

C-E. USNM 413001. Both from the Upper Campanian, basal phosphate bed of the

Annona Chalk at the Okay Quarry, Okay, Howard County, Arkansas.

Both x 1.

* Baculites rioturbioensis Hiinicken, 1965 Fig. 148

(= ?Eubaculites fide Klinger & Kennedy 1993: 190)

Hiinicken 1965: 58, pl. 2 (figs 1-2), pl. 3 (figs 7-8), pl. 7 (fig. 1), pl. 8

(figs 6-8).

Type. Holotype is the original of Hiinicken (1965, pl. 2 (figs 1-2), pl. 3

(fig. 7), pl. 7 (fig. 1), pl. 8 (fig. 7)) from Rio Turbio, in the south-west of

Santa Cruz Province, Argentina, CPC N3005.

Occurrence. Uppermost Campanian or basal Maastrichtian, southern

Argentina.

THE AMMONITE FAMILY BACULITIDAE 195

Baculites roedereri Collignon, 1931

(= B. yokoyamai Tokunaga & Shimizu, 1926 = B. besairiei)

Collignon 1931: 38, pl. 5 (fig. 10-10a), pl. 9 (fig. 17).

Type. Holotype by monotypy is the original of Collignon (1931, pl. 5

(fig. 10, 10a), pl. 9 (fig. 17)) from the Upper Coniacian of Mahagaga,

Madagascar, GD unregistered.

Occurrence. Upper Coniacian, Madagascar.

A B C

Fig. 146. Baculites reesidei Elias, 1933. USNM 507281, from USGS

Mesozoic locality 22182, Bearpaw Shale near Roy, Montana. x 1.

Baculites romanovskii Arkhangel'skij, 1912

Arkhangel'skij 1912: 74, pl. 3 (figs 14-15, 26).

Type. ?

Occurrence. ?

Baculites rotundus Reuss, 1845

(= Gen. et sp. indet. fide Diener 1925: 62; possibly an anisoceratid fide

Fritsch & Schlénbach 1872: 49).

* Baculites rugosus Cobban, 1962a Figs 149-151

Cobban 1962a: 131, pl. 27 (figs 1-9), pl. 28 (figs 1-11), text-fig. Ic.

Kennedy & Cobban 1976, pl. 10 (fig. 7).

Type. Holotype is the original of Cobban (1962a, pl. 27 (figs 8-9)) from the

Upper Campanian, lower part of the Monument Hill Bentonitic Member of

the Pierre Shale, Carter County, Montana, USNM 131120.

Occurrence. Upper Campanian, US Western Interior (Montana, South

Dakota, Wyoming, Colorado and New Mexico).

196

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 147. Baculites rex Anderson, 1958. CAS-28325.01, the holotype,

from locality 28325 (CAS), on south side of Crow Creek, in NE%

SECA S a OL Sa Reais ae

THE AMMONITE FAMILY BACULITIDAE 197

A B

Fig. 148. Baculites rioturbioensis Hiinicken, 1965. Cast of the holotype,

CPC N3005, from the Upper Campanian-Lower Maastrichtian of Rio

Turbio in the south-western part of Santa Cruz Province, Argentina.

sie,

198 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 149. Baculites rugosus Cobban, 1962a. A-B. Paratype. USNM 1311211 from

the lower part of the Monument Hill Bentonitic Member of the Pierre Shale at

USGS Mesozoic locality 23054 in Carter County, Montana.

ale

Baculites sanctae-crucis Pictet & Campiche, 1861

(See Lechites.)

THE AMMONITE FAMILY BACULITIDAE 199

Fig. 150. Baculites rugosus Cobban, 1962a. A-B. USNM 131121c, paratype.

C-D. USNM 131121d, paratype, from the Upper Campanian Exiteloceras jenneyi

zone, Monument Hill Bentonitic Member of the Pierre Shale, USGS Mesozoic

locality 23054 on the north flank of the Black Hills uplift, Carter County, Montana.

All x 1.

* Baculites schencki Matsumoto, 1959 Fig. 104A-C

(= B. capensis)

Matsumoto 1959: 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, 23a-c, 24-25. Matsumoto &

Obata 1963: 38, pl. 12 (figs 1-2), pl. 14 (fig. 1), text-figs 90, 131-133.

Type. Holotype is the original of Matsumoto (1959, pl. 32 (fig. 2a—c), text-

fig. 12a—b) from the Santonian of locality CIT 1034 in the Redding area,

California. UCLA 28830.

Occurrence. Santonian, California and Hokkaido.

200 ANNALS OF THE SOUTH AFRICAN MUSEUM

* Baculites schlueteri Moberg, 1885 Fig. 89

(= nomen dubium fide Kennedy & Christensen 1997: 111, fig. 27f)

Moberg 1885: 40, pl. 4 (fig. 14). Hagg 1954: 55. Kennedy & Christensen

1997: 111, fig. 27f.

Type. Lectotype by subsequent designation of Kennedy & Christensen

(1997: 111) is the original of Moberg (1885, pl. 4 (fig. 14)) from K6pinge,

Sweden.

Occurrence. Campanian, Sweden.

* Baculites scotti Cobban, 1958 Fig. 152

Cobban 1958: 660, pl. 90 (figs 1-9), text-fig. la-e, h. Scott & Cobban

1965: 2, map I-439. Gill & Cobban 1973: 7, fig. 2m. 7Hamama & Kassab

1990: 462, pl. 2 (figs 5-9) ?Kennedy & Cobban 1994a: 107, fig. 8.1-8.3,

8.12, 8.17-8.19. Larson e¢ al. 1997: 26.

Type. Holotype is the original of Cobban (1958, pl. 90 (figs 1-2), text-

fig. la) from the Middle Campanian Pierre Shale, north-east of Pueblo,

Colorado, USNM 108929.

Occurrence. Middle Campanian, US Western Interior (Colorado, Wyoming,

Montana, South Dakota, Nebraska, New Mexico, Utah), and very

doubtfully Egypt.

Baculites sheringomensis Crick, 1924

(= Eubaculites carinatus fide Klinger & Kennedy 1993: 218)

Crick 1924: 139, pl. 9.

Baculites sibiricus Glazunova, 1955

Glazunova 1955: 183, pl. 2 (figs 1, 3).

Type. ?

Occurrence. Siberia.

Baculites singularis Glazunova, 1955

Glazunova 1955: 187, pl. 2 (fig. 2).

Type. ?

Occurrence. Siberia.

Baculites sparsinodosus Collignon, 1969

(= ?B. menabensis)

Collignon 1969: 23, pl. 521 (figs 2052-2054). Klinger & Kennedy 1997,

figs 60, 6la.

Type. Holotype is the original of Collignon (1969, pl. 521 (fig. 2052)) from

the Lower Campanian of Gisement 302, Coupe Ampolypoly-Antsirasira-

Behamotra (Belo sur Tsiribihina), Madagascar, GD 12052.

Occurrence. Lower Campanian, Madagascar and possibly Zululand.

THE AMMONITE FAMILY BACULITIDAE 201

Fig. 151. Baculites rugosus Cobban, 1962a. A-B. USNM 131121h, paratype.

C-E. USNM 131121g, paratype. F. USNM 131121c, paratype. G. USNM 131121b, para-

type. H-I. USNM 131120, the holotype. All from the Monument Hill Bentonitic Member

of the Pierre Shale at USGS Mesozoic locality 23054 in Carter County, Montana. All x 1.

ANNALS OF THE SOUTH AFRICAN MUSEUM

202

Fig. 152

THE AMMONITE FAMILY BACULITIDAE 203

Baculites spillmani Conrad, 1858

(= Eubaculites carinatus fide Klinger & Kennedy 1993: 218)

Conrad 1858: 535, pl. 35 (fig. 24).

Baculites subanceps Haughton, 1925

Haughton 1925: 278, pl. 14 (figs 6-8). Haughton 1926: 24, pl. 3 (figs 6-8).

Matsumoto 1959: 130, pl. 34 (fig. 3), pl. 35 (fig. 1) (as Baculites aff.

B. anceps). Matsumoto & Obata 1963: 59, pl. 20 (fig. 3), text-figs 145-146,

156 (as B. anceps pacificus). Howarth 1965: 368, pl. 5 (fig. 3), pl. 6

(figs 6-7), pl. 7 (fig. 1), text-figs 4, 13-15. Klinger & Lock 1978: 77,

fig. Sa-l. Ward 1978: 1152, pl. 2 (figs 1-4, 8), text-fig. 6 (as B. anceps

pacificus). ?Matsumoto & Miyauchi 1984: 75, text-fig. 13 (as Baculites cf.

B. subanceps pacificus). Luger & Gréschke 1989: 400, pl. 49 (fig. 1), text-

fig. 1S5a—h. Olivero 1992: 56, pl. 1 (figs 21-22) (as B. aff. subanceps

pacificus). Klinger & Kennedy 1997, fig. 131.

Type. Lectotype by the subsequent designation of Howarth (1965: 368) is

the original of Howarth (1965, pl. 6 (fig. 6)), from the Upper

Campanian—Lower Ma astrichtian of Carimba, Angola, SAM-6829.

Occurrence. Upper Campanian (Lower Maastrichtian?), Angola, Cape

Province (Igoda), Egypt, ?Palestine, California, British Columbia,

Hokkaido, James Ross Island, Antarctica.

* Baculites subcircularis Anderson, 1958 Fig. 104H-K

(= ?Fresvillia)

Anderson 1958: 192, pl. 48 (fig. 7).

Type. Holotype by monotypy is the original of Anderson (1958, pl. 48

(fig. 7)) from the Campanian or Maastrichtian near the mouth of Briones

Creek, south of Brentwood, Contra Costa County, California, CAS type

collection .

Occurrence. Campanian or Maastrichtian, California.

Baculites sublaevis Griepenkerl, 1889

(= nom. nud. fide Kennedy 1986c: 191; see also Howarth 1965: 366)

Griepenkerl 1889: 409. |

Baculites subtilis Collignon, 1969

(= B. menabensis)

Collignon 1969: 18, pl. 519 (figs 2042-2044). Klinger & Kennedy 1997,

figs 59d-f, 6lc.

Fig. 152 (see facing page). Baculites scotti Cobban, 1958. A-B. OUMKT7905.

C-E. OUM KT7141. F-G. OUM KT7094. All from the Pierre Shale, Baculites

scotti zone, west-facing bluff, 1.6 km north-north-east of Oral in NW% NW4%

sec. 26, T. 7S., R. 7 E., Fall River County, South Dakota. All x1.

204

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 153. Baculites sweetgrassensis Cobban, 1951. A-B. USNM 43880.

C-E. USNM 433882. F-G. USNM 433884. H-J. USNM 433885. All from the

Middle Coniacian Inoceramus deformis zone at USGS Mesozoic locality 21422 near

Shelby, Toole County, Montana. All x 1.

Type. Holotype is the original of Collignon (1969, pl. 519 (fig. 2042)) from

the Lower Campanian of Gisement 304, Coupe Ampolypoly-Antsirasira-

Behamotra (Belo sur Tsiribihina), Madagascar, GD 12042.

Occurrence. Lower Campanian, Madagascar.

* Baculites suecicus Moberg, 1885 Fig. 91H

Moberg 1885: 34, pl. 4 (fig. 1). Hagg 1935: 59. Kennedy & Christensen

1997: 108, fig 26a.

Type. Holotype by monotypy is the original of Moberg (1885, pl. 4 (fig. 1))

from the Senonian of Kaseberga, Sweden.

Occurrence. Campanian, Sweden.

Baculites sulcatus Baily, 1855

Baily 1855: 457, pl. 11 (fig. 5c only). Woods 1906: 341, pl. 44 (fig. 4). Van

Hoepen 1921: 18, pl. 3 (figs 7-8). Spath 1922: 146. non Collignon 1931:

36, pl. 5 (figs 3, 3a, 4, 4a, 5, 5a, 13, 13a), pl. 9 (fig. 15). non Matsumoto &

Obata 1963: 46, pl. 12 (fig. 6), text-figs 94, 130 (as Baculites n. sp. (?) aff.

B. sulcatus). Klinger & Kennedy 1977: 75, fig. 3b-e, j-l. Klinger &

Kennedy 1997: 111, figs 63-66, 67a-j, 68-77, 78c.

Type. Lectotype by the subsequent designation of Matsumoto & Obata

(1963: 46) is the original of Baily (1855, pl. 11 (fig. 5c)), refigured by

Woods (1906, pl. 44 (fig. 4)) from the Lower Campanian of the Mzamba

Formation at the Mzamba River Estuary, Pondoland, BMNH C35625.

Occurrence. Lower Campanian, Pondoland, subsurface, Richards Bay,

Zululand.

THE AMMONITE FAMILY BACULITIDAE 205

Baculites superstes Pervinquiére, 1910

(= Bochianites fide Wiedmann, 1973: 311).

Pervinquiére 1910: 22, pl. 1 (figs 29-30).

Type. Holotype by monotypy is the original of Pervinquiére (1910, pl. 29

(figs 29-3)) allegedly from the Maastrichtian of Constantine.

SP unregistered.

Occurrence. The species was erroneously attributed to the Maastrichtian,

but is probably Barremian (Durand Delga 1954).

* Baculites sweetgrassensis Cobban, 1951 Fig. 153

Cobban 1951: 820, pl. 118 (figs 6-9), text-figs 1-3. Kauffman 1977: 270,

pl. 29 (fig. 9). Kennedy & Cobban 1991a: 70, pl. 14 (figs 24-25, 29-34,

38-42).

