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.

193 ANNOTATED CHECKLIST OF FOSSIL FISHES FROM THE SMOKY HILL CHALK OF THE NIOBRARA CHALK (UPPER CRETACEOUS) IN KANSAS

KENSHU SHIMADA1 AND CHRISTOPHER FIELITZ2

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;2Department of Biology, Emory & Henry College, P.O. Box 947, Emory, Virginia 24327

Abstract—The Smoky Hill Chalk Member of the Niobrara Chalk is an Upper Cretaceous marine deposit found in Kansas and adjacent states in North America. The rock, which was formed under the Western Interior Sea, has a long history of yielding spectacular fossil marine vertebrates, including fishes. Here, we present an annotated taxo- nomic list of fossil fishes (= non-tetrapod vertebrates) described from the Smoky Hill Chalk based on published records. Our study shows that there are a total of 643 referable paleoichthyological specimens from the Smoky Hill Chalk documented in literature of which 133 belong to chondrichthyans and 510 to osteichthyans. These 643 specimens support the occurrence of a minimum of 70 species, comprising at least 16 chondrichthyans and 54 osteichthyans. Of these 70 species, 44 are represented by type specimens from the Smoky Hill Chalk. However, it must be noted that the fossil record of Niobrara fishes shows evidence of preservation, collecting, and research biases, and that the paleofauna is a time-averaged assemblage over five million years of chalk deposition. Our historical review suggests that, whereas the number of described fish species from the Smoky Hill Chalk has de- clined in the last 75 years, the amount of research on fish taxa has constantly increased for the last three decades. We recognize three major periods of Smoky Hill Chalk paleoichthyological research: 1) between 1870 and 1879 due mostly to E. D. Cope’s effort in describing the majority of osteichthyan taxa; 2) between the late 1890s and into the early 1900s due primarily to efforts of new workers such as O. P. Hay, F. B. Loomis, A. Stewart, and S. W. Williston; and 3) since the mid-1960s. The recent resurgence in the study of fish taxa of the Smoky Hill Chalk appears to reflect the renewed interest in systematic ichthyology, biostratigraphy, and paleoecology.

INTRODUCTION (1990a) work is the lack of literature references and specific specimens that were used to base their occurrence data. The Niobrara Chalk is an Upper Cretaceous marine deposit occur- Because Russell’s (1988) taxonomic list included taxa based on ring in Kansas and adjacent states in North America. The rock formed unpublished data, and because Carpenter (1990, 2003) and Stewart’s under the Western Interior Sea, an epicontinental sea that extended in a (1990a) lists are not reproducible due to the lack of referred literature and generally north-south direction in the middle of the continent. The forma- specimens to support their occurrence data, uncertainty exists as to the tion consists of two calcareous stratigraphic members: the 20-m-thick Fort exact taxonomic composition of the vertebrate fauna in the Smoky Hill Hays Limestone Member gradationally overlaid by the 180-m-thick Smoky Chalk. Since Russell (1988), Stewart (1990a), and Carpenter’s (1990; 2003, Hill Chalk Member (Hattin et al., 1987; Hattin, 1982). The Fort Hays Lime- which is essentially based on his 1990 data) work, a number of papers stone yields vertebrate fossils, but only 12 taxa with small sample sizes are which describe vertebrates from the stratigraphic member were published known from the member to date (Shimada, 1996a; Shimada and Everhart, (e.g., Everhart et al., 2003; Hamm and Shimada, 2002b; Shimada, 1997f; 2003; Everhart, 2005b). On the other hand, the Smoky Hill Chalk is re- Stewart, 1996, 1999a). Also, some taxa represented in the Niobrara verte- garded as a fossil Lagerstätte and famous for yielding diverse and often brate fauna underwent taxonomic revisions in recent years. Therefore, we spectacular fossil marine vertebrates (Bottjer, 2002: Fig. 1). They include determined that a collective review of vertebrate fauna of the Smoky Hill chondrichthyans, osteichthyan fishes, and various tetrapods, such as ma- Chalk would make a significant contribution to the paleontology of the rine turtles, mosasaurs, plesiosaurs, pterosaurs, and birds (for overview, Niobrara Chalk (e.g., a review of Niobrara vertebrates in South Dakota by see Everhart, 2005c). In addition, there are some dinosaur remains from Martin et al., 1988). the Smoky Hill Chalk, which apparently represent post-mortem terrestrial In this paper, we focus on fossil fishes from the Smoky Hill Chalk materials washed out to sea (e.g., Carpenter et al. 1995; Everhart and Hamm, of Kansas (Figs. 1-4;Table 1). We here present an annotated taxonomic list 2005; Everhart and Ewell, in press). of fossil fishes described from the stratigraphic member base strictly on Fossil vertebrates from the Smoky Hill Chalk are documented in published records. We believe that our work illuminates the history of diverse scientific literature sporadically published since the mid-1800s (e.g., Niobrara fish research and serves as a useful research tool to those who Cope, 1868). The taxonomic composition of the vertebrate fauna, or part wish to study fish taxa from the formation. The list also offers a historical of the fauna, has been reviewed from time to time (e.g., Williston, 1900a; perspective of Niobrara fish research and an opportunity to evaluate the Stewart, 1900; Hay, 1903; Hussakof, 1908; Lane, 1945; Russell, 1988, current state of our knowledge of the Cretaceous ichthyofauna. 