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Caudal Fin Skeleton of the Late Cretaceous Lamniform Shark, Cretoxyrhina Mantelli, from the Niobrara Chalk of Kansas

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Caudal Fin Skeleton of the Late Cretaceous Lamniform Shark, Cretoxyrhina Mantelli, from the Niobrara Chalk of Kansas Lucas, S. G. and Sullivan, R.M., eds., 2006, Late Cretaceous vertebrates from the Western Interior. New Mexico Museum of Natural History and Science Bulletin 35. 185 CAUDAL FIN SKELETON OF THE LATE CRETACEOUS LAMNIFORM SHARK, CRETOXYRHINA MANTELLI, FROM THE NIOBRARA CHALK OF KANSAS KENSHU SHIMADA1, STEPHEN L. CUMBAA2, AND DEANNE VAN ROOYEN3 1Environmental Science Program and Department of Biological Sciences, DePaul University, 2325 North Clifton Avenue, Chicago, Illinois 60614; and Sternberg Museum of Natural History, Fort Hays State University, 3000 Sternberg Drive, Hays, Kansas 67601; 2Paleobiology, Canadian Museum of Nature, P.O. Box 3443, Station D, Ottawa, Ontario K1P 6P4, Canada; 3Department of Earth Sciences, Carleton University, 2240 Herzberg Laboratories, 1125 Colonel By Drive, Ottawa, Ontario K1S 5B6, Canada. Abstract—The caudal fin morphology of the Late Cretaceous lamniform shark, Cretoxyrhina mantelli (Agassiz), was previously inferred from scale morphology, which suggested that it was capable of fast swimming. A specimen from the Niobrara Chalk of western Kansas is described here and offers new insights into the morphology of the caudal fin of the taxon. The specimen preserves the posterior half of the vertebral column and a series of hypochordal rays. These skeletal elements exhibit features suggesting that C. mantelli had a lunate tail and a caudal peduncle with a lateral fluke. The specimen also supports the idea that the body form of C. mantelli resembled that of the extant white shark, Carcharodon carcharias (Linneaus). Given a total vertebral count in Cretoxyrhina mantelli of about 230, this specimen suggests that the transition between precaudal and caudal vertebrae was somewhere between the 140th and 160th vertebrae. The estimated total body length of the specimen described here ranges from 640 cm to 700 cm, marking the largest C. mantelli individual estimated to date. New skeletal data from the specimen further supports the view that C. mantelli was an active shark capable of fast swimming. INTRODUCTION insights into the shape of the shark’s tail. The purpose of this paper is 1) to describe the morphology of the specimen, and 2) to discuss the caudal fin Cretoxyrhina mantelli (Agassiz) was a Late Cretaceous lamniform morphology of C. mantelli and its paleoecological significance. For com- shark that lived in Cenomanian–Campanian seas worldwide, including the parative purposes, fossil specimens in the following institutions are referred Western Interior Seaway of North America (e.g., Cappetta, 1987; Siverson, to in this paper: Sternberg Museum of Natural History, Fort Hays State 1992, 1996; Shimada, 1997d). The species is represented chiefly by its University (FHSM), Hays, Kansas; and the vertebrate paleontology collec- teeth, but some reasonably complete skeletons of the species are known tion of the University of Kansas Museum of Natural History (KUVP), from the late Coniacian– Santonian portion of the Smoky Hill Chalk Mem- Lawrence. ber of the Niobrara Chalk in western Kansas (Shimada, 1997b). Those skeletal remains suggest that large individuals of C. mantelli measured about SYSTEMATIC PALEONTOLOGY 5 to 6 m in total length and possibly had a body form similar to the modern great shark, Carcharodon carcharias (Linnaeus) (Shimada, 1997b). The Class Chondrichthyes fossil record demonstrates that Cretoxyrhina mantelli fed on large marine Subclass Elasmobranchii vertebrates (e.g., teleosts, sea turtles, mosasaurs, and plesiosaurs: Shimada, Order Lamniformes Berg, 1958 1997c; Shimada and Everhart, 2004; Shimada and Hooks, 2004; Everhart, Family Cretoxyrhinidae Glikman, 1958 2004, 2005a) and possibly scavenged “bloat-and-float” carcasses of fully terrestrial vertebrates (e.g., nodosaurid dinosaurs: Everhart and Hamm, Genus Cretoxyrhina Glikman, 1958 2005). Cretoxyrhina mantelli (Agassiz, 1843) Despite the wealth of information about the paleobiology of Cretoxyrhina mantelli (e.g., Shimada, 1997a, 1997b, 1997c), there are Material—CMN 40906 (Figs. 1–5), a string of caudal and poste- rior precaudal vertebrae with hypochordal rays and placoid scales. still many unresolved questions. The morphology of its caudal fin is one such gap in our knowledge. The previously suggested morphology, and the Horizon and locality—The specimen was collected by G. F. Sternberg from the Niobrara Chalk (Upper Cretaceous: see Hattin, 1982) estimated starting point of the caudal fin in C. mantelli (Shimada, 1997b), were based on a series of assumptions. Shimada (1997b) demonstrated in western Kansas. It was sold to the Geological Survey of Canada, from which the museum originated, in 1912. that C. mantelli had keeled placoid scales with an average interkeel dis- tance of approximately 45 microns. Because this interkeel distance is com- Because exact biostratigraphic data were not available to constrain the age of the specimen within the Niobrara Chalk, chalk samples were parable to that in scales of some extant