Type. Holotype is the original of Cobban (1951, pl. 118 (figs 8-9)) from the

Middle Coniacian Marias River Shale on the Sweetgrass Arch, Toole

County, Montana, USNM 106983. |

Occurrence. Widely distributed in the Middle Coniacian of the US Western

Interior.

Baculites tanakae Matsumoto & Obata, 1963

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).

?Collignon 1969: 23, pl. 521 (fig. 2055), (as Baculites cf. tanakae).

?Summesberger 1979: 116, pl. 2 (figs 10-13), text-fig. 6 (as Baculites cf.

tanakai).

Type. Holotype is the original of Matsumoto & Obata (1963, pl. 17 (fig. 2),

text-fig. 106), from the Campanian along the main stream of the Haboro,

uppermost part of the Upper Yezo Group, Teshio Province (Hokkaido),

Campanian, GK H4288.

Occurrence. Campanian, Hokkaido, and possibly Austria and Madagascar.

Baculites tanakaeformis Collignon, 1970

(= B. increscens)

Collignon 1970: 2, pl. 607 (figs 2263-2265). Klinger & Kennedy 1997: 132,

fig. 84d-f.

Type. Holotype is the original of Collignon (1970, pl. 607 (fig. 2263)) from

Gisement 326, Middle Campanian of Coupe Ampolypoly-Antsirasira-

Behamotra (Belo sur Tsiribihina), Madagascar, GD 12263.

Occurrence. Middle Campanian, Madagascar.

* Baculites taylorensis Adkins, 1929 Figs 154-155

Adkins 1929: 204, pl. 5 (figs 9-11). Kennedy & Cobban 1993a: 93,

figs 10.1-10.9, 10.11-10.12, 10.16, 10.18-10.19, 11.1-11.2. Kennedy &

Cobban 1993d: 143, pl. 6 (figs 1-9), pl. 7 (figs 1-6, 10-13), text-fig. 8b, d.

206 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 154. Baculites taylorensis Adkins, 1929. A-B. USNM 411296.

C-E. USNM 411295. F-H. USNM 411299. I-K. USNM 411294.

L-N. USNM 411298. All from the basal phosphate bed of the Annona Chalk at the

Okay Quarry, Okay, Howard County, Arkansas. All x 1.

Type. Holotype is the original of Adkins (1929, pl. 5 (figs 10-11)) from the

Middle Campanian of Travis County, 7.5 miles north-east of Austin, Texas.

BEGAT collections.

Occurrence. Middle Campanian, Texas; Campanian, Arkansas.

Baculites teicheri Reyment, 1956

Reyment 1955: 15, pl. 1 (fig. 5a-b), text-fig. 2a. Reyment 1956: 50.

Type. Holotype is the original of Reyment (1955, pl. 1 (fig. 5a—b)) from the

Maastrichtian Nkoporo Shales, Amamgbala, Afikpo Division, Ogoja

Province, Nigeria, BMNH C47413.

Occurrence. Maastrichtian, Nigeria.

THE AMMONITE FAMILY BACULITIDAE 207

Fig. 155. Baculites taylorensis Adkins, 1929. A-B. TMM-21016, paratype.

C-E. TMM-21015, paratype. F-H. TMM-21014, the holotype. All from the Pecan Gap

Chalk, Campanian, along the old Austin-Manor highway on the eastern side of the Big

Walnut Creek Valley, 9 km approximately north-east of Austin, Travis County, Texas.

All-x. 1s

ANNALS OF THE SOUTH AFRICAN MUSEUM

208

Fig. 156

THE AMMONITE FAMILY BACULITIDAE 209

Baculites teres Forbes, 1846

(see Fresvillia)

Forbes 1846: 115, pl. 10 (fig. 5).

‘Euhomaloceras’ teshioense Shimizu, 1935

(nom. nud. fide Matsumoto & Obata 1963: 8).

*Baculites texanus Kennedy & Cobban, 1999 Fig. 156

Kennedy & Cobban 1999: 75, pls 6-7, text-fig. 7.

Type. Holotype is the original of Kennedy & Cobban (1999, pl. 6 (figs 4-6),

text-fig. 7a) from the Campanian Bergstrom Formation at USGS Mesozoic

locality 15535, USNM 475059.

Occurrence. Campanian, Texas, U.S. Gulf Coast Region, US Western

Interior (South Dakota and Colorado).

* Baculites thomi Reeside, 1927 Fig. 157

Reeside 1927b: 13, pl. 12 (figs 9-14). Reeside 1927b: 13 (pars), pl. 10

(figs 9-12) (as B. asper). Cobban & Kennedy 1991la: C5, pls 1-2, text-

fig. 2. Kennedy & Cobban 1991a: 74, text-fig. 26.

Type. Holotype is the original of Reeside (1927b, pl. 12 (figs 9-14)) from

the Santonian Elk Basin Sandstone Member of the Telegraph Creek

Formation, Big Horn County, Montana, USNM 73315.

Occurrence. Upper Santonian, but ranges into the Lower Campanian, US

Western Interior (Montana, Wyoming, New Mexico).

Baculites tippaensis Conrad, 1858

(= Eubaculites carinatus fide Klinger & Kennedy 1993: 218)

Conrad 1858: 334, pl. 35 (fig. 27).

* Baculites trifidilobatus Warren, 1930 Fig. 117I-K

(= B. codyensis fide Kennedy & Cobban 1991: 72). Warren 1930: 65, pl. 5

(figs 1-2, 11-12).

Type. Lectotype here designated is the original of Warren (1930, pl. 5

(figs 1-2)) from the Coniacian, Smoky River Shales, Little Smoky River,

Alberta, Canada, GMUA Ct 462.

Occurrence. Coniacian, Alberta.

Fig. 156 (see facing page). Baculites texanus Kennedy & Cobban, 1999.

A. USNM 475064, paratype. B, D-E. USNM 475061, paratype. C. USNM 475062,

paratype. All from the Bergstrom Formation, Upper Campanian, south-south-west

of Manda, Travis County, Texas.

210

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 157. Baculites thomi Reeside, 1927b. A-C. USNM 433907.

D. USNM 433905. E. USNM 433906. All from the Santonian Desmoscaphites

erdmanni zone near the top of the Marias River Shale at USGS Mesozoic

locality 21419, 12.8 km west of Shelby in the NE% sec. 31, T. 32 N., R. 3 W.,

Toole County, Montana. All x 1.

THE AMMONITE FAMILY BACULITIDAE 211

Baculites tuberculata d’Orbigny, 1842

(= B. incurvatus Dujardin, 1837, fide Schliter 1876: 142; Kennedy 1984:

143-4).

D’Orbigny 1842: 565.

Fig. 158. Baculites undatus Stephenson, 1941. A-C. USNM 449487, from the

Upper Campanian Nostoceras hyatti zone fauna of the Coon Creek Tongue of the

Ripley Formation at Coon Creek, McNairy County, Tennessee. x 1.

Baculites uedae Matsumoto & Obata, 1963

Matsumoto & Obata 1963: 40, pl. 20 (figs 5-7), pl. 21 (figs 1, 3, 6), text-

figs 91-92, 121-129.

212 ANNALS OF THE SOUTH AFRICAN MUSEUM

Type. Holotype is the original of Matsumoto & Obata (1963, pl. 20 (fig. 6))

from the Santonian, main stream of the Detofutamata River, Chikubetsu

area, Teshio Province, Hokkaido, GK H4794.

Occurrence. Santonian, Hokkaido.

*Baculites undatus Stephenson, 1941 Figs 158-159

Stephenson 1941: 405, pl. 79 (figs 5-10). Cobban 1973: 459, figs 2-5.

Cobban 1974: 6, text-fig. 3. Cobban et al. 1992: A7. Kennedy & Cobban

1993c: 424, figs 12.2, 14.18, 14.19, 14.21, 15.13, 15.25-15.27. Kennedy

& Cobban 1994b: 1297, figs 14.31-14.32, 15.5-15.7. Cobban & Kennedy

1994c: B8, pl. 8 (figs 9-11), pl. 9 (figs 1-6). Larson et al. 1997: 31.

Type. Holotype is the original of Stephenson (1941, pl. 79 (figs 5-7)) from

the Campanian Nacatoch Sand near Chatfield, Texas, USNM 77245.

Occurrence. Upper Campanian to Lower Maastrichtian, Texas, Colorado

and New Mexico; also recorded from Tennessee, Mississippi, Delaware and

New Jersey (Owens et al. 1970: 32) and Arkansas, and recently by Cobban

et al. (1992) from the US Western Interior.

*Baculites undulatus d’Orbigny, 1850 Fig. 182H

D’Orbigny 1850: 19, 20, no. 21. Fritsch & Schlénbach 1872: 49. Roman &

Mazeran 1913: 11, pl. 4 (figs 6-8). Matsumoto & Obata 1963: 28, pl. 8

(fig. 4), pl. 9 (figs 1-5), pl. 11 (figs 2-3), text-figs 62-71. Wright 1979:

287, pl. 1 (figs 6-8), pl. 7 (fig. 11). Szdsz 1986: 121, pl. 1 (figs 1-2) (as

B. aff. undulatus). Breton & Bavent 1985: 102, figs 1-3. Nishida ef al.

1992, pl. 6 (figs 4-5). Kennedy et al. 1989: 101, fig. 311. Vasicek 1993:

184, pl. 3 (figs 2-3) (as Baculites cf. undulatus). Summesberger & Kennedy

1996: 135, pl. 18 (figs 8-13, 17-20, 22-25) (as Baculites cf. undulatus).

Type. Holotype by monotypy is the original of d’Orbigny (1850: 190),

figured by Roman & Mazeran (1913, pl. 4 (fig. 6)) and Sornay (1955), from

the Upper Turonian of Uchaux, Vaucluse, France; MNHP, d’Orbigny

Collection no. 6796.

Occurrence. Upper Turonian, England, France, Bohemia, Hokkaido, Trans-

Pecos, Texas and New Mexico, and possibly Lower Turonian, Romania.

Baculites vaalsensis Kennedy & Jagt, 1995

Kennedy & Jagt 1995: 282, figs 4.1-4.18, 5.1-5.17, 6.1-6.9. Kennedy et

al. 1997b: 11, figs 9-10.

Type. Holotype is the original figured’ specimen of Van der Weijden (1943,

pl. 13 (fig. 5)) from the Hervian of Emma Colliery, shaft 1, near Treebeek,

The Netherlands.

Fig. 159 (see facing page). Baculites undatus Stephenson, 1941. A-C. USNM 182430

from the Nacatoch Sand, 1.1 mile north-west of Chatfield, Navarro County, Texas.

D-F. USNM 18243, from the same locality as A-C. G-I. USNM 182427 from the

Nacatoch Sand at USGS Mesozoic locality 762 near Chatfield, Texas.

J. USNM 182437 from the Pierre Shale at USGS Mesozoic locality D1353,

Grand County, Colorado. x 1.

THE AMMONITE FAMILY BACULITIDAE 213

Fig. 159

214 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 160. Baculites vertebralis Lamarck, 1801. A-B. BMNHC70591, from Valognes,

France. C-D. NHMW 7460 from Fresville. Both from the Upper Maastrichtian Calcaire

a Baculites of the Cotentin Peninsula, Manche, France. Both x 1.

THE AMMONITE FAMILY BACULITIDAE 215

Occurrence. Lower Campanian, north-eastern Belgium, and adjacent parts

of The Netherlands, Germany, northern Aquitaine (France) and New Jersey.

Baculites valognensis Bohm, 1891

(nom. dub. fide Howarth 1965: 366; Kennedy 1986a: 61; 1986c: 191)

Bohm 1891: 50, pl. 1 (fig. 13a—b). Nowak 1908: 335, pl. 14 (figs 6-7), text-

figs 1-4 (p. 331), text-figs 6a, 7a—b, 9, 12 (p. 337). Birkelund 1979: 53.

Type. ?

Occurrence. Senonian, northern Alps; in Denmark Birkelund (1979: 53)

recorded it from the Upper Maastrichtian (as B. vertebralis).

Baculites vanhoepeni Venzo, 1936

Venzo 1936: 116 (58), pl. 10 (6) (figs 11-12) (as B. vagina var. van

hoepeni). Klinger & Kennedy 1977: 73, figs 2g-k, 3a-i, 4a—c, 5c. Klinger

& Kennedy 1997: 138, figs 79m, 91-115.

Type. Lectotype, by the subsequent designation of Klinger & Kennedy

(1977: 73), is the original of Venzo (1936, pl. 10 (6) (fig. 1la—b)) from

‘False Bay’, Zululand. DSGUB 1 GO 242.

Occurrence. Middle-?Upper Campanian, Zululand and Natal (Durban

subsurface).

Baculites ventroplanus Collignon, 1969

(= B. menabensis)

Collignon 1969: 20, pl. 520 (figs 2048-2050), p. 24, pl. 521

(figs 2056-2058). Klinger & Kennedy 1997, fig. 58d-f.

Type. Holotype is the original of Collignon (1969, pl. 520 (fig. 2048)) from

the Lower Campanian of Gisement 303, Coupe Ampolypoly-Antsirasira-

Behamotra (Belo sur Tsiribihina), Madagascar, GD 12048.

Occurrence. Lower Campanian, Madagascar.

* Baculites vertebralis Lamarck, 1801 Figs 160-161

Lamarck 1801: 159. Schliiter, 1876: 143 (pars), pl. 39 (figs 12-13), pl. 40

(figs 4, 5, non 6). non Moberg 1885: 38, pl. 4 (figs 8-9). ?Pervinquiére

1907: 93 (pars), non pl. 4 (fig. 9). Pervinquiére 1910: 21. non Taubenhaus

1920: 11. non ?Picard 1929: 444, pl. 9 (figs 5-8), text-figs 3-6. Hagg 1947:

93 (pars). Mikhailov 1951: 48, pl. 1 (figs 4-6), text-figs 15a—b, 16a-b.