1993; Carpenter, 1990, 2003; Stewart, 1990a; Everhart, 2005c). Russell (1988) presented the first systematic list of vertebrate taxa from the Smoky METHODS Hill Chalk based on published literature supplemented with unpublished data. His study showed that the vertebrate paleofauna includes about 110 The occurrences of fossil vertebrates from the Niobrara Chalk have been described in various media. For the purposes of this study, we only taxa, of which about 60% are non-tetrapod vertebrates (= ‘fishes’) and 40% tetrapods. Recently, Carpenter (2003) examined the vertebrate bios- considered published papers that include specific catalogue numbers of specimens housed in public institutions. Data based on pictorial guidebooks tratigraphy of the Smoky Hill Chalk, which was based on his previous work (Carpenter, 1990) and partially similar to Stewart’s (1990a) work. (e.g., Liggett, 2001; McKinzie, 2002), unpublished dissertations and the- ses (e.g., Shimada, 1994d; Fielitz, 1999), as well as magazines, news me- One of the major shortcomings of Carpenter (1990, 2003) and Stewart’s 194 nology, Rapid City; UNSM, University of Nebraska State Museum, Lin- coln; USNM, United States National Museum of Natural History (Smithsonian Institution), Washington, D. C.; UW-Madison, University of Wisconsin at Madison (Geology Museum, Department of Geology and Geophysics); YPM, Yale University, Peabody Museum of Natural History, New Haven, Connecticut. SYSTEMATIC PALEONTOLOGY

Class Chondrichthyes Subclass Holocephali Order Chimaeriformes Obruchev, 1953 Family Callorhynchidae Garman, 1901 Genus Edaphodon Buckland, 1838 Edaphodon sp. FIGURE 1. George F. Sternberg (far left), prominent Niobrara fossil hunter, Referred material–BMNH P10343, dorsal fin spine (Stahl, 1999, excavating skeleton of Xiphactinus audax Leidy, 1870 (FHSM VP-333), in June fig. 143). of 1952 from Smoky Hill Chalk of Gove County, Kansas (photograph courtesy of FHSM). Note that this Xiphactinus skeleton contains skeleton of Gillicus arcuatus Additional remarks (at order level)–Ichthypriapus hubbsi (Cope, 1875) (FHSM VP-334) inside its abdominal cavity, and is probably the most Hibbard, 1942, initially placed in “chimaeroid,” is now considered to be frequently illustrated specimen of this taxon (e.g., Bardack, 1965a, pl. 1, fig. A; nomen dubium. The type specimen (KUVP 1136) is bony and considered Schwimmer et al., 1997, fig. 1A; Bottjer, 2002, fig. 19.5; Everhart, 2005c, fig. 5.1). to be part of partial skeleton of Protosphyraena gladius (Cope, 1873) dia, and other forms of non-technical publications (e.g., Sternberg, 1900; (KUVP 465), with which was discovered associated (Schultze et al., 1982, Anonymous, 1947; Walker, 1982) are not included. We chose to include p. 4; Stahl, 1999, p. 154). data that are in print in published abstracts for professional scientific meet- Subclass Elasmobranchii ings as long as referable catalogue numbers of specimens are given. Al- Order incertae sedis though a formal description of such specimens is pending, Figure 2 is pre- sented to substantiate their existence. Family Ptychodontidae Jaekel, 1898 We use a widely accepted taxonomic scheme (e.g., Cappetta, 1987; Genus Ptychodus Agassiz, 1835 Nelson, 1994) combined with information in some more recently published Ptychodus anonymus Williston, 1900a taxonomic literature (e.g., Taverne and Gayet, 2005). For each species, we list referred catalogue numbers, anatomical parts represented in each speci- Referred material–AMNH FF 19553, 201 associated teeth, sev- men, and their literature citation. When a particular specimen is referred to eral vertebral centra, pieces of calcified cartilages, and oral denticles (part in multiple papers, we limit the list of references to literature that includes of this specimen, including a set of associated denticles, is curated in LACM the best description of the specimen and/or the most recent publication as LACM 138527); FHSM VP-14854, teeth and oral denticles (Everhart dealing with the specimen. Whereas a complete synonymy list is not given, et al., 2003; Everhart and Caggiano, 2004; Everhart, 2005c, p. 63). nominal taxa are given for each species. Wherever appropriate, additional Notes–The taxonomy of Ptychodus anonymus is problematic be- taxonomic notes and other relevant information are given as “Notes” for cause the type materials of this taxon (Williston, 1900a, pl. 11, figs. 5–8, each species. Issues related to higher taxonomic levels of species or species 16–18, 20–22, 24; Williston, 1900b, pl. 29, figs. 5–8, 16–18, 20–22, 24), groups are discussed as “Additional remarks.” that were collected from pre-Niobrara deposits (“Benton” formation), ap- The following institutional abbreviations are used in this paper: pear to include teeth that belong to P. mammillaris (Williston, 1900b, pl. AMNH FF, American Museum of Natural History, Fossil Fish collection, 29, figs. 5, 6, 8, 16, 18: see Herman, 1977, p. 59). Williston (1900a, p. 32; New York, New York; BMNH, British Natural History Museum, London; 1900b, p. 241) and Stewart (1990a) noted the occurrence of