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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 und anthropolegische Ergebnisse einer Forschungreise im westlichen und zentralen Sild Afrika ausgefihrt in den Jahren 1903-1905 4 (15). Denkschriften der medizinisch-naturwissenschaftlichen Gesellschaft zu Jena 16: 269-270. 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) By HERBERT CHRISTIAN KLINGER & WILLIAM JAMES KENNEDY The South African Museum forms part of Iziko Museums of Cape Town The Annals of the South African Museum publishes original research articles, revisions and review articles in anthropology, archaeology, palaeontology, geology, entomology, herpetology, ornithology, and marine and freshwater biology. In order to be considered for publication, manuscripts should deal, at least in part, with material from the collections of the South African Museum. Other contributions are also considered provided at least one of the authors is a staff member of the Museum. In the case of descriptions of new species not already part of the Museum's collections, the holotype and, if possible, part of the paratype series must be deposited in the South African Museum. Authors whose contributions do not meet with these criteria should contact the Editorial Committee prior to final preparation and submission of their manuscript. All articles are refereed by three referees of international standing. Each paper accepted is published as a separate part at irregular intervals as material becomes available; parts are priced individually. Editorial committee Dr P. A. Hulley (Chairman) Dr H. C. Klinger Elizabeth Louw (Editor) ANNALS OF THE SOUTH AFRICAN MUSEUM Publisher: South African Museum, P.O. Box 61, Cape Town 8000 South Africa Please direct enquiries (including copyright) and manuscripts to the Editor. Please direct orders and subscriptions to the Publications Office. e-mail: elouw@samuseum.ac.za BACK COPIES Orders for back copies of the journal are welcome. For a full list of papers published since 1898, please write to the above address or visit our web-site: http://nv1.samuseum.ac.za/collect/infosci/pub/prclist.htm OUT OF PRINT 1, 2(1-3, 5-8), 3(1-2, 4-5, 8, t.-p.i.), 4(1), 5(1-3, 5, 7-9), 6(1, 2, t.-p.i.), 7(1-4), 8, 9(1-2, 7), 10(1-3), 11(1-2, 5, 7, t.-p.i.), 14(1-3), 15(4-5), 24(2, 5), 27, 31(1-3), 32(5), 33, 36(2), 43(1), 45(1), 49(1), 67(5, 11), 84(2) ISBN 0 86813 179 2 D8540 Printed in South Africa by The Rustica Press, Old Mill Road, Ndabeni, Western Cape 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) By 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 © onlin Sa eee ene ce ee aeteree a erercle's wise ees caine ds alae sian cnet see ee aio saeemobaanctares 31 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.) Lechites Criobaculites fae} a os | = 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 “UBNYOUsee] oy) SuLINp ‘9Z76] ‘yeds Sainooqny snues oy} JO uoNGLsIp o1yde1Z0eH ad V snoqyiussqny “J =q'] SsnjouLoraw] “yz =] xajduis ‘qJ=5 ousva “J = A Snayjuukgn] “J =Q'y Snjoursmsayy] *J =17] Xajduns -y = § puispa "J = A sSnpuups-qJ=.) . 7 31 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 12 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 = Z =) eo) an) > @ a | -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 = eo pa Se Z = 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 = ~ = rn 7 ) 4 ss re Z Z “3 rn r ) rs 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 | 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 | 17 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 WL 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 0 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 C B. lomaensis ae E. occidentalis | a B. rectus me Motes) Of B. regina ee 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 JI E. carinatus Ord») Adowond B. baculus OLD) T. columna Olds) 27 Fig. 16. Stratigraphic distribution of species of the genera Baculites, Trachybaculites Eubaculites and Fresvillia in the Maastrichtian in Europe. 28 ANNALS OF THE SOUTH AFRICAN MUSEUM MAASTRICHT. iB. ambatryensis E L U | B. subanceps Tee -- B. vicentei E 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} 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. 