Hagg 1954: 55 (pars). Glazunova 1955: 182, pl. 3 (figs 1-3, 6). Naidin &

Shimanskiy 1959: 183, pl. 3 (figs la—b, 3), text-fig. 17-1. non Giers 1964:

256, text-figs 1-2. Tzankov 1982: 17, pl. 3 (fig. 4, 4a). Balan 1982: 209,

pl. 19 (figs 8-9). Kennedy 1986a: 57, pl. 11 (figs 6-11), pl. 12 (figs 1-6),

text-figs 3a—-d, 7d-f, 8. Kennedy 1986c: 187, pl. 19 (figs 1-4, 7-10), pl. 20

(figs 3-5), pl. 28 (figs 2, 7-10, 14-16), pl. 29 (figs 1-15), pl. 30 (figs 1-9),

text-figs lla-b, 12. Kennedy 1986e: 1012, pl. 1 (figs 8-9). Kennedy 1986d,

fig. 10e. Ward eft al. 1991: 1183, fig. 2. Birkelund 1993: 53, pl. 13

(figs 1-11), pl. 16 (figs 1-5), text-fig. 6a, 8d-i. Ward & Kennedy 1993: 51,

ANNALS OF THE SOUTH AFRICAN MUSEUM

216

SRS

Fig. 161

THE AMMONITE FAMILY BACULITIDAE ZAG

figs 43.5-43.7, 43.14, 43.15. (as Baculites cf. B. vertebralis). Kennedy &

Christensen 1997: 110, fig. 261, I-m. Kennedy et al. 1997a: 18, figs 15e-j,

m-o, 16a-f, i-k, 17-18.

Type. Lectotype by subsequent designation of Kennedy (1986a: 57) is the

original of Faujas-Saint-Fond (1799, pl. 21 (figs 2-3)). It is the holotype by

monotypy of B. faujasii Lamarck, 1822, which is thus a subjective

synonym.

Occurrence. Upper Maastrichtian, France, Belgium, Denmark, southern

Sweden, northern Germany, Poland, southern Russia, Maryland, U.S.A.,

and possibly North Africa and Palestine. According to Ward et al. (1991:

1183, fig. 2), B. vertebralis possibly already occurs in the Lower

Maastrichtian of Zumaya.

Baculites vertebralis var. syriacus Conrad, 1852

Conrad 1852: 227, pl. 20 (fig. 121) (as B. syriacus). Picard 1929: 445, pl. 9

(figs 9-10) (as B. vertebralis var. syriacus).

Type. According to Dr S. Rothmann, Jerusalem, most of Conrad’s types are

in the Peabody Museum, Yale University.

Occurrence. Maastrichtian, Palestine.

Baculites vicentei Stinnesbeck, 1986

(= Trachybaculites)

Stinnesbeck 1986: 203, pl. 9 (fig. 4), pl. 10 (figs 3-6), text-fig. 23a-c.

* Baculites yokoyamai Tokunaga & Shimizu, 1926 Fig. 105B-R

Tokunaga & Shimizu 1926: 195, pl. 22 (fig. 5a—b), pl. 26 (fig. 11).

Reyment 1958: 7, pl. 1 (figs 1-2), text-figs 1-2) (as B. ovatus Say?).

Matsumoto & Obata 1963: 30, pl. 8 (fig. 5), pl. 10 (figs 1-6), pl. 11 (figs 1,

4-5), pl. 12 (fig. 3), pl. 14 (fig. 4), text-figs 72-87. Cobban & Scott 1972:

48, pl. 20 (figs 15-21) (as Baculites cf. yokoyamai). Hattin & Siemers,

1978, text-fig. 7.2 (as Baculites cf. yokoyamai). Cobban & Hook 1979: 13,

pl. 4 (figs 9-10). Renz 1982: 105, pl. 34 (figs 3-6), text-fig. 80 (as

B. inornatus). Cobban & Hook 1983: 7, pl. 1 (figs 1-7). Cobban 1983: 16,

pl. 14 (figs 6-8). Cobban 1984: 14, pl. 1 (figs 5-6). Cobban 1986, fig. 3h-i.

Kennedy 1988: 110, pl. 23 (figs 8-10), text-fig. 29c. Kennedy & Cobban

1988: 608, figs 3: 1, 2, 7, 13-14, 18-19. Cobban 1990, pl. 9 (figs 16-22).

Kennedy & Cobban 1991a: 69, pl. 13 (figs 4-10, 17-21, 24-28, 34-37,

41-42), text-fig. 22a. Kennedy et al. 1989: 101, fig. 31e-h. Summesberger

1992: 124, pl. 8 (figs 10-11). Summesberger & Kennedy 1996: 135, pl. 18

(figs 5-7, 16). Klinger & Kennedy 1997: 21, figs 7-11, 12a-i.

Fig. 161 (see facing page). Baculites vertebralis Lamarck, 1801. A-C. IRSNB 10283

(IG 4285 ex Bosquet Collection) from the Upper Maastrichtian Calcaire de Kunraed of

Kunrade, Limburg, The Netherlands. D-F. IRSNB 10284 (IG 8261 ex De Jaer Collection).

Horizon and locality as in A-C. G-I. IRSNB 10278 (IG 6521). Horizon and locality as

above. J-L. IRSNB 10282 (IG6521 ex Ubaghs Collection). Horizon and locality as above.

All x 1.

218

ANNALS OF THE SOUTH AFRICAN MUSEUM

Type. Holotype by monotypy is the original of Tokunaga & Shimizu (1926,

pl. 22 (fig. Sa—b), pl. 26 (fig. 11)) from the lower Futaba Beds in the upper

reaches of the Sakurazawa in Oriki, Hirono-mura, Fukushima prefecture,

north-east Honshu. The type was destroyed during WW II (see Matsumoto

& Obata 1963: 31). A neotype has been designated by Matsumoto & Obata

(1963, pl. 10 (fig. 5)) and is from the Coniacian along the Bannosawa,

Hokkaido, GK H4580.

Occurrence. Lower Turonian to Lower Coniacian, US Western Interior

(Wyoming, Montana, South Dakota, Colorado, Kansas, New Mexico and

Texas); Trans-Pecos, Texas, mid-Turonian of Austria; and Coniacian of

Hokkaido, Venezuela, Zululand and India (Klinger in prep.).

Baculites zhuravlevi Grabovskaya, 1984

Grabovskaya 1984: 84, pl. 12 (figs 4-6), text-fig. 8b-g.

Type. Holotype is the original of Grabovskaya (1984, pl. 12 (fig. 4)) from

the Campanian of Saghalin, BPI 951/20.

Occurrence. Campanian, Saghalin.

Genus Boehmoceras Riedel, 1931

* Boehmoceras arculus (Morton, 1834) Fig. 162A-F

Morton 1834: 44, pl. 15 (figs 1-2). Morton 1834: 45 (as Hamites arculus

var. A). Riedel 1931: 692, pl. 78 (figs 3-6) (as B. léscheri). Ulbrich 1971,

pl. 5 (fig. 4) (as B. loescheri). Summesberger 1979: 119, pl. 2 (figs 15-16,

18), text-figs 9-12) (as B. ldscheri). Kennedy & Wright 1983: 866 (as

Boehmoceras). Kennedy 1983, pl. 2 (fig. 1) (as Boehmoceras sp.).

Schonfeld 1985, pl. 2 (fig. 6). Kennedy 1987: 777, pl. 82 (figs 4-16), text-

fig. 2 (as B. loescheri). Immel 1987: 131 (as B. loescheri). Kennedy &

Cobban 1991b: 182, figs 6: 2, 8, 8: 9-15, 18-22, 9: 1-2, 11-52, 10: 20-21,

24-26, 12: 3. Kennedy & Christensen 1993: 154, figs 3, 41.

Type. Morton (1834) illustrated two specimens; Richards (1968) referred to

two to five co-types; a lectotype was designated by Kennedy & Christensen

(1993: 155) and is the specimen figured as their fig. 3, ANSP collections,

from the “Older Cretaceous strata, Greene County, Alabama’.

Occurrence. Upper Santonian, Germany, Austria, France, Sweden, Gulf

Coast Region, USA.

* Boehmoceras krekeleri (Wegner, 1905) Fig. 163

Wegner 1905: 210, pl. 8 (fig. 2) (as Ancyloceras krekeleri). Riedel 1931:

691. pl. 77 (figs 3-5), pl. 78 (figs 1-2). Summesberger 1979: 118, pl. 2

(fig. 14), text-figs 7-8. Kennedy & Wright 1983: 866. Schonfeld 1985, pl. 2

(fig. 4). Immel 1987: 131. Kennedy 1987: 778, text-fig. 3a—b. Kennedy &

Christensen 1993: 154, figs 2d—-h, 4c, d, h, j.

Type. Holotype by monotypy is the original of Wegner (1905, pl. 8 (fig. 2))

from the Santonian of the Recklinghauser Mergel in the mine Waldersee

near Horneburg, Munster Basin, Germany. Originally deposited in the

THE AMMONITE FAMILY BACULITIDAE 219

Fig. 162. A-F. Boehmoceras arculus (Morton, 1834). A-C. USNM 434130.

D-F. USNM 434120. Both from the Upper Santonian of USGS Mesozoic locality 31555,

Tombigbee Sand Member of the Eutaw Formation, collections from spoil heaps for lock and

dam at Columbus, about 16 m below top of Member, Columbus, Lowndes County,

Mississippi. G-I. Baculites capensis Woods, 1906, USNM 434089, from the Upper

Santonian of USGS Mesozoic locality 31555. All x 1.

220 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 163. Boehmoceras krekeleri (Riedel, 1931). A-B, E. UPST L1 (Université

Paul Sabatier, Toulouse), from the Santonian of Sougraine, Corbieres, Aude,

France. C-D. UPST PMA7-1, Santonian of Chemin de Croutets, Corbiéres,

Aude. Both x 1.

Museum fur Naturkunde, Berlin; apparently the types are lost (see Kennedy

& Christensen 1993: 154).

THE AMMONITE FAMILY BACULITIDAE

Fig. 164. Boehmoceras loescheri Riedel, 1931. All SP unregistered, ex Arnaud

Collection. A-C from Assize N1, Charmant (Charente); D-G, H-J from Assize

M2, Nieul-le Virouil (Charente-Maritime).

All x 1.

221

222 ANNALS OF THE SOUTH AFRICAN MUSEUM

Occurrence. Upper Santonian, northern Germany, France and Gosau,

Austria; subsurface deposits in Sweden.

* Boehmoceras loescheri Riedel, 1931 Fig. 164

(= B. arculus (Morton, 1834)

Riedel 1931: 692, pl. 78 (figs 3-6).

Type. Riedel’s (1931) types were lodged in the former Preussische

Geologische Landesanstalt, later the Zentrale Geologische Anstalt. They are

from various localities in the Upper Santonian of the Munster Basin. As yet,

no lectotype has been designated.

Occurrence. Upper Santonian, northern Germany, Gosau, Austria and

northern Aquitaine, France.

Genus Criobaculites Klinger & Kennedy, 1997

* Criobaculites trabeatus (Morton, 1834) Fig. 165A-D, G-J

Morton 1834: 45, pl. 15 (fig. 3). Cobban & Kennedy 1995: 23,

fig. 17.32-17.35.

Type. Holotype by monotypy is the original of Morton (1834, pl. 15 (fig. 3))

from the Maastrichtian Prairie Bluff Chalk of Alabama, according to

Johnson (1905: 27), in the collections of the Academy of Natural Sciences,

Philadelphia, ANSP 72865.

Occurrence. Maastrichtian, Prairie Bluff Chalk, Alabama.

Criobaculites? sp. :

Cobban & Kennedy (1992c: 684, fig. 1.1-1.4, 3.1 (as Baculites sp.)).

Occurrence. Maastrichtian, Fox Hills Formation, South Dakota.

Genus Eubaculites Spath, 1926

Eubaculites ambindensis Collignon, 1971

(= E. latecarinatus)

Collignon 1971: 13, pl. 646 (fig. 2393). Klinger & Kennedy 1993, fig. 48.

Type. Holotype is the original of Collignon (1971, pl. 646 (fig. 2393)) from

the ‘Lower’ Maastrichtian, zone of Pachydiscus gollevillensis and

Pachydiscus neubergicus of Gisement 504, Mont Ambinda-Mikoboka

(Manera), Madagascar, GD 12393.

Occurrence. Lower Maastrichtian, Madagascar.

Fig. 165 (see facing page). A-D, G-J. Criobaculites trabeatus (Morton, 1834).

ANSP 72865, from the Maastrichtian Prairie Bluff Chalk at Prairie Bluff, Alabama.

E-F. Criobaculites sp. USNM 449803, from the Trail City Member of the Fox Hills

Formation on the Pat Peterson Ranch in the NE% sec. 7, T. 20 N., R. 25 E., Corson

County, South Dakota. K-L. Boehmoceras arculus (Morton, 1834). Lectotype, ANSP

collection, from the ‘older’ Cretaceous strata of Greene County, Alabama, that is, the

Tombigbee Sand Member of the Eutaw Formation. A-F, K-L x 1; G-J x 2.

223

THE AMMONITE FAMILY BACULITIDAE

Fig. 165

224 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 166. Eubaculites occidentalis (Meek, 1862). USNM 1363, one of Meek’s

syntypes and refigured by Matsumoto (1959, pl. 42 (fig. 2)) from “‘Komooks’,

Vancouver Island. x 1.