P. anonymus BSPHG, Bayerische Staatssammlung für Paläontologie und Historishe from the Smoky Hill Chalk of Kansas (specimens unspecified). Everhart Geologie, München, Germany; CM, Carnegie Museum of Natural His- (2005c, fig. 4.7) erroneously noted an associated tooth set of P. anonymus tory, Pittsburgh, Pennsylvania; CMC VP, Cincinnati Museum Center, Ver- (FHSM VP-2223) as “P. mortoni.” tebrate Paleontology collection, Cincinnati, Ohio; CMN, Canadian Mu- Ptychodus cf. P. latissimus Agassiz, 1843 seum of Nature (formerly National Museum of Canada (NMC)), Ottawa, Referred material–FHSM VP-14853, tooth (Everhart et al., 2003; Ontario; CMNH, Cleveland Museum of Natural History, Cleveland, Ohio; Fig. 2A). DMNH, Denver Museum of Nature and Science, Colorado; ESU, Empo- Notes–The exact taxonomic placement of the specimen (Fig. 2A) is ria State University (Johnston Geology Museum), Kansas; FFHM, Fick uncertain within the genus Ptychodus because the total intraspecific mor- Fossil and History Museum, Oakley, Kansas; FHSM VP, Fort Hays State phological range of teeth in most Ptychodus species is poorly known. There University, Sternberg Museum of Natural History, Vertebrate Paleontol- is a possibility that the tooth may belong to one of the other Ptychodus ogy collection, Hays, Kansas; FMNH, Field Museum of Natural History, species described from the Smoky Hill Chalk of Kansas. Chicago, Illinois; KUIP, University of Kansas Museum of Natural History, Invertebrate Paleontology collection, Lawrence; KUVP, University of Kan- Ptychodus martini Williston, 1900a sas Museum of Natural History, Vertebrate Paleontology collection, Referred material–FHSM VP-2121, tooth (Hamm, 2002; Hamm Lawrence; LACM, Natural History Museum of Los Angeles County, Cali- and Shimada, 2004); KUVP 55277 (holotype), tooth set (for review, see fornia; MCZ, Museum of Comparative Zoology, Harvard University, Cam- Schultze et al., 1982, p. 13; Hamm and Shimada, 2004). bridge, Massachusetts; NAMAL, North American Museum of Ancient Life, Notes–Hamm and Everhart (1999) reported the occurrence of 170 Lehi, Utah; NJSM, New Jersey State Museum, Trenton; ROM, Royal associated teeth of Ptychodus martini, but the specimen remains Ontario Museum, Toronto, Canada; SDNHM, San Diego Natural History undescribed. Everhart et al. (2003) noted the occurrence of P. martini from Museum, California; SDSMT, South Dakota School of Mines and Tech- 195

FIGURE 2. Representative specimen of fossil fishes from Smoky Hill Chalk of Kansas that are substantiated only by published abstracts (i.e., pending formal descriptions by their authors). A, tooth of Ptychodus cf. P. latissimus (FHSM VP-14853, occlusal view, scale = 10 mm: Everhart et al., 2003); B, tooth of Cretalamna appendiculata (one of ca. 120 associated teeth in LACM 128126, lingual view, scale = 10 mm: Shimada, 2005b); C, tooth of Pseudocorax laevis (FHSM VP-13960, lingual view, scale = 5 mm: Hamm et al., 2003); D, tooth of Squalicorax volgensis (from fossiliferous rock specimen curated as FHSM VP-644, lingual view, scale = 1 mm: Beeson and Shimada, 2004); E, tooth of Rhinobatos sp. (from fossiliferous rock specimen curated as FHSM VP-644, top = occlusal view, bottom = lingual view, scale = 1 mm: Beeson and Shimada, 2004); F, tooth of Palaeobalistum sp. (from fossiliferous rock specimen curated as FHSM VP-644, top = occlusal view, bottom = profile view, scale = 1 mm: Beeson and Shimada, 2004); G, vomer of Albulidae (from fossiliferous rock specimen curated as FHSM VP-644, lingual view, scale = 1 mm: Beeson and Shimada, 2004); H, left mandible of Coelacanthiformes (part of LACM 131958, lingual view, scale = 10 mm: Stewart et al., 1991; Everhart, Everhart, and Stewart, 1995; Everhart, 2005c, p. 101). the Smoky Hill Chalk of Kansas (specimens unspecified). Everhart (2005c, apparently occurred in pre-Niobrara deposits (“Benton” formation: Williston, p. 65) erroneously reported the type specimen of P. martini (KUVP 55277) 1900b, p. 241, 243), dismissing the occurrence of P. mammillaris from as “FHSM VP-55271.” the Smoky Hill Chalk on this ground. Russell (1988) listed Ptychodus occidentalis Leidy, 1868, in the Ptychodus mortoni Mantell, 1839 Smoky Hill Chalk vertebrate fauna. However, P. occidentalis is a Referred material–FHSM VP-15013, tooth (Everhart et al., 2003); Cenomanian–Turonian taxon (see Welton and Farish, 1993, p. 64; Shimada KUVP 55269, tooth set; KUVP 55270, tooth set (for review, see Schultze et al., 2006), and the general area where Leidy’s (1868) original material et al., 1982, p. 14); KUVP 59041, partial skeleton (Stewart, 1980; was collected (“a few miles east of Fort Hays, Kansas,” p. 207) lacks expo- Schwimmer et al., 1997, p. 77; Everhart and Caggiano, 2004, p. 129; sures of the Smoky Hill Chalk, but those of the Greenhorn and Carlile Everhart, 2005c, p. 54). formations (Cenomanian–Turonian). This fact eliminates the Niobrara Notes–Stewart (1980) used preserved calcified vertebrae in record of P. occidentalis. Ptychodus mortoni (KUVP 59041) to suggest that Ptychodontidae is a Hattin (1996) described a cirriped-bearing coprolite-like specimen neoselachian shark (note: recent studies suggest that Ptychodontidae may (KUIP 286929) interpreted to be a possible Ptychodus regurgitate. be a hybodontiform (i.e., non-neoselachian) shark: Cuny et al., 2003, 2004, Kauffman (1972) suggested that breakage in inoceramid shells in the 2005). Niobrara Chalk is due to Ptychodus predation (see also Stewart, 1988). Bardack (1965b, p. 9) noted FHSM “10688-288” (uncertain catalogue Ptychodus polygyrus Agassiz, 1839 number) and VP-1652 as “Ptychodus sp.” Everhart and Caggiano (2004, Referred material–FHSM VP-76, VP-2123, and VP-15008, iso- p. 129) listed the following specimens to consist of “associated Ptychodus lated teeth (Caggiano and Everhart, 2003; Everhart et al., 2003; Everhart, tooth sets” (species unspecified): FHSM VP-2222, VP-2223, and VP- 2005c, p. 65); KUVP 55237, tooth (Williston, 1900b, p. 240; catalogue 14854. number based on Schultze et al., 1982, p. 14). Additional remarks (at genus level)–Russell (1988) cited Herman Order Lamniformes Berg, 1958 (1977, p. 59) for the occurrence of Ptychodus mammillaris Agassiz, 1839, Family Mitsukurinidae Jordan, 1898 from the Smoky Hill Chalk, which was based on teeth described by Williston Genus Scapanorhynchus Woodward, 1889 (1900b, pl. 29, figs. 2, 3, 5, 6, 8, 16, 18). However, those Ptychodus teeth Scapanorhynchus raphiodon (Agassiz, 1844) 196 Referred material–FHSM VP-13961, five associated teeth (Hamm 1997d; 1997e; 1997f; Everhart, 2000); KUVP 13522, and 64541–64549, and Shimada, 2002a, 2002b); KUVP 13522, one tooth (Williston, 1900a, isolated teeth (Williston, 1900a, 1900b; for review, see Schultze et al., 1982); 1900b; for review, see Schultze et al., 1982). ROM 44656, tooth set; UNSM 1216, tooth set with some vertebrae Notes–Williston’s (1900a, pl. 8, fig. 2, pl. 14, fig. 4; 1900b, pl. 26, (Shimada, 1997c, 1997e). fig. 2, pl. 32, fig. 4) Scapanorhynchus raphiodon tooth probably from the Notes–Until Glikman (1958) established the genus Cretoxyrhina, Niobrara Chalk of Kansas is considered to be a tooth of Cretoxyrhina this taxon was referred to Oxyrhina Agassiz, 1843 (e.g., Eastman, 1895; mantelli (KUVP 13522: Schultze et al., 1982, p. 7). Williston, 1900a, p. 36; Sternberg, 1907b) or Isurus Rafinesque, 1810 (e.g., Williston, 1900b, p. 246; Lane, 1945). Although further studies are Family Odontaspididae Müller and Henle, 1839 needed, Shimada’s (2005a) phylogenetic analysis suggested that Genus Johnlongia Siverson, 1996 Cretoxyrhina may be closely allied to Alopiidae Bonaparte, 1838, based on Johnlongia sp. the anatomical data taken from Niobrara specimens (e.g., Shimada, 1997d). Applegate (1970, p. 398) noted that KUVP houses an “Isurus [= Referred material–FHSM VP-15545, tooth (Everhart et al., Cretoxyrhina] mantelli” specimen that preserves scales associated with a 2004; Shimada, Ewell et al., 2004). skull. He reported it as “no. 199,” but Shimada (1997d, p. 647) was not Notes–Shimada, Ewell et al. (2004) noted that FHSM VP-15545 is able to locate such a specimen. Bardack (1965b, p. 9) noted FHSM VP- morphologically unique and may represent a new taxon within the genus 2081 and VP-2714 as “Isurus sp.” Shimada (1995, 1996b) discussed some Johnlongia. They also noted that its occurrence represents the skeletal specimens of C. mantelli from the Niobrara Chalk of Kansas but stratigraphically youngest Johnlongia specimen in the world. did not refer to any specific catalogue numbers. Shimada and Hubbell (2001) Additional remarks (at family level)–Russell (1988, p. 25) listed discussed small symmetrical teeth of C. mantelli from the Niobrara Chalk “Odontaspis sp.” in his faunal list of the Smoky Hill Chalk based on his of Kansas described by Eastman (1895). Shimada (2002) emended the personal communication with J. D. Stewart. However, the presence of the tooth type identification of C. mantelli proposed by Shimada (1997c), who genus Odontaspis from the stratigraphic unit cannot be confirmed because reconstructed the Cretoxyrhina dentition based on articulated tooth sets Russell (1988) did not refer to any specific specimen. from the Smoky Hill Chalk. Family Cretoxyrhinidae Glikman, 1958 In the Niobrara fossil record, putative bite marks of Cretoxyrhina Genus Cretalamna Glikman, 1958 mantelli are found on bones surfaces of mosasaurs (Everhart, 2005c, p. 50, 51, figs. 4.2, 4.3; Everhart, 2005e) and elasmosaurid plesiosaurs Cretalamna appendiculata (Agassiz, 1843) (Everhart, 2005a) as well as hadrosaurid and nodosaurid dinosaurs (Hamm Referred material–FHSM VP-14851, tooth; FHSM VP-14852, and Everhart, 2001; Everhart and Hamm, 2005; Everhart and Ewell, in tooth (Hamm et al., 2003); LACM 128126, partial skeleton including ca. press). Shimada et al. (2002) and Shimada and Everhart (2004) described 120 teeth (Shimada, 2005b; Fig. 2B). a Xiphactinus audax specimen from the Smoky Hill Chalk (ESU 1047: Notes–Williston (1900a, p. 37; 1900b, p. 247) suggested that this see below) that has an embedded tooth of Cretoxyrhina mantelli (unnum- species occurs from the Niobrara Chalk of Kansas but did not specify the bered). Everhart (2005c, p. 49–52, fig. 4.1; 2005e) discussed mosasaur specimens (and they may be lost: see Schultze et al., 1982). specimens from the Smoky Hill Chalk of Kansas (FHSM VP-13284 and Additional remarks (at genus level)–The genus has been com- VP-13742) with one or more embedded C. mantelli teeth (unnumbered). monly referred to Cretolamna, but as pointed out by Siverson (1999), the original spelling of the genus by Glikman (1958) is Cretalamna, and that Family Anacoracidae Casier, 1947 