29 5, 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 = c : >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’). iL 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) I 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). I A E L i U B 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 B E I Cc = Cae E I D ie U Edin a Unset E E 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 L E U I A | 0 10 E L U I B 0 1 E L U C E L GT UY. L D Bd ad Mls 2 E L U I | E 0 1 el 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 ee B 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 A E B L 0 me L E U I C j 0 1 E | ; ee D 0 1 E L I U E 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 E L Pee A ; E L eS nel B : ik OG C E it D : U sm erg! E &®&% io U png! E 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. E L i U C 5 im) E L I U D i 0 5 E it jk 7 U E 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). A E L U L | iis Sea E E L OI Uae ne E B E 1 Ry’ ane C 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 A 5 E iG U B E ; I Cc Eas 0 5 E naa L ae U D 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 | B L U (¢ L U D 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 o) 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 E \ L a B 0 10 E 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. 35 THE AMMONITE FAMILY BACULITIDAE "I x “BIpU] WoYINOS JO UOHeUWIO,| INAepNeA ey) WoIy WOE “GYIISOHNING jesaye] Oy] qld YIM suouIoeds JUSUILUIO [eIO}L] UT UOTIETIeA *(QPRT oH ‘saqio d ‘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 es No @\ \ \ i O Lechites \/_ Sciponoceras @ ae ©. Baculites ©} Eubaculites ] oo @ 70 @ MAXIMUM RECORDED WHORL HEIGHT 50 @® 0@ lo ef a) & @) 4Q @ } ® @ 2 @ @ ae OL” © 30 @ @ @ o 8 ee 20 @ @® © =e Oe @ @ ® @ ©@@ 1Q 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. nN (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. 2 | k 2 Size distribution in Baculites capensis | | | 5. @ indicates distribution of specimens with preserved apertures. NUMBER OF SPECIMENS —_ ee 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 73 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 F 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. TE 78 yyy yf, YG, Ui; Z Y yy 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. 81 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. 83 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. 85 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. 91 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. le 93 THE AMMONITE FAMILY BACULITIDAE . 70 1g F 94 ANNALS OF THE SOUTH AFRICAN MUSEUM A 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. 2 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 96 oh HO basis be ‘I Ay P, 135 toe 73 18 F 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. 98 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 Os ecg etnies < SD om x oO ~ © oO Om a. TAH Ka ie A} ase. ° na RAS =o HG av Ae e223 3 Os Q QAVAN © Sage i) x Vevs 5S Oy @z= © OSS .n bo} 0 SE gazes aeeov 6A oS 1 ie) -~ 82 Sf oH se Sige) 6y (ey S556 as ¥ oS E-o S88 SO) Sao Qa 8 = 6 wv’eE cO MD op on. on) oO # aq fy Ay A, 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 98 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 TX TV “eles uojsurysem ‘purysy Blong ‘spurjsy ueny ueg ‘eIZ10eH Jo Weng Woy ‘70'8ZZ-SVD *Z981 “YSOIAI SNIDULOUI SaNNIDg *J-H “eIUIOsTTED ‘ asnoy IIW OT, By) 1 peoy Ipjoquinyy wo yseID oo1YD 1g 0} peor ANUNOD suoye SoTIW 9° “Jeet OOTYD JO Yue ysea “AyunoD ayng WO ‘70°L6619-SVO “9S8I ‘ASPIL SisuaooIyo sarynovg *J-q *BunoTuCo 1B SVD pur (DET :6S61) ClouNsyeW Aq UOATZ eJEp AjTBOO] “10°1899-SWO ‘*8S6I ‘UOsIopuy xau SalNoDg “Q-V “ROI “3Iq 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. a : reat y i io ‘ eet e L = . i : ' > t en: = ; \. r. ; ba i} ' ey i : in ; vs 2) Qe ‘ ie = 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 (c) Scientific names, but not their vernacular derivatives e.g. Therocephalia, but therocephalian Punctuation should be loose, omitting all not strictly necessary. Reference to the author should 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. WOON YAN HERBERT CHRISTIAN KLINGER & WILLIAM JAMES KENNEDY 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)