Baculites binodosus Noetling, 1897

(= ?Eubaculites fide Klinger & Kennedy 1993)

Noetling 1897: 77, pl. 23 (fig. 3). Klinger & Kennedy 1993, fig. 56. Fatmi

& Kennedy 1999: 659, figs 6.3-6.7, 13.19-13.22, 16.3

Type. Holotype by monotypy is the original of Noetling (1897, pl. 23

(fig. 3)) from the Maastrichtian of Des Valley near Khattan, Baluchistan,

India, GSI 3097.

Occurrence. Maastrichtian, Baluchistan, India.

THE AMMONITE FAMILY BACULITIDAE 225

Fig. 167. Eubaculites occidentalis Meek, 1862. CAS-66773.01 from Fresno

County, Coalinga Quadrangle, from c. 50 ft above contact with massive sandstone,

west of Los Gatos Creek, California. x 1.

226 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 168. Fresvillia constricta Kennedy, 1986a. IRSNB 10254 (ex Leriche

Collection, IG 19859) from the Upper Maastrichtian Calcaire 4 Baculites of

Fresville, Manche, France. x1.

Eubaculites carinatus (Morton, 1834)

Morton 1834: 44, pl. 13 (fig. 1). D’Orbigny 1847, pl. 1 (figs 3-7) (as

Baculites lyelli). Conrad 1858: 334, pl. 35 (fig. 27) (as B. tippaensis).

Conrad 1858: 335, pl. 35 (fig. 24) (as B. spillmani). Binckhorst 1861: 42,

pl. 5d (fig. 3a-d) (as B. anceps). Gabb 1864: 80 (ars), pl. 14 (fig. 29-29a);

non pl. 17 (fig. 27, 27a), non pl. 14 (fig. 27b) (as B. chicoensis). Kossmat

1895: 156 (60) (pars), pl. 19 (5) (fig. 14a-b) only (as B. vagina n. var.

simplex). Kossmat 1895: 157 (61) (ars), pl. 19 (5) (fig. 16, ?non fig. 15 (as

B. vagina var. otacodensis). Steinmann 1895: 89, pl. 6 (fig. 4), text-

figs 8-10) (as B. vagina). Kossmat 1897, pl. 6 (fig. 4) (as B. vagina).

Wetzel 1930: 90, pl. 10 (figs 3-4) (as B. vagina). Crick 1924: 139, pl. 9

(figs 1-3) (as B. sheringomensis). Crick 1924: 140, pl. 9 (figs 4-5) (as

Baculites cf. vagina var. otacodensis). Crick 1924: 140, pl. 9 (figs 6-7) (as

Baculites cf. vagina var. simplex). Basse 1931: 20, pl. 2 (figs 6-10) (as

Baculites vagina.) Spath 1940: 49 (pars) (as E. otacodensis). ?0\sson 1944:

104, pl. 16 (figs 3-5), text-fig. 1 (as B. lyelli). Wright 1957: L218, text-

fig. 245, 6a—-b (as Eubaculites otacodensis). Matsumoto 1959: 166, pl. 43

(fig. 6), pl. 44 (figs 1-3), text-figs 84a—b, 85a-b) (as E. ootacodensis).

THE AMMONITE FAMILY BACULITIDAE 220

Fig. 169. Fresvillia teres (Forbes, 1846). BMNH C51152, holotype, the original of

Forbes (1846, pl. 10 (fig. 5)) from the Upper Maastrichtian Valudavur Formation,

Pondicherry, South India. x 1.

Leanza 1964: 95, pl. 1 (figs 1-5), text-fig. 1 (as Eubaculites argentinicus).

Rossi de Garcia & Comacho 1965: 72, pl. 1 (fig. 1) (as Eubaculites sp.).

Brunnschweiler 1966: 27, pl. 1 (figs 9-14), text-figs 9-11 (as

E. ootacodensis). Brunnschweiler 1966: 29, pl. 1 (fig. 7), pl. 2 (figs 1-14),

text-figs 12-14 (as E. vagina). Brunnschweiler 1966: 31, pl. 2 (figs 15-17),

pl. 3 (figs 1-7), text-fig. 15 (as E. kossmati). Brunnschweiler 1966: 32, pl. 3

(figs 8-12), text-fig. 16 (as E. multicostatus). Collignon 1971: 15, pl. 645

(figs 2388-2389) (as B. simplex). Riccardi 1974: 388, pl. 1 (figs 1-7), pl. 2

(figs 1-4), pl. 3 (figs 1-6), pl. 4 (figs 1-7), text-fig. 2 (as E. ootacodensis).

Hinicken & Covacevich 1975: 149, pl. 1 (figs 5-12), pl. 2 (figs 4-9), pl. 3

(figs 1-8), pl. 4 (figs 1-8), pl. 5 (figs 1-4), text-figs 6-28 (as E. lyelli).

Klinger 1976: 90 (pars), pl. 39 (fig. 1), non 3; pl. 41 (figs 1-2), pl. 42

(figs 3, 8), ?non pl. 43 (fig. 1), text-fig. llc (as E. ootacodensis). Kennedy &

Summesberger 1986: 197, pl. 14 (figs 1-5) (as E. lyelli). Kennedy 1986e:

1016, pl. 1 (figs 1-3), pl. 2 (figs 3-8), pl. 3 (figs 2-8, 13-21) (as E. lyelli).

Kennedy 1986c: 194 (as E. ootacodensis). Henderson et al. 1992:

150, figs 6f, g, 14-16, 17a-c, g-j, 18-20). Kennedy & Henderson 19972,

228 ANNALS OF THE SOUTH AFRICAN MUSEUM

text-fig. 5a—c, I-p. Klinger & Kennedy 1993: 218, figs 7a—e, 21-30, 3la-g,

32-35, 36e-f, 37-38, 42a, 52g-h. Kennedy & Cobban 1993b: 62, fig. 3b-i,

I-s, 5. Ward & Kennedy 1993: 53, figs 43.8-43.9, 43.13. Cobban &

Kennedy 1995: 26, figs 14.1, 14.5-14.7, 15.4, 15.6-15.8, 16.13-16.15,

16.23-16.24, 16.28-16.30, 17.52-17.59, 18.1-18.44.

Type. Holotype by monotypy is the original of Morton (1834, pl. 13 (fig. 1))

from the Maastrichtian Prairie Bluff Chalk of Alabama, ANSP’72866.

Occurrence. Lower and Upper Maastrichtian with records from the Biscay

Region and northern Spain, the Netherlands, southern France, Austria,

Zululand, South India, Western Australia, Chile, California, Texas,

Missouri, Mississippi, Alabama and New Jersey.

‘Eubaculiceras’ compressum Brunnschweiler, 1966

(= Eubaculites simplex)

Brunnschweiler 1966: 36, pl. 4 (figs 15-17), pl. 5 (figs 1-3), text-fig. 21.

Type. Holotype is the original of Brunnschweiler (1966, pl. 4 (figs 15-17),

text-fig. 21) from the Upper Maastrichtian, Miria Formation, Western

Australia, CBC 2724.

Occurrence. Upper Maastrichtian, Western Australia.

‘Eubaculiceras’ fastigiatum Brunnschweiler, 1966

(= Eubaculites simplex)

Brunnschweiler 1966: 37, pl. 5 (figs 7-9), text-fig. 22.

Type. Holotype by monotypy is the original of Brunnschweiler (1966, pl. 5

(figs 7-9)), text-fig. 22) from the Upper Maastrichtian, Miria Formation,

Western Australia, CBC 2726.

Occurrence. Upper Maastrichtian, Western Australia.

Fig. 170 (see facing page). A-B. Sciponoceras roto Cieslinski, the original of Collignon’s

(1964: 8, pl. 319 (fig. 1361)) S. baculoides from the Lower Cenomanian, zone of

Mantelliceras martimpreyi, from Gisement 478 (5 km west of Beraketa, Manera), and

Gisement 485, hills of Vohipaly, lower level, Sakaraha. C-D. Hemiptychoceras ambiguus

(Collignon, 1964), the holotype and original of Collignon’s (1964, pl. 319 (fig. 1475))

Baculites (?) (Protobaculites nov. subgen.) ambiguus from the Lower Cenomanian, Zone of

Mantelliceras martimpreyi, Gisement 478, 5 km west of Beraketa (Manera) (see Kennedy &

Wright 1994). E-F. S. roto, the original of Collignon’s (1964: 8, pl. 319 (fig. 1362))

S. baculoides, horizon and locality as in A and B. G-J. S. roto, the original of Collignon’s

(1964: 8, pl. 319 (fig. 1359)) S. baculoides, horizon and locality as in A and B.

H-I. S. roto, the original of Collignon’s (1964: 8, pl. 319 (fig. 1360)) S. baculoides; horizon

and locality as in A and B. K-L. S. roto, the original of Collignon’s (1964: 35, pl. 325

(fig. 1452)) Sciponoceras baculoides, horizon and locality as in A and B.

M-N. Sciponoceras antanimangaensis (Collignon, 1964), the original and holotype of

Collignon’s (1964: 34, pl. 325 (fig. 1451)) Lechites antanimangaensis from the Lower

Cenomanian, zone of Mantelliceras mantelli and M. newboldi from Gisement 362, Signal

Antanimanga I (Mandabe), Madagascar. All in the collections of the Départment des

Sciences de la Terre, Université de Bourgogne, Dijon, France. A-I x 2; K-N ~x 1.

THE AMMONITE FAMILY BACULITIDAE 229

Fig. 170

230 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 171

THE AMMONITE FAMILY BACULITIDAE 231

Fig. 172. Lechites sp. Extreme smooth form of L. gaudini. A-C. BMNH C93267i.

D-F. BMNH C93268. Upper Albian, S. dispar zone, Upper Greensand phosphate bed,

Dorsetshire Gap, 800 m west-south-west of Melcombe Park Dairy Farm, Melcombe,

Hersey, Dorset. All x 1.

Fig. 171 (see facing page). Lechites gaudini. A-E. Macroconch, BMNH C4755

from the Cambridge Greensand, Upper Albian, Stoliczkaia dispar zone, Cambridge.

F. Microconch from the Gault Clay, Upper Albian S. dispar zone, Folkestone, Kent.

G-I. BMNH C34738a, the original of Spath (1941, pl. 72 (fig. 4)).

J-L. BMNH C4754a, the original of Spath (1941, pl. 72 (fig. 7)).

M-O. BMNH C4756, the original of Spath’s (1941, text-fig. 243e-g) L. moreti.

P-R. BMNH C4754a, the holotype of L. communis Spath (1941: 666,

text-fig. 244a-c). S-U. BMNH C34738b.

All x 1.

232 ANNALS OF THE SOUTH AFRICAN MUSEUM

A baad IR LL -M NO

Fig. 173. Tuberolechites regifex Cooper & Kennedy, 1977. A-D. SMC B38412,

paratype. E-G. SMC B97583, paratype. H-K. SMC B97668, paratype.

L-O. SMC 97675, the holotype. All from the Upper Albian, Stoliczkaia dispar

zone, Cambridge Greensand, Cambridge.

All x 1.

Eubaculites kossmati Brunnschweiler, 1966

(= E. carinatus)

Brunnschweiler 1966: 31, pl. 2 (figs 15-17), pl. 3 (figs 1-7), text-fig. 15.

Type. Holotype is the original of Brunnschweiler (1966, pl. 2 (fig. 15-17),

text-fig. 15) from the Upper Maastrichtian, Miria Formation, Western

Australia, CBC 2704.

Occurrence. Upper Maastrichtian, Western Australia.

Eubaculites labyrinthicus (Morton, 1834)

Morton 1834: 44, pl. 13 (fig. 10) (as Baculites labyrinthicus). Stoliczka

1866: 199, pl. 90 (fig. 14, non fig. 15) (as B. vagina var. otacodensis.

Kossmat 1895: 157, pl. 19 (5) (fig. 15, non fig. 16) (as B. vagina var.

otacodensis). Cottreau 1922: 180 (72), pl. 9 (17) (fig. 11-11b) (as B. vagina

var. otacodensis). Spengler 1923: 54, pl. 4 (fig. 9) (as B. vagina). Besairie

THE AMMONITE FAMILY BACULITIDAE 233

1930: 222, pl. 26 (figs 12-13) (as B. vagina). Klinger 1976: 87 (pars),

pl. 42 (fig. 4) (as E. vagina). Kennedy & Henderson 1992: 716, text-

fig. Sd-k. Klinger & Kennedy 1993: 194, figs 1-6, 7f-h, 36a-d, 52a-c.

Cobban & Kennedy 1995: 23, figs 14.2, 15.2-15.3, 17.36-17.51.

Type. There are two syntypes, the figured specimen in Morton (1834, pl. 13

(fig. 10)), ANSP 72868, and an unfigured specimen, ANSP 72869, both

from the Prairie Bluff Chalk of Alabama. Cobban & Kennedy (1995: 23)

designated Morton’s figured specimen as the lectotype.

Occurrence. Lower Maastrichtian, Alabama, New Jersey, South India,

Zululand, and Madagascar.

A B

Fig. 174. Lechites? sp. OUMKY303, Utatur Group. Odium, South India. Ex

Geological Survey of India Collections, GSI142 L3/184. This appears to be the

‘Baculites gaudini’ of Stoliczka (1865: 199). x 1.