usage is followed here. Genus Pseudocorax Priem, 1897 Genus Cretoxyrhina Glikman, 1958 Pseudocorax laevis (Leriche, 1906) Cretoxyrhina mantelli (Agassiz, 1843) Referred material–FHSM VP-13959, tooth; FHSM VP-13960, tooth (Hamm et al., 2003; Fig. 2C). Referred material–CMN 40906, caudal and posterior precaudal Notes–Applegate (1970, p. 395) noted that FHSM has some teeth vertebrae with hypochordal rays and placoid scales (Shimada, Cumbaa et of Pseudocorax affinis (Agassiz, 1843), but he did not specify the speci- al., 2004, Shimada et al., this volume); FFHM 1972.196, tooth (Everhart, mens. Shimada (1997e) reported 20 associated teeth of “cf. Pseudocorax 2005a, p. 18); FHSM VP-75, tooth set; FHSM VP-323, partial skeleton; sp.” (KUVP 55060: the catalogue number refers to associated Cretoxyrhina FHSM VP-2184, partial skeleton; FHSM VP-2187, nearly complete skel- skeletal elements that are enclosed in the same matrix). Stewart (1990a) eton; FHSM VP-2224, tooth; FHSM VP-12850, tooth; FHSM VP-13105, and Hamm (2001) noted the occurrence of P. laevis (specimens unspeci- abnormal tooth; FHSM VP-13106, tooth; FHSM VP-13284 and VP-13285, fied). two separate teeth embedded in a series of mosasaur vertebrae (FHSM VP-13283) (Shimada, 1993a, 1993b, 1994a, 1994c, 1997a, 1997b, 1997c, Genus Squalicorax Whitley, 1939 1997d, 1997e, 1997f; Siverson, 1996, text-fig. 4; Everhart, Everhart, and Squalicorax falcatus (Agassiz, 1843) Shimada, 1995; Blanco-Piñón et al., 2005; Everhart, 2005c, p. 60, 61; Referred material–AMNH FF 1908, four teeth (holotype of Shimada et al., this volume); FHSM VP-644, teeth (the catalogue number Galeocerdo hartwellii Cope, 1872: Hussakof, 1908, p. 29, fig. 6); CMC is assigned to an entire fossiliferous rock specimen which yielded the teeth: VP-5722, partial skeleton including at least 104 teeth (Everhart, 2005c, p. Beeson and Shimada, 2004); FHSM VP-13747, tooth tip embedded in a 61; Shimada and Cicimurri, 2005); FHSM VP-644, teeth (the catalogue series of vertebrae with partial ribs of a mosasaur (FHSM VP-13746: number is assigned to an entire fossiliferous rock specimen which yielded Everhart, 2004a); FHSM VP-14010, partial skeleton (Corrado et al., 2003; the teeth: Beeson and Shimada, 2004); FHSM VP-13220, tooth; FHSM Everhart, 2005c, p. 59, 61); KUVP 213, teeth; KUVP 247, partial skel- VP-13222, three teeth (Shimada, 1997e, p. 929); KUVP 34092, partial eton; KUVP 1094, tooth (the catalogue number refers to a series of mosa- skeleton (Stewart, 1978, 1999b); KUVP 34916, vertebrae and teeth saur vertebrae in which the tooth is embedded within); KUVP 49490, pec- (Shimada and Cicimurri, 2005); KUVP 64511, 64515–64520, 64523, toral fin; KUVP 13520, tooth set; KUVP 13523, isolated teeth; KUVP 64525, 64527–64529, 64531, 64532, 64534–64536, 64538, and 64540, 13536, tooth; KUVP 13589, isolated teeth; KUVP 40337; isolated teeth; isolated teeth (Williston, 1900a, 1900b; as “S. cf. kaupi” in Schultze et al., KUVP 55060, fragmentary skeleton; KUVP 57294, tooth; KUVP 66120, 1982, p. 11, 12); LACM 16056, nearly complete neurocranium with teeth tooth; KUVP 68979, fragmentary skeleton; KUVP 68980, tooth; KUVP and jaw fragment (Shimada and Cicimurri, 2005; cf. Compagno, 1988, 69102, partial skeleton; KUVP 69707, isolated teeth; KUVP 84872, tooth; 1990); LACM 120090, partial skeleton (Schwimmer et al., 1997, p. 76; KUVP 86418, tooth; KUVP 112807, tooth (Shimada, 1993b, 1997c; 197 Shimada and Cicimurri, 2005); LACM 135929, partial skeleton with at Referred material–FHSM VP-644, “at least 70 teeth” (the cata- least 42 teeth; LACM 143536, ca. 20 teeth, placoid scales, and calcified logue number is assigned to an entire fossiliferous rock specimen which cartilage pieces; LACM 143537, partial skeleton with at least 70 teeth; yielded the teeth: Beeson and Shimada, 2004; Fig. 2E). USNM 425665, nearly complete skeleton (Shimada, 2000; Shimada and Notes–Stewart (1990a, p. 24) and Everhart (2005c, table 13.1; Cicimurri, 2001, 2005; cf. Frickhinger, 1995, p. 180); UW-Madison NS 2005d) noted the occurrence of Rhinobatos sp. and Rhinobatos incertus, 1509.34, partial skeleton with at least 50 teeth (Shimada and Cicimurri, respectively, from the Smoky Hill Chalk (specimens unspecified). 2005). Additional remarks (at subclass level)–Stewart (1978, 1999b) Notes–Galeocerdo hartwellii Cope, 1872, described only from the inferred certain enterospirae (KUVP 25868, and 43068–43075) from the Niobrara Chalk of Kansas, is a synonym of Squalicorax falcatus. Everhart Smoky Hill Chalk to be selachian in origin. Stewart and Carpenter (1990) (2004b) reported a tooth of S. falcatus associated with a plesiosaur skel- and Hattin (1996) described several coprolites of unknown origin (possi- eton (USNM 9468). Everhart (2005c) illustrated a tooth of S. falcatus bly “piscine origin”: Stewart and Carpenter, 1990, p. 205). As noted by (unnumbered specimen: fig. 4.4) and putative bite marks of S. falcatus on Everhart (2005c, p. 62), there is an undescribed species of Galeorhinus a mosasaur (fig. 4.5). Everhart (2005c, p. 61) erroneously reported USNM Blainville, 1816 (Carcharhiniformes Compagno, 1973: Triakidae Gray, 425665 as “USNM 423665.” 1851), in the Smoky Hill Chalk (J. D. Stewart, personal commun. 