fast-swimming sharks, Shimada (1997b) inferred that C. mantelli was also capable of fast swimming. This taken from the matrix for stratigraphically diagnostic foraminiferans. Our result shows that four taxa dominate the assemblage. Heterohelix globulosa evidence, combined with the fact that C. mantelli had a conical head, led Shimada (1997b) to consider that the species had a stout fusiform body (Ehrenberg), with a Campanian – Maastrichtian range, is the most abun- dant. The other key taxa are: Archeoglobigerina cretacea (d’Orbigny), with a lunate caudal fin for efficient hydrodynamic propulsion. Given that the fossil species had a total vertebral count of approximately 230, Shimada with a range from middle Coniacian–early Maastrichtian; Rugoglobigerina rugosa (Plummer), known from the early Campanian; and Whiteinella (1997b) suggested that the caudal fin of C. mantelli could have begun at about 133rd vertebra, on the basis of comparisons with some extant fast- centennialensis Frerichs, which occurs in late Santonian–early Campanian strata (Pessagno, 1967; Frerichs et al., 1975; Frerichs, 1979). The pres- swimming lamniform sharks with lunate tail fins (e.g., lamnids: Lamna Cuvier, Isurus Rafinesque, and Carcharodon Smith). ence of R. rugosa, which makes up about 10% of the individuals in our sample, strongly suggests that the Cretoxyrhina specimen came to rest on The Canadian Museum of Nature (CMN) in Ottawa, Ontario, houses a putative Cretoxyrhina mantelli specimen, CMN 40906, which consists the chalky ocean bottom in the early Campanian. The shark specimen is thus from the upper part of the Smoky Hill Chalk. If so, whereas C. mantelli of a vertebral column and some additional skeletal elements (Fig. 1). CMN 40906 is noteworthy because of its large size and features that provide new is known from early Campanian deposits (e.g., Siverson, 1992), CMN 186 FIGURE 1. Photograph of CMN 40906, string of vertebrae with hypochordal rays of Cretoxyrhina mantelli (Agassiz) (anterior to the left; scale bar = 30 cm). FIGURE 2. Line drawing of CMN 40906 (cf. Fig. 1), string of vertebrae (“v”) with hypochordal rays (hcr) of Cretoxyrhina mantelli (Agassiz) (anterior to the left; scale bar = 30 cm). 40906 represents the youngest Cretoxyrhina specimen documented thus far from the Smoky Hill Chalk of Kansas (see Stewart, 1990; Everhart, 2005b, table 13.1). It should be noted that museum records indicate that the specimen was found in Gove County, Kansas, but strata from the lower Campanian (the uppermost Niobrara Chalk) are not known to occur there. Other fossil specimens purchased by the museum from the Sternberg family at the same time list G. F. Sternberg as the collector, and Logan County as the source. Because outcrops with lower Campanian strata do occur just west of Gove County in Logan County (Hattin, 1982), perhaps the collection data are incorrect. Description—A total of 109 vertebral centra are physically preserved in the specimen (Fig. 1). The anteriormost ninety-five of these are clearly in anatomical sequence and are in a good state of preservation. We refer to these as “v1” through “v95”, counting sequentially from the anteriormost centrum in the specimen (Fig. 2). The v-numbers are placed in quotations to highlight the fact that they are artificial assignments. There are impres- sions of four vertebrae near “v95” which are otherwise missing. Fourteen smaller vertebrae, generally in poor condition, are on the slab, but are largely FIGURE 3. Close-up view of vertebral centra (“v23”-“v25”: see Fig. 2) in CMN disarticulated, and have not been assigned v-numbers. If the four missing 40906 (scale bar = 2 cm). vertebrae and the 14 smaller vertebrae are counted, CMN 40906 consists 187 FIGURE 4. Close-up view of putative caudal fin base, showing connection between vertebral column and hypochordal rays in CMN 40906 (scale bar = 10 cm; cf. Figs. 1, 2). FIGURE 5. Placoid scales (four examples) of Cretoxyrhina mantelli (Agassiz) from CMN 40906 (scale bar = 50 micronsm). Sample locations are relative to vertebral positions in Figure 2. A, Scale from dorsal to “v24”: top, apical view (anterior to the top); bottom, anterior view. B, Scale from above, or ventral to “v66”: left, oblique view (anterior to the top left); right ,anterior view. C, Scale location same as A: lateral view (anterior to the left). D, Scale location same as B: apical view (anterior to the right). 188 FIGURE 6. Example of articulated vertebrae of Cretoxyrhina mantelli (Agassiz) (anterior to the left; scale bar = 10 cm; v 73–v82 in FHSM VP-2187: see Shimada, 1997b). A, vertebrae in right(?) lateral view (note that this surface was against ocean floor when the shark skeleton was buried); B, vertebrae in dorsal(?) view (note post- burial distortion that resulted in lateral flattening of vertebrae). of a total of 113 vertebrae. the tail. The centra (Fig. 3) are amphicoelous and asterospondylic with nu- A few patches of articulated placoid scales and many disarticulated merous concentric lamellae around the primary double-cone calcification scales were found in the chalk matrix surrounding the specimen (Fig. 5). (see Welton and Farish, 1993). The centra are well calcified, and the “double- The scales are of two types. The most common type (Figs.
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