234 ANNALS OF THE SOUTH AFRICAN MUSEUM

Eubaculites latecarinatus (Brunnschweiler, 1966)

Brunnschweiler 1966: 33, pl. 3 (figs 13-14), pl. 4 (figs 1-5), text-

figs 17-18. Brunnschweiler 1966: 35, pl. 4 (figs 11-14), text-fig. 20

(as Giralites quadrisulcatus). Collignon 1971: 18, pl. 646 (fig. 2393) (as

Eubaculites ambindensis). Collignon 1971: 15, pl. 645 (fig. 2390) (as

Baculites occidentalis). Klinger 1976: 91, pl. 40 (figs 1-2), pl. 41 (fig. 3),

pl. 42 (figs 2, 6), pl. 43 (figs 3-4), text-fig. 1ld-e. Klinger & Kennedy

1980: 296, figs 2-4, 5d. Henderson et al. 1992: 159, figs 221-n, 23n-p.

Klinger & Kennedy 1993: 238, figs 26a, 39-41, 42b-c, 43-49, 50a, 53a.

Kennedy et al. 1997a: 20, figs 15a—-d, k-1, 16g—h, 16.

Type. Holotype is the original of Brunnschweiler (1966, pl. 4 (figs 2-4))

from the Lower Maastrichtian, nodule bed at the top of the Korojong

Calcarenite, Western Australia, CBC 2718.

Occurrence. Lower Maastrichtian, Western Australia, Madagascar,

Zululand, and offshore Alphard Group, southern Cape Province; and Upper

Maastrichtian, Severn Formation, Maryland.

Eubaculites multicostatus Brunnschweiler, 1966

(= Eubaculites carinatus)

Brunnschweiler 1966: 32, pl. 3 (figs 8-12), text-fig. 16.

Type. Holotype is the original of Brunnschweiler (1966, pl. 3 (figs 10-12),

text-fig. 16) from the Upper Maastrichtian, Miria Formation, Western

Australia, CBC 2710.

Occurrence. Upper Maastrichtian, Western Australia.

* EFubaculites occidentalis (Meek, 1862) Figs 107A-D, 166-167

Meek 1862: 316. Anderson & Hanna 1935: 24, pl. 8 (figs 3-4). Usher

1952: 98, pl. 28 (fig. 1), pl. 31 (fig. 19), text-fig. 4. Matsumoto 1959: 150,

pl. 35 (figs 2a-d, 3a-d), pl. 36 (fig. la-d), pl. 41 (fig. la-d), pl. 42

(figs la-c, 2a-c), text-figs 64, 65a—b, 66, 67-71. Obata & Matsumoto 1963:

82, pl. 23 (fig. 3), pl. 25 (fig. 1), pl. 26 (figs 1-3), pl. 27 (figs 2-5, 8), text-

figs 172-186. Jones 1963: 28, pl. 11 (fig. 13). non Collignon 1971: 15,

pl. 645 (fig. 2390) (= E. simplex). ?Dundo 1971, pl. 1 (fig. 10) (as B. cf.

occidentalis). Ward 1978: 1153, pl. 2 (figs 5-6, 9), text-fig. 2. Haggart

1991, pl. 5 (fig. 5). (All as Baculites occidentalis.)

Type. Meek (1862) based this species on several specimens; as yet, no

lectotype has been designated (see Matsumoto 1959: 150).

Occurrence. Upper Campanian, California, Baha California

(Mexico), British Columbia, southern Alaska, Honshu, and possibly Arctic

Russia.

Fig. 175 (see facing page). Pseudobaculites natosini (Robinson, 1945). USNM 458244 from

the Bearpaw Shale at USGS Mesozoic locality D1442 in the NW% SW sec. 9, T. 27 N.,

R. 20 E., Blaine County, Montana. x 0.74

THE AMMONITE FAMILY BACULITIDAE 235

Fig. 175

ANNALS OF THE SOUTH AFRICAN MUSEUM

236

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Ul BOSPA Artfedo] SIOZOSAW SOSA Je SYS STMT Oy) WO EYZYSpWNSN “V “(Sr6T “UOsUIgoY) mIsoMmU sajynovgopnasq “OL “31d

THE AMMONITE FAMILY BACULITIDAE 237

‘Giralites’ quadrisulcatus Brunnschweiler, 1966

(= Eubaculites latecarinatus)

Brunnschweiler 1966: 35, pl. 4 (figs 11-14), text-fig. 20.

Type. Holotype is the original of Brunnschweiler (1966, pl. 4 (figs 12-14),

text-fig. 20) from the Lower Maastrichtian, Korojong Calcarenite, Western

Australia, CBC 2722.

Occurrence. Maastrichtian, Western Australia.

‘Cardabites’ scimitar Brunnschweiler, 1966

(= Eubaculites simplex)

Brunnschweiler 1966: 38, pl. 5 (figs 16-21), text-fig. 24.

Type. Holotype is the original of Brunnschweiler (1966, pl. 5 (figs 16-18),

text-fig. 24) from the Upper Maastrichtian, Miria Formation, Western

Australia, CBC 2729.

Occurrence. Upper Maastrichtian, Western Australia.

Eudaculites simplex (Kossmat, 1895)

Kossmat 1895: 156 (pars), pl. 19 (fig. 13a—-b), non pl. 19 (fig. 14a—-b) (as

B. vagina var. simplex). non Collignon 1938: 105. Hiinicken 1965: 64, pl. 2

(figs 3-4), pl. 3 (figs 5-6), pl. 5 (fig. 5), pl. 6 (figs 6-7), pl. 8 (figs 2, 5)

(as Baculites sp.). Brunnschweiler 1966: 34, pl. 4 (figs 6-10), text-fig. 19

(as Giralites simplex). Brunnschweiler 1966: 36, pl. 4 (figs 15-17),pl. 5

(figs 1-3), text-fig. 21 (as Eubaculiceras compressum). Brunnschweiler

1966: 37, pl. 5 (figs 7-9), text-fig. 22 (as Eubaculiceras fastigiatum).

Brunnschweiler 1966: 38, pl. 5 (figs 12-15), text-fig. 23 (as Cardabites

tabulatus). Brunnschweiler 1966: 38, pl. 5 (figs 16-21), text-fig. 24

(as Cardabites scimitar). Collignon 1971: 15, pl. 545 (fig. 2390)

(as B. occidentalis). Klinger 1976: 92 (pars) pl. 42 (fig. 10), pl. 43 (fig. 2,

14) (as Eubaculites compressum). Henderson et al. 1992: 153,

figs 21, 22a-k, o-p, 23a-f, k-m, 24-25. Kennedy & Henderson 1992, text-

fig. 6d-f. Klinger & Kennedy 1993: 252, figs 7i-k, 50b, 51, 52d-f, i-n,

53b, 54-55.

Type. Lectotype by the subsequent designation of Kennedy (1986c: 195) is

the original of Kossmat (1895, pl. 19 (figs 13a-b)) from the Maastrichtian

Arrialoor Group of Otacod, South India, GSI 14819.

Occurrence. Maastrichtian, South India, Western Australia, Patagonia and

Zululand.

‘Cardabites’ tabulatus Brunnschweiler, 1966

(= Eubaculites simplex)

Brunnschweiler 1966: 38, pl. 5 (figs 12-15), text-fig. 23.

Type. Holotype is the original of Brunnschweiler (1966, pl. 5 (figs 10-12),

text-fig. 23) from the Upper Maastrichtian, Miria Formation, Western

Australia, CBC 2727.

Occurrence. Upper Maastrichtian, Western Australia.

238 ANNALS OF THE SOUTH AFRICAN MUSEUM

THE AMMONITE FAMILY BACULITIDAE 239

* Eubaculites vagina (Forbes, 1846) Figs 39-41

Forbes 1846: 114, pl. 10 (fig. 4) (as Baculites vagina). D’Orbigny 1847,

pl. 3 (figs 3-6) (as Baculites ornatus). Stoliczka 1866: 198 (pars), pl. 91

(figs 1-6). Kossmat 1895: 155 (59), pl. 19 (5S) (fig. 17) (as Baculites vagina

(a) Typische Form). Crick 1898: 78, pl. 17 (fig. 5). non Basse 1931: 20, pl.

2 (figs 6-10). Collignon 1971: 15, pl. 645 (fig. 2391), p. 18, pl. 646

(fig. 2394). Klinger 1976: 87, pl. 35 (figs 1-4), pl. 36 (figs 1-4), pl. 37

(figs 1-5), pl. 38 (figs 1-3), pl. 39 (fig. 2), pl. 42 (fig. 1, non figs 4, 7 , 9,

11), pl. 43 (figs 5-12). Kennedy 1977, text-fig. 31.1-31.2. Klinger &

Kennedy 1980: 299, text-fig. Sa-c. Kennedy 1989, text-fig. 17i-j. Kennedy

& Henderson 1992: 712, pl. 5 (figs 10-13), pl. 7 (figs 1-18), pl. 8

(figs 1-12), pl. 9 (figs 1-13), pl. 10 (figs 1-3), text-fig. 1c, f. Klinger &

Kennedy 1993: 203, figs 8-20.

Type. Lectotype by the subsequent designation of Kennedy & Henderson

(1992: 713) is the original of Forbes (1846, pl. 10 (fig. 4)) from the Upper

Maastrichtian Valudavur Formation of Pondicherry, South India,

BMNH C49726.

Occurrence. Maastrichtian of South India and Madagascar.

Genus Fresvillia Kennedy, 1986

* Fresvillia constricta Kennedy, 1986 Fig. 168

Kennedy 1986a: 62, pl. 14 (figs 39-42), text-fig. 10a.

Type. Holotype by monotypy, is the original of Kennedy (1986a, pl. 14

(figs 39-42), text-fig. 10a) from the Upper Maastrichtian Calcaire a

Baculites, Fresville, Manche, France, IRSNB 10254 (ex Leriche collection

IG 19859).

Occurrence. Upper Maastrichtian, France.

* Fresvillia teres (Forbes, 1846) Fig. 169

Forbes 1846: 115, pl. 10 (fig. 5). D’Orbigny 1850: 215. Stoliczka 1866:

197 (ars), pl. 90 (fig. 12 only, non 13). Kossmat 1895: 155 (59). Kossmat

1897: 64. Spath 1953: 16 (all as Baculites teres). Matsumoto 1959: 163,

pl. 45 (figs 5-6), text-figs 82a—-c, 83 (as ?Baculites aff. B. teres). Jones

1963: 29, pl. 16 (figs 10-12, 14), text-fig. 14 (as Baculites aff. B. teres).

Kennedy 1986a: 62. Stinnesbeck 1986: 204 (as ?B. teres). Kennedy &

Henderson 1992: 718, pl. 5 (figs 14-17), text-fig. 2c.

Fig. 177 (see facing page). A-F. Pseudobaculites nodosus Cobban, 1952.

A-C. USNM 108209, the holotype from USGS Mesozoic locality 21096.

D-F. USNM 108214, paratype, from USGS Mesozoic locality 23100.

G-M. Pseudobaculites wyomingensis Cobban, 1952. G-I. USNM 108217, the holotype from

USGS Mesozoic locality 21748. J. USNM 433904 from USGS Mesozoic locality 4991.

K-L. USNM 433903 from USGS Mesozoic locality D3797. M. USNM 108219. All from

the Upper Coniacian Scaphites depressus zone in the Cody Shale of Fremont County,

Wyoming. All x 1.

240

ANNALS OF THE SOUTH AFRICAN MUSEUM

ait it a

THE AMMONITE FAMILY BACULITIDAE 241

Type. Holotype by monotypy is the original of Forbes (1846, pl. 10 (fig. 5))

from the Maastrichtian of the Valudavur Formation, south India,

BMNHC51152 (ex Geological Society Collection R10890).

Occurrence. Maastrichtian of South India, California and Alaska, and

possibly Chile.

Genus Lechites Nowak, 1908

* Lechites antanimangaensis Collignon, 1964 | Fig. 170M-N

(? =Sciponoceras fide Cooper & Kennedy 1977: 652-3)

Collignon 1964: 34, pl. 325 (fig. 1451).

Type. Holotype by monotypy is the original of Collignon (1964, pl. 325

(fig. 1451)) from the Lower Cenomanian of Signal Antanimanga I

(Mandabe), Madagascar. GD 11451.

Occurrence. Lower Cenomanian, Madagascar.

Lechites campichei Renz, 1968

(= L. gaudini fide Cooper & Kennedy 1977: 651-652)

Pictet & Campiche 1861: 112, pl. 55 (fig. 9) (as B. gaudini). Boule et al.

1907: 64, pl. 15 (fig. 1) (as B. gaudini). Spath 1941: 665, pl. 72 (fig. 5) (as

L. gaudini). Renz 1968: 82, pl. 17 (figs 8a—c, 9a-c, 10a—b), text-fig. 29m.

Type. Holotype is the original of Pictet & Campiche (1861, pl. 55 (fig. 9)),

refigured by Renz (1968, pl. 17 (fig. 9)), from the Upper Albian of Ste

Croix, Switzerland, MGL L40020.

Occurrence. Upper Albian, Switzerland, Cambridge Greensand of England

and Madagascar.

Lechites comanchensis (Adkins, 1920)

Adkins 1920: 51, pl. 2 (figs 20-22). Adkins 1928: 207, pl. 20 (fig. 9) (as

Cyrtochilus ?comanchensis). Clark 1965: 23, pl. 3 (figs 1-2, 5-11), pl. 4

(fig. 16), text-fig. 6a.

Type. Holotype is the original of Clark 1965, pl. 3 (figs 8-11), BEG 20997.

Occurrence. Upper Albian of Texas.