2002– 2005). Squalicorax kaupi (Agassiz, 1843) Referred material–AMNH FF 1909, two teeth (holotype of Class Osteichthyes Galeocerdo crassidens Cope, 1872; as “Corax falcatus” in Hussakof, 1908, Subclass Actinopterygii p. 29); FHSM VP-2213, partial jaws with teeth (Shimada, 1994b; Everhart, Superorder Neopterygii 2005c, p. 58, 61; Shimada and Cicimurri, 2005); FHSM VP-13219, tooth (Squalicorax cf. S. kaupi: Shimada, 1997e, p. 929); KUVP 55188, 26 Order Pycnodontiformes Berg, 1940 teeth and some calcified cartilage pieces; KUVP 55190, vertebrae and teeth Family Pycnodontidae Owen, 1846 (Shimada and Cicimurri, 2005); KUVP 64501–64510, 64512–64514, Genus Micropycnodon Hibbard and Graffham, 1945 64521, 64522, 64524, 64526, 64530, 64533, 64537, and 64539, teeth Micropycnodon kansasensis (Hibbard and Graffham, 1941) (as “C. falcatus” in Williston, 1900a, 1900b; for review, see Schultze et al., 1982, p. 10, 11); LACM 128007, vertebrae and teeth (Shimada and Referred material–KUVP 1019 (holotype), partial skull; KUVP Cicimurri, 2005); YPM 56409, two teeth (Everhart, 2005c, p. 55). 7030, partial skeleton (Dunkle and Hibbard, 1946; Everhart, 2005c, p. Notes–Galeocerdo crassidens Cope, 1872, described only from the 98). Niobrara Chalk of Kansas, is a synonym of Squalicorax kaupi. Compagno Notes–Hibbard and Graffham (1941) initially described the genus (1988, p. 404; 1990, p. 373, 374) described a partial skeleton of Squalicorax as Pycnomicrodon, which was later modified to Micropycnodon because (LACM 16056), which he referred to it as “S. ‘falcatus’ (possibly = S. the former name was preoccupied (Hibbard and Graffham, 1945). The pristodontus).” The specimen could be S. kaupi based on this description. two referred specimens were originally reported as from the Fort Hays Lime- Shimada and Cicimurri (2005) described a S. kaupi specimen consisting stone, but they are now thought to have come from the Smoky Hill Chalk of 42 teeth, placoid scales, and some calcified cartilage pieces (KUVP 229) (Schultze et al., 1982, p. 22; but see also Stewart, 1990a, p. 21). that could have come from the Niobrara Chalk of Kansas. Everhart (2005c, Genus Hadrodus Leidy, 1873 fig. 4.4) illustrated a tooth of S. kaupi (unnumbered specimen). Hadrodus marshi Gregory, 1950 Squalicorax pristodontus (Agassiz, 1843) Referred material–YPM 1950 (holotype), partial skull (Gregory, 1950). Referred material–FHSM VP-15009, tooth; FHSM VP-15010, Family Nursallidae Blot, 1987 tooth (Hamm et al., 2003; Everhart, 2005c, p. 58; Shimada and Cicimurri, this volume). Genus Palaeobalistum Blainville, 1818 Notes–Everhart (2005c) illustrated a tooth of Squalicorax Palaeobalistum sp. pristodontus (unnumbered specimen: fig. 4.4) and erroneously reported Referred material–FHSM VP-644, teeth (the catalogue number is FHSM VP-15011 (tooth of S. kaupi: see Shimada and Cicimurri, this vol- assigned to an entire fossiliferous rock specimen which yielded the teeth: ume) as a tooth of S. pristodontus (p. 58). Beeson and Shimada, 2004; Fig. 2F). Additional remarks (at order level)–Stewart (1990a, p. 24) noted Squalicorax volgensis (Glikman in Glikman and Shvazhaite, 1971) the occurrence of “an undetermined pycnodont that is not Micropycnodon Referred material–FHSM VP-644, tooth (the catalogue number and is probably not Hadrodus” (specimens unspecified). is assigned to an entire fossiliferous rock specimen which yielded the tooth: Beeson and Shimada, 2004; Fig. 2D). Order Semionotiformes Aramboug and Bertin, 1958 Notes–Based on literature (e.g., Siverson, 1996; Cappetta and Case, Family Lepisosteidae Cuvier, 1825 1999; Cicimurri, 2001), Squalicorax volgensis in FHSM VP-644 repre- Genus Lepisosteus Lacépède 1803 sents the youngest geologic record of the taxon (Shimada and Cicimurri, this volume). Lepisosteus sp. Additional remarks (at genus level)–Until Whitley (1939) estab- Referred material–KUVP 36243, fragmentary skeleton and scales lished the genus Squalicorax, the taxon was commonly referred to Corax (Wiley and Stewart, 1977; Everhart, 2005c, p. 99). Agassiz, 1843. Everhart (2004a, 2005c, p. 53) discussed mosasaur speci- mens (FHSM VP-3 and VP-13746) from the Smoky Hill Chalk of Kansas Order Pachycormiformes Berg, 1940 with putative Squalicorax bite marks. Family Pachycormidae Woodward, 1895 Order Rajiformes Berg, 1940 Genus Protosphyraena Leidy, 1857 Protosphyraena gladius (Cope, 1873) Family Rhinobatidae Müller and Henle, 1838 Genus Rhinobatos Linck, 1790 Referred material–AMNH FF 1849 (holotype), pectoral fin; AMNH FF 2064, pectoral fin (Hay, 1903, p. 24, 25; Hussakof, 1908, p. Rhinobatos sp. 93); KUVP 439, 465, 49505, 60620, 66692, and 84867, specimens that 198 include or consist of pectoral fin (Stewart, 1988); KUVP 60692, pectoral Additional remarks (at genus level)–Species of Protosphyraena fin and additional skeletal elements (Schwimmer et al., 1997, p. 77); LACM were initially referred to the genus Erisichthe Cope, 1873 (see also Cope, 126520, pectoral fin (Stewart, 1988). 