Fig. 178 (see facing page). Sciponoceras baculoides (Mantell, 1822). A-B. The

lectotype (larger specimen), a microconch, and a paralectotype (smaller specimen),

the original of Mantell (1822 pl. 23 (fig. 6)), BMNH C8612 from the Lower Chalk of

Sussex. C. Paralectotype BMNH C36576a, the original of Mantell (1822, pl. 23

(fig. 7)) from the same locality and horizon as A and B. D-E. SMC B21312,

a microconch from the Lower Chalk of Ventnor, Isle of Wight. F-H. SMC B21321, a

macroconch from the same locality and horizon as D and E. I-J. SMC B21320, a

macroconch. Details as for D and E.

Alias

ANNALS OF THE SOUTH AFRICAN MUSEUM

242

Fig. 179

THE AMMONITE FAMILY BACULITIDAE 243

* Lechites communis Spath, 1941 Fig. 171P-R

(= L. gaudini fide Cooper & Kennedy 1977: 648)

Spath 1941: 666, text-fig. 244a-e. Renz 1968: 81, pl. 17 (figs 6a—c, 13a-b),

text-fig. 29c-d. Nagy 1971: 17, pl. 1 (figs 7-8). Horvath 1983, pl. 1

(fig. 5). Mitiu 1984: 85, pl. 2 (figs 12-13).

Type. Holotype is the original of Spath (1941, text-fig. 244a-c) from the

Upper Albian Cambridge Greensand, BMNH C4754b.

Occurrence. Upper Albian, Cambridge Greensand, England, Switzerland,

Hungary.

Lechites fasciatus Scholz, 1971

Scholz 1971: 431, figs 1-2.

Type. ?

Occurrence. Upper Albian, Hungary.

* Lechites gaudini (Pictet & Campiche, 1861) Figs 171-172

Pictet & Campiche 1861: 112, pl. 55 (figs 5-9) (as B. gaudini). Bose 1923:

157 (as Baculites cfr. gaudini). ?Collignon 1933: 73, pl. 5 (fig. 8, 8a). Spath

1941: 662, pl. 72 (figs 4-7, 9-10), text-fig. 242 (cum. synon.). Spath 1941:

66, text-fig. 244 (as L. communis). Breistroffer 1947: 94 (as L. gaudini

var. raricosta). ?Wiedmann & Dieni 1968: 64, pl. 6 (fig. 10), text-fig. 37

(as L. italicus). Renz 1968: 82, pl. 17 (figs 11-12, 14-15), text-figs 29b,

g—-h, k. (as L. vraconensis). ?Scholz 1971: 431, figs 1-2 (as L. fasciata).

Nagy 1971: 17, pl. 1 (figs 5-6). Cooper & Kennedy 1977: 644, figs 1

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

Scholz 1978, pl. 3 (figs 1, 8). Scholz 1979: 12, pl. 1 (figs 1-9), text-

fig. 5a-—b. Chiriac 1981: 63, pl. 3 (fig. 6a-c), text-fig. 20. Horvath 1983,

pl. 1 (fig. 2). Mitiu 1984, pl. 3 (figs 6-9). Immel & Seyed-Emami 1985:

112, pl. 7 (fig. 11). Delamette 1988, pl. 11 (fig. 6). Immel 1987: 128.

Follmi 1989: 120, pl. 4 (fig. 1) (as Lechites aff. gaudini). Henderson 1990:

129, fig. 12a—m. Ivanov 1991, pl. 4 (fig. 11). Moncrieff & Kelly 1993: 5.

Gale et al. 1996: 577, figs 22a-c, 27e-i, l-o. Klinger & Kennedy 1997: 7,

figs 2a-c, 3.

Fig. 179 (see facing page). Sciponoceras baculoides (Mantell, 1822). A-D. BMNH C422, a

macroconch, the original of Crick (1896: 80) (see also Wright & Kennedy 1995, pl. 95

(fig. 5)) from the phosphatic Middle Cenomanian, A. rhotomagense zone, Turrilites acutus

subzone fauna of the Chalk Basement Bed at Chardstock, Devon. E-F. BMNH C44005a, a

macroconch and a syntype of Baculites obliquatus J. de C. Sowerby (1828, pl. 592 (fig. 2

left)) (see also Wright & Kennedy 1995, pl. 95 (fig. 3)) from the Lower Chalk, Middle

Cenomanian, A. rhotomagense zone, T. costatus subzone, Hamsey near Lewes, Sussex.

G-H. SMC B21311, macroconch, horizon as for E and F, Ventnor, Isle of Wight.

I. BGS-GSM37740, horizon and locality as for G and H (see also Wright & Kennedy 1995,

pl. 97 (fig. 1)). J-K. BGS-GSM92186, horizon and locality as for G and H (see also Wright

& Kennedy 1995, pl. 96 (fig. 7)). All x 1.

ANNALS OF THE SOUTH AFRICAN MUSEUM

244

Fig. 180

THE AMMONITE FAMILY BACULITIDAE 245

Fig. 181. Sciponoceras bohemicum (Fritsch, 1872). A-C. BGS-GSM108888 from

the Chalk Rock, Upper Turonian, Hitch Wood near Hitchin, Hertfordshire.

D-F. BGS-Zr7803 from the Chalk Rock, Upper Turonian, Kensworth near

Dunstable, Bedfordshire. Both x 1.

Fig. 180 (see facing page). Sciponoceras baculoides (Mantell, 1822). Muséum d’histoire

Naturelle Paris Collections, ex D’Orbigny Collection no. 6136. These specimens are the

basis of the illustrations of D’Orbigny (1842, pl. 138 (figs 6-11)). A-U are from the Middle

Cenomanian of Rouen, Seine Maritime, France; V-Y, is, by its preservation from the same

horizon at Cap Blanc Nez, Boulonnais, France. All x 1.

Fig. 182 (see overleaf). A-G, I-M. Sciponoceras bohemicum (Fritsch, 1872).

A-C. BGS-GSM115260 from the Chalk Rock, Upper Turonian, Kensworth near Dunstable,

Bedfordshire. D-F. BGS-Yc7286, horizon and locality as A-C. G. SMC B4142, horizon as

A-C, from Cuckhamsley, Farnborough, Berkshire; the original of Woods (1896, pl. 2

(fig. 9))). I-K. BGS-Yc7523c from the Chalk Rock, Upper Turonian, Hitch Wood near

Hitchin, Hertfordshire. L-M. BGS-Yc7303. Horizon and locality as A-C. H. Baculites

undulatus d’Orbigny, 1850. The holotype, Muséum d’Histoire Naturelle, Paris Collection,

ex d’Orbigny Collection, from the Upper Turonian of Uchaux, Vaucluse, France.

A-G, I-M x 1, H x 2.

Fig. 183 (see overleaf). Sciponoceras bohemicum (Fritsch, 1872). A-C. BMNH C33427,

Upper Turonian, Chalk Rock, Hill End, near Hitchin, Hertfordshire. D. GPIB80. The

original of Schliter’s (1876, pl. 39 (fig. 1)) Baculites cf. bohemicum from the Upper

Turonian Scaphiten-Planer of Oerlinghausen, Teutoburgerwald, Germany.

E-G. BMNH C79507, Upper Turonian, Chalk Rock, Hitch Wood, near Hitchin,

Hertfordshire. H-J. Horizon and locality as for A-C. K-M. BGS-Zr7804, Upper

Turonian, Chalk Rock, Kentworth, near Dunstable, Bedfordshire. N-P. BMNH C73688.

Locality as for K-M. All x 1.

ANNALS OF THE SOUTH AFRICAN MUSEUM

246

Fig. 182

247

THE AMMONITE FAMILY BACULITIDAE

Fig. 183

248

ANNALS OF THE SOUTH AFRICAN MUSEUM

Type. Lectotype by the subsequent designation of Spath (1941: 663) is the

original of Pictet & Campiche (1861, pl. 55 (fig. 5a-c)) from the upper

Albian of St Croix, Switzerland, MGL L21271.

Occurrence. Upper Albian, England, France, Switzerland, Hungary,

Romania, Sardinia, North Africa, Madagascar, Zululand, South India,

Hokkaido, Mexico and Antarctica.

Lechites imlayi Leanza, 1970

(= Rossalites fide Aguirre Urreta & Riccardi 1988: 600)

Leanza 1970: 211, fig. 10.1-10.3.

Lechites italicus Wiedmann & Dieni, 1968

(= 2L. gaudini fide Cooper & Kennedy 1977)

Wiedmann & Dieni 1968: 64, pl. 6 (fig. 10), text-fig. 37.

Type. Holotype by monotypy is the original of Wiedmann & Dieni (1968,

pl. 6 (fig. 10), text-fig. 37) from the condensed Upper Albian of Orosei,

Sardinia (coll. Sardegna), IGP 59.

Occurrence. Upper Albian, Sardinia.

* Lechites moreti Breistroffer, 1936 Fig. 171M-O

Pictet & Campiche 1861: 112, pl. 55 (fig. 10-11) (as B. gaudini).

Breistroffer 1936: 66. Spath 1941: 665, pl. 72 (fig. 8), text-fig. 243.

Breistroffer 1947: 62. Wiedmann 1962a: 184, pl. 10 (fig. 1), text-fig. 45.

Renz 1968: 81, pl. 16 (figs 10a—c, 12-13), text-fig. 29a, i. Scholz 1979: 14,

pl. 1 (fig. 10), text-fig. 5c. (as L. gaudini moreti). Chiriac 1981: 63, pl. 3

(figs 7-9). Horvath 1983, pl. 2 (fig. 11). Delanoy & Latil 1988, pl. 1

(fig. 4).

Type. Lectotype by subsequent designation Spath (1941: 665) is the original

of Pictet & Campiche (1861, pl. 55 (fig. 10a—d)), refigured by Renz (1968,

pl. 16 (fig. 10), text-fig. 29a, 1) from the Upper Albian of Ste Croix,

Switzerland, MGL L40016.

Occurrence. Upper Albian, Cambridge Greensand, England, France,

Switzerland, Spain, Sardinia, Hungary, North Africa.

Lechites raricostatus Breistroffer, 1947

(= L. gaudini fide Cooper & Kennedy 1977: 645)

Pictet & Campiche 1861: 112, pl. 55 (fig. 8) (as B. gaudini). Collignon

1929: 70, pl. 7 (fig. 12) (as B. gaudini). Breistroffer 1947: 94 (as

Lechites gaudini var. raricostata). Renz 1968: 82, pl. 16 (fig. 1la-—b),

pl. 17 (fig. 7a-c), text-fig. 291. Nagy 1971: 18 (as Lechites aff.

raricostatus).

Type. Lectotype is the original of Pictet & Campiche (1861, pl. 55 (fig. 8)),

by subsequent designation of Renz and refigured by him (Renz 1968, pl. 17

(fig. 7), text-fig. 291) from the Upper Albian of St Croix, Switzerland,

MGL L40021.

THE AMMONITE FAMILY BACULITIDAE 249

Fig. 184. A-I. Sciponoceras glaessneri Wright, 1963. R. Henderson Collection L.842 from

the Middle Cenomanian of Bathurst Island, Northern Australia. All x 1.

250 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 185. Sciponoceras gracile (Shumard, 1860). Microconchs. A-B. USNM 411539

from Texcrete Quarries, Dallas County. C-E. USNM 411542 from the same locality.

F-H. USNM 411541 from Newton Branch, 3.75 miles south of Britton on old

Britton-Midlothian Road, Ellis County. I-J. USNM 411540, from 1.5 to 1.8 miles south-

east of Britton on Rogers Farm, Ellis County. All specimens from the Upper Cenomanian

Sciponoceras gracile zone fauna of the Britton Formation in north-east Texas. All x 1.

Occurrence. Upper Albian, Cambridge Greensand, England, Switzerland,

Hungary and Madagascar.

Lechites (?) sanctaecrucis (Pictet & Campiche, 1861)

(= ?Anisoceratid; see also Spath 1939: 573; 1941: 661 footnote)

Pictet & Campiche 1861: 109, pl. 55 (fig. la-h, 2-4) (as Baculites

sanctae-crucis). Tavani 1942: 30 (116) (as Lechites cfr. sanctae-crucis).

Type. Lectotype, by the subsequent designation of Spath (1939: 573) is

the original of Pictet & Campiche (1861, pl. 55 (fig. la—c)) from the

Middle? Albian of Switzerland.

Occurrence. Middle Albian of Switzerland and, doubtfully, Somalia.

Lechites varicostatus Chiriac, 1981

Chiriac 1981: 64, pl. 3 (figs 10, 1la-c).

Type. Holotype is the original of Chiriac (1981, pl. 3 (figs 10, 11la-c)),

from the Upper Albian, probably dispar zone of Southern Dobrogea,

IGG 16925.

Occurrence. Upper Albian, Romania.

Lechites vraconensis Renz, 1968

(= L. gaudini fide Cooper & Kennedy 1977)

Renz 1968: 82, pl. 17 (fig. 1la-c, 12a—b, 14a-b, 15a-—c), text-fig. 29b, g—h,

THE AMMONITE FAMILY BACULITIDAE 251

Type. Holotype is the original of Renz (1968, pl. 17 (fig. lla-c), text-

fig. 29g-h) from the Upper Albian of La Vraconne, Switzerland,

MGL L40024.

Occurrence. Upper Albian, Switzerland.

Subgenus Lechites (Tuberolechites) Cooper & Kennedy, 1977

* Lechites (Tuberolechites) regifex Cooper & Kennedy, 1977 Fig. 173

Cooper & Kennedy 1977: 654, fig. 8, 1-15. Scholz 1979: 15, pl. 1

(figs 11-16) (as L. gaudini nodosus).

Type. Holotype is the original of Cooper & Kennedy (1977, fig. 8 (1-4))

from the Upper Albian of the Cambridge Greensand, England, SM B97675.

Occurrence. Upper Albian, Cambridge Greensand, England, Salazac

(Gard), France, and Hungary.

Genus Pseudobaculites Cobban, 1952

* Pseudobaculites natosini (Robinson, 1945) Figs 175-176

Robinson 1945: 52, pl. 1 (figs 5-6). Cobban & Kennedy 1994a: C2,

pls 1-2. Larson et al. 1997: 37.

Type. Holotype by monotypy is the original of Robinson (1945, pl. 1

(figs 5-6)) from the Bearpaw Shale, south-western Saskatchewan,

GSC 9119.

Occurrence. Upper Campanian and Lower Maastrichtian, southern

Saskatchewan, Montana, Wyoming, Colorado.

* Pseudobaculités nodosus Cobban, 1952 Fig. 177A-F

Cobban 1952: 759, pl. 110 (figs 1-10, 17-18). Kennedy & Cobban 1991a:

78, pl. 17 (figs 9-14), text-fig. 27.

Type. Holotype is the original of Cobban (1952, pl. 110 (figs 1-4)) from the

Upper Coniacian, 152 m above base of the Cody Shale near Lander,

Fremont County, Wyoming, USNM 108209.

Occurrence. Upper Coniacian of western Wyoming only.

* Pseudobaculites wyomingensis Cobban, 1952 Fig. 177G-M

Cobban 1952: 760, pl. 110 (figs 11-16, 19-21). Kennedy & Cobban 1991a:

79, pl. 14 (figs 35-37), pl. 17 (figs 15-23).

Type. Holotype is the original of Cobban (1952, pl. 110 (figs 11-13)) from

the Upper Coniacian, about 457 m above the base of the Cody shale near

Dubois, Fremont County, Wyoming, USNM 108217.

Occurrence. Upper Coniacian, US Western Interior (Wyoming, Utah and

possibly south-eastern Colorado).

252 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 186

THE AMMONITE FAMILY BACULITIDAE 253

Fig. 187. Sciponoceras kossmati (Nowak, 1908), the original of Matsumoto & Obata (1963,

pl. 4 (fig. 1)), Kyushu University Collections, GK H4335, from the Upper Cenomanian

Mikasa Formation, Ikushumbets Valley, Ishikari Province, Hokkaido, Japan. x 1.

Fig. 186 (see facing page). Sciponoceras gracile (Shumard, 1860). Macroconchs.

A-D. USNM 411544 from Texcrete Quarries, Dallas County. E. USNM 41537 from 1.5 to

1.8 miles south-east of Britton on Rogers Farm, Ellis County. F. USNM 411543 from

Texcrete Quarries, Dallas County. G. USNM 411538 from the same locality as E. All the

specimens are from the Upper Cenomanian Sciponoceras gracile zone fauna of the Britton

Formation in north-east Texas. All x 1.

254 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 188. A-E. Sciponoceras kossmati (Nowak, 1908). A-D. The original of

Matsumoto & Obata (1963, pl. 6 (fig. 4)), Kyushu University Collections,

GK H4337, ‘approximately basal Turonian’ Mikasa Formation, Ikushumbets Valley,

Ishikari Province, Hokkaido, Japan. E. Dorsal view of the original of Matsumoto &

Obata (1963, pl. 5 (fig. 3)), Kyushu University Collections, GK H4342, from the

same horizon and locality as A-D. F-H. Sciponoceras baculoides (Mantell, 1822),

the original of Matsumoto & Obata (1963, pl. 1 (fig.:3)), Kyushu University

Collections H5453 from the Cenomanian of the D-sawa, a tributary of the

Ikushumbets Valley, Ishikari Province, Hokkaido, Japan.

All x 1.

THE AMMONITE FAMILY BACULITIDAE 255

D E F

Fig. 189. A-C. Sciponoceras ?baculoides (Mantell, 1822). CAS-61818.01, the

holotype of Cyrtochilus major Anderson, 1958, from Shasta County, Cottonwood

District, Hayes Gulch, 2 miles north-west of Gas Point, near-basal beds of Upper

Cretaceous Series. D-F. Sciponoceras ?kossmati (Nowak, 1908). CAS-31097.01, the

holotype of Cyrtochilus stylus Anderson, 1958, from Tehama County, on Dry Creek

(Redbluff-Beegum road), 4.6 miles W of Rosewood, a few hundred feet above the

local base of Upper Cretaceous, 0.1 miles east of basal Chico conglomerate.

Both x 1.

Genus Sciponoceras Hyatt, 1894

* Sciponoceras baculoides (Mantell, 1822) Figs 178-180

Mantell 1822: 123, pl. 23 (figs 6-7) (as Hamites baculoides). ?Collignon

1933: 72, pl. 5 (fig. 7) (as Baculites baculoides). Matsumoto 1959: 104,

pl. 31 (fig. la-d), text-fig. 2a—b (cum. synon.). Matsumoto & Obata 1963:

9, pl. 1 (figs 1-4), pl. 2 (figs 1-3), text-figs 3, 26-32. non Henderson 1973:

81, figs 4a-d, 6, nos 4a-c, 5a-c, 7a—c. Forster 1975: 166, pl. 4 (fig. 6),

text-fig. 36. non Wiedmann & Schneider 1979: 656, pl. 4 (figs 2-4), text-

fig. 5. Marcinowski 1980: 252, pl. 3 (figs 17-20). Chiriac 1981: 64, pl. 4

(figs 1-5), text-fig. 21. Szdsz 1982, 35, pl. 2 (figs 2a—b, 3a—-b, 4a-b), pl. 3

(fig. 2a—b). Kennedy & Juignet 1983: 19 (figs 11(a)-(y), 12(a)-(bb),

13(a)-(w), 14(a)-(n) (cum. synon.). Marcinowski & Walaszczyk 1985, pl. 1

(fig. 3). Kaplan et al. 1983, pl. 5 (fig. 2). Wright & Kennedy 1987: 177,

256

ANNALS OF THE SOUTH AFRICAN MUSEUM

pl. 37 (fig. 13). Blaszkiewicz & Szymakowska 1989: 266, pl. 519 (fig. 2).

Delamette & Kennedy 1991: 462, figs 17.6, 17.7, 17.14, 17.15. Young

(undated) 1979: 23, pl. 1 (figs 41-45), text-fig. 9j (as Sciponoceras sp. cf.

baculoides). Summesberger 1992: 123, pl. 8 (figs 7-9). Thomel 1992, pl. 6

(fig. 7), pl. 10 (fig. 2), pl. 11 (figs 1-3), pl. 19 (fig. 4). Kennedy 1994: 234,

pl. 10 (figs 7-9), pl. 12 (figs 1-6). Wright & Kennedy 1995: 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. Klinger & Kennedy 1997: 11,

fig. 2d-f.

Type. Lectotype by the 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)), BMNH 8612.

Occurrence. This species has been recorded from the Lower to the Upper

Cenomanian, but many of these records cannot be substantiated. According

to Wright & Kennedy (1995: 319), S. baculoides occurs ‘in flood abundance

at the top of the Middle Cenomanian T. costatus Subzone of the

Cc D

Fig. 190. Sciponoceras roto Cieslinski, 1959. A-B. BGS-GSM70774 from the

phosphatic Lower Cenomanian Mantelliceras mantelli zone fauna near Warminster,

Wiltshire. C-D. BMNH C88900 from the same horizon and locality.

E. BGS-GSM70778 from the same horizon and locality. F. BMNH C24423 from the

same horizon and locality. G. Muséum d’Histoire naturelle de Genéve Collections

no. 1316, from the Cenomanian of the Rochers de Fiz, Haute Savoie, France.

All x 1.

THE AMMONITE FAMILY BACULITIDAE 257

Fig. 191. Trachybaculites columna (Morton, 1834). A-C. ANSP 72867a.

D-E. ANSP 72867b. F-H. ANSP 72867e. J, N-O. ANSP 72867c.

K-M. ANSP 72867d. All syntypes and from the Maastrichtian Prairie Bluff Chalk at

Prairie Bluff, Alabama. All x 2.

258 ANNALS OF THE SOUTH AFRICAN MUSEUM

A. rhotomagense Zone in the Lower Chalk from Kent to the Isle of Wight,

and in the phosphatized faunas of the Chalk Basement bed of comparable

age at Punfield Cove and Buckland Newton, Dorset. It ranges to the lower

Upper Cenomanian C. guerangeri Zone, occurring in the phosphatized

faunas of the Chalk Basement Bed of Dorset and Somerset, and C Bed of the

Cenomanian Limestone in Devon.’ It is also known from West and Central

Europe, North Africa, southern India, California, ?Texas, Hokkaido,

Mozambique, Zululand and Madagascar.

* Sciponoceras bohemicum (Fritsch, 1872) Figs 181-183

Fritsch 1872: 49, pl. 13 (figs 23-25, 29, 30); Fritsch 1875: 78, fig. 63 (as

Baculites faujasii var. bohemica). Wright 1979: 285, pl. 1 (figs 3-5), pl. 7

(figs 10, 12) (cum. synon.). Wright & Kennedy 1981: 115. Wright &

Kennedy 1981: 115, pl. 31 (figs 4-6, 8, 10-11, ?7), pl. 32 (figs 9-10,

12-15) (as S. bohemicum anterius). Wright & Kennedy 1981: 116, pl. 31

(fig. 9) (as S. bohemicum bohemicum). Szasz 1982: 35, pl. 2 (figs 2a—b,

3a-b, 4a-b), pl. 3 (fig. 2a-b). Wright & Kennedy 1987: 177, pl. 37

(figs 7-8). Summesberger 1992: 123, pl. 8 (figs 7-9). Metzdorf 1992: 293,

pl. 3 (figs 5, 7). Santamaria Zabala 1992: 235 (as S. bohemicum anterius);

1992: 236, pl. 2 (fig. 7). Summesberger & Kennedy 1996: 134, pl. 18

(figs 4, 26) (as S. bohemicum).

Type. Lectotype by the subsequent designation of Wright (1979: 285) is the

original of Fritsch (1872, pl. 13 (fig. 25a-c)) from the Turonian Priesen

Beds of Lenesic, near Laun, Bohemia.

Occurrence. Turonian, northern and central Europe, Spain.

Sciponoceras cucullatum Collignon, 1964

Collignon 1964: 38, pl. 326 (fig. 1458). Klinger & Kennedy 1997: 12,

figs 2g-l, n-o, 4-5, 6a-d.

Type. Holotype by monotypy is the original of Collignon (1964, pl. 326

(fig. 1458)) from the Lower Cenomanian, west of the falls of Mahaboka,

Manera, Madagascar, GD 11458.

Occurrence. Lower Cenomanian of Madagascar; Cenomanian, Zululand.

* Sciponoceras glaessneri Wright, 1963 Fig. 184

Wright 1963: 599, pl. 81 (figs 2-3).

Type. Holotype is the original of Wright (1963, pl. 81 (fig. 2)) from the

Middle Cenomanian of Meadinga, Bathhurst Island, GDUA F.15852.

Occurrence. Middle Cenomanian, Bathhurst Island, Australia.

Fig. 192 (see facing page). Nodose Baculites sp. from the Upper Campanian of Oron,

Israel. A-C. SAM-PCI8582. D-F. SAM-PCI8583. G-I. SAM-PCI8585.

J-L. SAM-PCI8586. M-O. SAM-PCI8581.

All x 1.

THE AMMONITE FAMILY BACULITIDAE 259

260 ANNALS OF THE SOUTH AFRICAN MUSEUM

Sciponoceras gracile (Shumard, 1860) Figs 185-186

Shumard 1860: 596 (as B. gracilis). non Boule et al. 1907: 44. ?non

Collignon 1931: 94 (54), pl. 9 (5) (fig. 28). Cobban & Scott 1972: 47, pl. 17

(figs 9-29), text-fig. 18 (cum. synon.). Kauffman & Powell 1977: 100,

pl. 10 (fig. 4). ?Cooper 1978: 70, fig. 12b-d. Wright & Kennedy 1981: 112,

pl. 31 (figs 1-3), pl. 32 (figs 8, 11), text-figs 38a—q (cum. synon.). Kennedy

et al. 1981: 30, pl. 8 (figs 9-10), pl. 10 (figs 1-4). Kennedy & Juignet 1983:

22, figs 18(a)-(d), 32(i)-(p) (cum. synon.). Wright & Kennedy 1987: 177,

pl. 37 (figs 3-4). Kennedy 1988: 108, pl. 20 (figs 1-14, 17-20), text-fig. 38.

?Konecny & Vasi¢ek 1987: 91, pl. 1 (fig. 3), pl. 5 (fig. 2) (as Baculites cf.

gracile). Cobban et al. 1989: 61, figs 94a-b, 95g-n, 96a-b.

Type. Shumard’s types are lost (fide Cobban & Scott 1972: 47) and, as yet,

no neotype has been designated.

Occurrence. Upper Cenomanian of southern England, France, southern

Germany, Czechoslovakia, the United States, Mexico, and perhaps Angola.

Records from Madagascar (Boule et al. 1907; Collignon 1931) are dubious,

as is that from Cameroon (Solger 1904; see Reyment 1955: 15).

Sciponoceras intermedium Matsumoto & Obata, 1963

Matsumoto 1959: 109, pl. 30 (figs 2a-c, 3a-b), pl. 31 (?fig. 4), text-

figs 7a—b, 8-11 (as S. aff. bohemicum). Matsumoto & Obata 1963: 24, pl. 8

(figs 1-3, 6), pl. 11 (fig. 6), pl. 12 (figs 4-5), text-figs 50-59, 61.