1877c). The taxonomic validity of the following three Protosphyraena species from the Niobrara Chalk of Kansas is uncertain: P. dimidiata (Cope, Protosphyraena nitida (Cope, 1872) 1878) (AMNH FF 1635, holotype consisting of partial skull and vertebra: Referred material– AMNH FF 1634, partial skull (Hay, 1903); Hay, 1903, p. 16–20; Hussakof, 1908, p. 92); P. recurvirostris Stewart, AMNH FF 1893, partial skull (listed as the holotype by Hussakof, 1908, p. 1898d (see also Stewart, 1900; KUVP 373, holotype consisting of ros- 93: cf. AMNH FF 2121); AMNH FF 1894, partial pectoral fin; AMNH FF trum: Schultze et al., 1982, p. 24); and P. sequax Hay, 1903 (based on 2105 (holotype of Erisichthe penetrans Cope, 1877c), rostrum (Hussakof, Felix’s (1890) specimen of P. nitida skull: Hay, 1903, p. 20–22). These 1908, p. 94); AMNH FF 2121 (holotype), partial skull and pectoral fin species are based on fin-less specimens, where the taxonomic identifica- (Hay, 1903; Everhart, 2005c, p. 89); FHSM VP-3251, pectoral fin (Stewart, tion of Protosphyraena species generally requires the knowledge about 1988); KUVP 374, 417, 505, 12490, 49515, 55588, 55589, 60603, 60615, the morphology of pectoral fins (e.g., Stewart, 1988). In addition, no re- and 64495, specimens that include or consist of pectoral fin (Stewart, 1988); ferred material can be relocated for P. occidentalis Stewart, 1900, from the KUVP 49514, pectoral fin; KUVP 125302, pectoral girdle and fin; LACM Smoky Hill Chalk of western Kansas, and it is presently treated as 129752, complete skull and fins (Everhart, 2005c, p. 92, fig. 5.6); LACM Protosphyraena sp. (see Schultze et al., 1982, p. 24). 133270, pectoral fin (Schwimmer et al., 1997, p. 77); NJSM 15021, par- Crook (1892, fig. 1.9) illustrated a Protosphyraena specimen (cata- tial skull; NJSM 15839, partial skull (Everhart, 2005c, p. 92). logue number unspecified). Stewart (1898a, p. 28) described a new, un- Notes–Erisichthe nitida Cope, 1872, E. penetrans Cope, 1877c, named species of Protosphyraena on the basis of a premaxillary specimen. Pelecopterus chirurgus Cope, 1875, Protosphyraena chirurgus Cope, He thought it might be a synonym of either P. penetrans (Cope, 1877c) or 1877c, Newton (1878) and Felix’s (1890) P. penetrans, and P. obliquidens “P. xiphoides” (a probable misspelling of P. ziphioides (Cope, 1877c), a Loomis, 1900, are all interpreted to be junior synonyms of P. nitida (e.g., species originally described as an Erisichthe taxon but now considered as a see Hay, 1903). Martinichthys taxon: see below). Sternberg (1913) described an unnamed species of Protosphyraena based on a complete set of pectoral fins. Hussakof Protosphyraena perniciosa (Cope, 1874) (1908, p. 95) listed AMNH FF 1646 (fragmentary pectoral fin) as Referred material–AMNH FF 1609 (paratype of Pelecopterus “Protosphyraena sp.” McClung (1908, pl. 13) reported the only known chirurgus Cope, 1875), partial pectoral girdle (described as Protosphyraena caudal fin of Protosphyraena (KUVP 55500: Everhart, 2005c, p. 93) in nitida by Hussakof, 1908, p. 94); AMNH FF 1620, pectoral fin (Hay, 1903, the Niobrara fossil record. Bardack (1965b, p. 9) noted FHSM VP-1659, p. 9–16; Stewart, 1988; listed as P. tenuis (Loomis, 1900) by Hussakof, VP-1660, VP-2024, and VP-2025 as “Protosphyraena sp.” Everhart 1908, p. 94); AMNH FF 1850 (holotype), pectoral fin spine; AMNH FF (2005c) noted that at least 14 specimens of Protosphyraena are housed in 1895, fragmentary pectoral fin; AMNH FF 1901, pectoral girdle and fin; YPM (e.g., YPM 42137, 42138, 42152, 42200, and 42285: p. 89), and he AMNH FF 2009, pectoral girdle (Hussakof, 1908, p. 94); AMNH FF 2058, illustrated four Protosphyraena teeth (unnumbered specimens: fig. 5.4). pectoral fin (Stewart, 1988); FHSM VP-80, VP-4510, and VP-7623, speci- mens that include or consist of pectoral fin (Stewart, 1988); FHSM VP- Order Amiiformes Hay, 1929 10266, pectoral fin; FHSM VP-12059, pectoral fin base (Schwimmer et Family incertae sedis (see Grande and Bemis, 1998) al., 1997, p. 77, 78); KUVP 356, 363, 366, 369, 372, 424–426, 431, 432, Genus Paraliodesmus Dunkle, 1969 440, 13588, 13706, 40294–40296, 49422, 49511, 49512, 49550, 49551, 55493, 55590, 55591, 56601–56603, 56611, 60604, 64496, and 64497, Paraliodesmus guadagnii Dunkle, 1969 specimens that include or consist of pectoral fin (Stewart, 1988); KUVP Referred material–FMNH PF14383, three nearly complete skel- 49530, hypural plate; KUVP 55500, caudal fin (McClung, 1908); KUVP etons (Grande and Bemis, 1998, p. 11); USNM 21083 (holotype), partial 56614, pectoral girdles and fin (McClung, 1908; for review, see Schultze skeleton (Dunkle, 1969; Bardack, 1976; Everhart, 2005c, p. 99). et al., 1982); KUVP 67877, caudal skeleton (Arratia and Lambers, 1996, Notes–The holotype (Stewart, 1990b, 1990c) is on a valve of the p. 192); LACM 126004 and 126005, specimens that include or consist of inoceramid bivalve, Platyceramus platinus (Logan, 1898). Stewart (1996, pectoral fin; UNSM 2-9-9-37, pectoral fin (Stewart, 1988). p. 390) discussed another Paraliodesmus guadagnii specimen found with Notes–AMNH FF 1609, now identified as P. perniciosa, was ini- an unnamed holocentrid fish (KUVP 82576). Grande and Bemis (1998, p. tially referred to as Pelecopterus chirurgus by Cope (1875), a synonym of 618) noted that Paraliodesmus guadagnii possibly belongs to the subfam- Protosphyraena nitida. Hay (1903, p. 8) pointed out that the specimen is ily Caturoidea Owen, 1860. not Pelecopterus chirurgus. Stewart (1988) misspelled Protosphyraena Additional remarks (at order level)–Bardack (1976) referred to perniciosa as “P. pernicosa” (Everhart, 2005c, p. 91, 92). Everhart (2005c, an undescribed amioid (“SMM 11324,” old catalogue number used at fig. 5.5) illustrated a pectoral fin of P. perniciosa (unnumbered