Type. Holotype is the original of Matsumoto & Obata (1963, pl. 8

(fig. la—d)), Upper Turonian, from the basal part of the Upper Yezo Group,

along the Pombets, a tributary of the [kushumbets River, Hokkaido,

GK H5386.

Occurrence. Upper Turonian, Hokkaido.

* Sciponoceras kossmati (Nowak, 1908) Figs 187, 188A-E

(= ?S. gracile (pars) fide Wright & Kennedy 1981: 114)

Kossmat 1895: 154, pl. 19 (5) (fig. 18a-d) (as Baculites n. sp. aff.

bohemicum). Nowak 1908: 348 (as Baculites kossmati). Anderson 1958:

188, pl. 11 (fig. 5, 5a) (as Cyrtochilus stylus). Matsumoto 1959: 106, pl. 31

(figs 2a—b, 3), text-figs 4a-—b, 5a-b, 6a—b. 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.

Immel 1979: 631, pl. 3 (fig. 4) (as S. cf. kKossmati). Immel 1987: 128 (as S.

cf. kossmati).

Type. Holotype is the original of Kossmat (1895, pl. 19 (5) (fig. 18a-d))

from the Lower Trichonopoly Group of India, GSI.

Fig. 193 (see facing page). Nodose Baculites sp. from the Upper Campanian and Lower

Maastrichtian of Israel. A-C. SAM-PCI8576. N-P. SAM-PCI8577.

Q-S. SAM-PCI8578. All from the Lower Maastrichtian of the upper part of the Mishash

Formation, Phosphate Member, Oron. D-F. SAM-PCI8587. K-M. SAM-PCI8584.

D-M. All from the Upper Campanian, Oron. All x 1.

261

THE AMMONITE FAMILY BACULITIDAE

262 ANNALS OF THE SOUTH AFRICAN MUSEUM

THE AMMONITE FAMILY BACULITIDAE | 263

Occurrence. Lower Turonian, California, rarely in the Middle Cenomanian;

common in uppermost Cenomanian and basal Turonian of Hokkaido; the

Indian holotype was recorded from the Lower Trichonopoly Group

of Garudamungalum (Lumachelles), which dates it as late Turonian;

Germany.

* Sciponoceras major (Anderson, 1958) Fig. 189A-—C

(= S. baculoide fide Matsumoto 1959: 104)

Anderson 1958: 189, pl. 38 (fig. 6) (as Cyrtochilus major).

Type. Holotype by monotypy is the original of Anderson (1958, pl. 38

(fig. 6)) from Hayes Gulch, Cottonwood District, Shasta County, California.

CAS type collection.

Occurrence. Cenomanian, California.

Sciponoceras matsumotoi Inoma, 1980

Inoma 1980: 171, pl. 2 (figs 4-6), text-fig. 4.

Type.

Occurrence. Cenomanian, Hokkaido.

Sciponoceras orientale Matsumoto & Obata, 1963

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.

Type. Holotype is the original of Matsumoto & Obata (1963, pl. 6

(figs 1-2)) from the Turonian, upper part of the Middle Yezo Group,

Abeshinai-Saku area, Teshio Province, Hokkaido, Geological Institute,

University Tokyo, GT I-3160a.

Occurrence. Approximately Lower and Middle Turonian of Hokkaido.

Sciponoceras pervinquierei Breistroffer, 1940

Pervinquiére 1907: 92, pl. 4 (fig. 8) only (as B. baculoides). Breistroffter

1940: 29 (as Cyrtochilus Pervinquierei = invalid under article 13(a) of

ICZN; = S. roto).

Type. Holotype by the original designation of Breistroffer (1940: 29, p. 36

footnote) is the original of Pervinquiére (1907, pl. 4 (fig. 8)) from the

Vraconian of Mechtat Mergueb, Tunisia.

Occurrence. Lower Cenomanian, Tunisia.

Fig. 194 (see facing page). A-C. Trachybaculites? furcillatus (Blanckenhorn, 1905).

SAM-PCI8573 from the Lower Maastrichtian of Israel. D-R. Baculites ?anceps.

D. SAM-PCI8560. E, I-K. SAM-PCI8561. F-H. SAM- PCI8551. O. SAM-PCI8550.

P-R. SAM-PCI8549, all from the Lower Maastrichtian, upper part of Mishash Formation,

Phosphatic Member, Nahal Ashosh. S-X. Smooth Baculites from the upper Lower

Campanian of Nahal Qidron, Judean Desert, Israel. All x 1.

264 ANNALS OF THE SOUTH AFRICAN MUSEUM

A B Cc

H ! J

Fig. 195. Smooth Baculites. A-C. SAM-PCI8571 from the upper Lower Campanian

of Nahal Qidron, Judean Desert. D-F. SAM-PCI8569. G-J. SAM-PCI8567, both

from the lower Upper Campanian of the Arad Region, east-central Israel. All x 1.

* Sciponoceras roto Cieslinski 1959 Figs 170A-B, E-L, 190

Cieslinski 1959: 89, pl. 4 (fig. 10a—c), text-fig. 14. Kennedy 1971: 10, pl. 3

(fig. 7). Hancock et al. 1972, pl. 81 (fig. 8a-c). Wiedmann & Schneider

1979: 657. Immel 1979: 632, pl. 4 (fig. 5) (as S. cf. roto). Kennedy et al.

1979: 10 (as S. cf. roto). Marcinowski 1980: 254, pl. 3 (figs 14-15).

Wiedmann & Boess 1984: 501, fig. 6ls—t. Immel & Seyed-Emami 1985: 112

(as Baculites cf. roto). Immel 1987: 128 (as S. cf. roto). Blaszkiewicz &

Szymakowska 1989: 266, pl. 566 (fig. 2). Delamette & Kennedy 1991: 460,

figs 17.8-17.13, 17.16-17.23. Wright & Kennedy 1995: 315, pl. 94

(figs 13-19), pl. 95 (fig. 4), pl. 98 (fig. 28), text-figs 131j-1, n. Klinger &

Kennedy 1997: 17, fig. 2p.

Fig. 196 (see facing page). Baculites vaalsensis Kennedy & Jagt, 1995. A-C. Paratype,

IRSNB 10462b, from the Hervian of Aachen. D. IRSNBIG5425, from the Hervian of

Battice-Croix Polinard. E-F. IRSNB 10460, from the Vaals Formation of Aachen.

G. IRSNBIG 5425, locality as for 4. H-J. Collections of the Rheinisch-Westfalische

Technische Hochschule, Aachen, from the Vaals Formation of Aachen. K-M. IRSNBIG,

locality as for 4. N. Paratype IRSNB 10462c, locality as for 1-3. O-Q. IRSNBIG5425,

locality as for 4. All x 1.

265

THE AMMONITE FAMILY BACULITIDAE

. 196

ANNALS OF THE SOUTH AFRICAN MUSEUM

266

Fig. 197

THE AMMONITE FAMILY BACULITIDAE 267

Type. Cieslinski (1959) based this species on nine syntypes; no lectotype has

as yet been designated. The figured specimens are housed in the collections

of the Geological and Palaeontological Institute, University of Warsaw.

Occurrence. Lower Cenomanian of southern England, Germany, Spain,

Zululand and Mangyschlak; according to Marcinowski (1980: 254) it ranges

through the whole Cenomanian of Poland.

Sciponoceras santacrucense Leanza, 1970

Leanza 1970: 212, fig. 11 (1-7). Riccardi & Aguirre Urreta 1988, pl. 2

(figs 3-4). Klinger & Kennedy 1997, fig. 6e.

Type. Holotype is the original of Leanza (1970, fig. 11 (1-3)) from the

upper reaches of the section at Puesta Los Alamos, Santa Cruz Province,

Argentina, CPC 4326.

Occurrence. Turonian, Argentina.

Sciponoceras subbaculoides (Geinitz, 1875)

Geinitz 1875: 281, pl. 63 (fig. 1). Horvath 1983, pl. 3 (figs 15-16).

Type. Lectotype by the subsequent designation of Wright & Kennedy (1981:

114) is the original of Geinitz (1875, pl. 63 (fig. 1)) from the lower Planer at

Oberau, Bohemia.

Occurrence. ?Upper Cenomanian, Czech Republic (Bohemia) and Hungary.

* Sciponoceras stylus (Anderson, 1958) Fig. 189D-F

(= S. kossmati fide Matsumoto 1959: 106; S. gracile fide Cooper 1978: 70)

Anderson 1958: 188, pl. 11 (fig. 5, 5a) (as Cyrtochilus stylus).

Type. Holotype is the original of Anderson (1958, pl. 11 (fig. 5, 5a)) from

the Lower Cenomanian of Dry Creek, west of Rosewood, northern Tehama

County, California, CAS type collection.

Occurrence. Lower Cenomanian, California.

Genus Trachybaculites Cobban & Kennedy, 1995

* Trachybaculites columna (Morton, 1834) Fig. 191

Morton 1834: 44, pl. 19 (fig. 8). Johnson 1905: 27. Stephenson 1941: 405,

pl. 76 (figs 1-4). Matsumoto 1959: 161, pl. 30 (fig. 1), pl. 34 (fig. 4), pl. 43

(fig. 4), text-figs 80a—b, 81. Cobban & Kennedy 1992c: 682, figs 1.7-1.9,

3.2. Cobban & Kennedy 1995: 29, figs 10.1, 10.3, 13.4-13.6, 14.3, 14.9,

17.1-17.4, 17.17-17.31. Kennedy & Jagt, 1998: 161, pl. 2 (figs 1-6).

Fig. 197 (see facing page). Baculites vaalsensis Kennedy & Jagt, 1995. A-C. Collections

of the Rheinisch-Westfalische Technische Hochschule, Aachen, from the Vaals Formation of

Aachen. D, E-G. IRSNBIG 5425, from the Hervian of Battice-Croix Polinard.

H. IRSNB 10459a from the Vaals Formation of Vaals, the Netherlands. I. IRSNBIG5425,

locality as for D. All x 1.

268 ANNALS OF THE SOUTH AFRICAN MUSEUM

Type. Lectotype, by subsequent designation of Cobban & Kennedy (1995:

29) is ANSP 72867a from the Prairie Bluff Chalk, Prairie Bluff, Alabama.

Occurrence. Maastrichtian, Prairie Bluff Chalk of Alabama and Mississippi,

Fox Hills Formation, California, and the Netherlands.

Trachybaculites vicentei Stinnesbeck, 1986

Stinnesbeck 1986: 203, pl. 9 (fig. 4), pl. 10 (figs 3-6), text-fig. 23a-c.

Type. Holotype is the original of Stinnesbeck (1986, pl. 10 (fig. 3)) from the

Maastrichtian of west of La Gloria, San Vicente near Talcahuano, Central

Chile, GPIB V3/240.

Occurrence. Maastrichtian, Central Chile.

ACKNOWLEDGEMENTS

Part of this discussion was presented at the Ammonoid Symposium in

London in 1991. Financial assistance to Klinger by the Foundation for Research

Development, South Africa, is gratefully acknowledged. Samantha Black,

Jacque Blaeske, Ingrid Klinger and Madel Joubert provided technical assistance

with the illustrations for the symposium presentation and for this printed

version. Kennedy acknowledges the financial support of the Natural

Environment Research Council (United Kingdom) and the technical! assistance

of the staff of the Geological Collections, Oxford University Museum of Natural

History, and the Department of Earth Sciences, Oxford.

Amongst the colleagues who supplied us with material, loan of specimens

and valuable advice, we have to single out Dr W. A. Cobban, U.S. Geological

Survey, Denver, Neal L. and Peter L. Larson of the Black Hills Institute of

Geological Research, Hill City, and Dr N. H. Landman, American Museum of

Natural History, New York. Without their assistance, we would never have

been able to study the Baculitidae in such detail. Other colleagues to whom we

are indebted in various ways are Dra M. B. Aguirre Urretta (Buenos Aires), Mr

H. Breitkreutz (Bielefeld), Prof. M. R. Cooper (Durban-Westville), Dr N.

Eldridge (New York), Dr J. Gorzyca-Skala (Wrociaw), Dr J. Haggart

(Vancouver). Dr M. K. Howarth (London), Prof. H. Immel and Dr G. Schairer

(Munich), Dr J. Jagt (Maastricht), Dr Z. Lewy (Jerusalem), Dr K. Larsson

(Lund), Dr U. Leppig (Freiburg), Prof. E. Mancini and Dr E. ‘Puckett,

Tuscaloosa, Prof. R. Marcinowski, Drs I. Walaszczyk and M. Machalski

(Warsaw), Prof. Emeritus T. Matsumoto (Fukuoka), Dr R. Ragaini (Pisa),

Prof. H. Remy (Bonn), Dr S. Ritzkowski (G6ttingen), Dr A. C. Riccardi (La

Plata), Dr H. Summesberger (Vienna), Dr C. Sarti (Bologna), Dr F. Thackeray

(Pretoria), Prof. J. Thierry (Dijon), Dr G. Thomel (Nice), Prof. Z. Vasitek

(Ostrava), Prof. P. Ward (Seattle), Prof. G. E. G. Westermann (Hamilton) and

Prof. K. Young (Austin). If we have left out anyone we apologize but are

neverthless grateful for their assistance.

We thank the Natal Parks Board, especially Mr Ricky Taylor for access to

areas under their control and for assistance with transportation around the Lake

St Lucia and False Bay areas during the field season of 1992.

THE AMMONITE FAMILY BACULITIDAE 269

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

(particularly Articles 22 and 51).

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

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

etc. 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): Nickles, p. 163, fig. 301.

1955 Leda bicuspidata (Gould): Nickles, 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 Tabies 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 sumames 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.