specimen). FHSM) on a bowl-shaped shell of the inoceramid bivalve, Volviceramus grandis (Conrad, 1875). He also noted that fragmentary amioid remains in Protosphyraena tenuis (Loomis, 1900) “KU 24201” and “24202” are similar to “SMM 11324.” Referred material– AMNH FF 205 and 211, specimens that in- clude or consist of pectoral fin (Stewart, 1988); AMNH FF 2061, fragmen- Order Asarotoformes Schaeffer, 1968 tary pectoral fin (Hussakof, 1908, p. 94); KUVP 380, 384, 385, and 13587, Family Asarotidae Schaeffer, 1968 pectoral fin (Stewart, 1900; for review, see Schultze et al., 1982, p. 24, Genus Asarotus Schaeffer, 1968 25); KUVP 426, 427, 429, 430, 49529, 49549, 55586, 55587, 55617, Asarotus arcanus Schaeffer, 1968 60616, 70216, and 84807, specimens that include or consist of pectoral fin (Stewart, 1988); KUVP 433, pectoral fin; KUVP 125303, pectoral girdle Referred material–AMNH FF 5555 (holotype), disarticulated skull and fin (Schwimmer et al., 1997, p. 77); KUVP 49419, posterior part of and anterior portion of body. skeleton including caudal fin of “P. cf. P. tenuis”; KUVP 119500, hypural Additional remarks (at subclass level)–Everhart (2005c, tables plate (Arratia and Lambers, 1996, p. 192); LACM 135837, pectoral fin 5.1, 13.1) listed the presence of an aspidorhynchiform fish, Belonostomus (Schwimmer et al., 1997, p. 77). (Lambe, 1902) (Aspidorhynchidae Nicholson and Lydekker, 1889), in the Notes–Stewart’s (1900) specimens of Protosphyraena penetrans Smoky Hill Chalk (specimens unspecified; see also Stewart, 1900, p. 24, (Cope, 1877c) are interpreted to belong to P. tenuis (see Schultze et al., 29). 1982). 199 Division Teleostei Stewart, 1900. Crook (1892, fig. 1.8) illustrated an I. ctenodon specimen Order Ichthyodectiformes Bardack and Sprinkle, 1969 as I. anaides (catalogue number unspecified). Family Ichthyodectidae Crook, 1892 Genus Xiphactinus Leidy, 1870 Genus Gillicus Hay, 1898b Xiphactinus audax Leidy, 1870 Gillicus arcuatus (Cope, 1875) Referred material–AMNH FF 993, nearly complete skeleton (de- Referred material–AMNH FF 2326 (neotype), cranial elements scribed as Portheus molossus Cope, 1871, in Hussakof, 1908, p. 88); (Bardack, 1965a); AMNH FF 7377, partial skull and scales (Hussakof, AMNH FF 1665 (holotype of P. molossus), partial skull; AMNH FF 1682 1908, p. 74); AMNH FF 8582, 8586–8589, and 8603, cranial elements (holotype of P. thaumas Cope, 1870), pectoral fin rays and caudal verte- (Bardack, 1965a); AMNH FF 8563, mandible (Nelson, 1973, fig. 6A); brae; AMNH FF 1732, skull (described as P. molossus in Hussakof, 1908, CMNH 10118, neurocranium (Bardack, 1965a); FHSM VP-334 (referred p. 88); AMNH FF 1673, 1706, 1747, 2373, 7382, 8112, 8115, 8261, 8272, to “FH 5026” by Bardack, 1965a, and “SMM 5026” by Bardack, 1976), and 8590, cranial elements and/or vertebrae; AMNH FF 1719, cranial ele- nearly complete specimen inside abdominal cavity of Xiphactinus audax ments and pectoral girdle; AMNH FF 2332, fin ray fragments; AMNH FF (FHSM VP-333) (Shimada and Everhart, 2004; Everhart, 2005c, p. 72, 8443, nearly complete skeleton (Bardack, 1965a, Nelson, 1973, fig. 3C); fig. 5.1); KUVP 3, 80, 127, 129, 133–135, 138, 143, 146, 339, 346, 478, AMNH FF 1903 (holotype of P. lestrio Cope, 1873), partial skull and ver- 738, 965, 10321, 11667, and 11669, cranial elements and/or pectoral girdle; tebrae (Hussakof, 1908, p. 88); AMNH FF 7296, pelvic girdle and fins; USNM 21081, cranial elements (Bardack, 1965a). AMNH FF 7318, skull (described as P. molossus in Hussakof, 1908, p. Notes–Portheus arcuatus Cope, 1875, is a junior synonym of 88); AMNH FF 7350, premaxilla and maxilla (Schwimmer et al., 1997; Gillicus arcuatus. The holotype could not be located, so Bardack (1965a) described as P. thaumas in Hussakof, 1908, p. 90); AMNH FF 322199, selected AMNH FF 2326 as a neotype. McClung (1908) described the nearly complete skeleton (Osborn, 1904; see Everhart, 2005c, p. 76); occurrence of G. arcuatus from the Niobrara Chalk of Kansas (specimens BMNH P.11125, composite skeleton (Woodward, 1913); CM 1189, neu- unspecified). rocranium and vertebrae; CM 4101 and 4102, composite of two individu- als; CMN 8151, composite skeleton; CMN 8152 and 8152A, anterior por- Genus Ichthyodectes Cope, 1870 tion of fish; CMNH 10421, various skeletal parts; DMNH 1667, nearly Ichthyodectes ctenodon Cope, 1870 complete skeleton (Bardack, 1965a; Everhart, 2005c, p. 72, 73); ESU 1047, Referred material–AMNH FF 1611, partial skull (described as nearly complete skull and vertebrae (Shimada et al., 2002; Shimada and “Ichthyodectes anaides Cope, 1872” in Hussakof, 1908, p. 76); AMNH Everhart, 2004); FHSM VP-333 (Fig. 1; referred to “FH 5026” by Bardack, FF 1622, neurocranium (Bardack, 1965a); AMNH FF 1628, vertebrae (de- 1965a), nearly complete specimen containing Gillicus arcuatus (FHSM scribed as “I. anaides” in Hussakof, 1908, p. 76); AMNH FF 1633, man- VP-334) inside its abdominal cavity (Schwimmer et al., 1997, fig. 1A; dible; AMNH FF 1743, neurocranium, maxilla, and pectoral girdle (de- Shimada and Everhart, 2004; Everhart, 2005c, p. 72, fig. 5.1); FHSM VP- scribed as “I. multidentatus Cope, 1872” in Hussakof, 1908, p. 76); AMNH 8395, cranial elements; FMNH UF973, nearly complete skeleton (Bardack, FF 1810, neurocranium and jaws (Bardack, 1965a); AMNH FF 1905, jaws, 1965a, 1965b, p. 9); KUVP 1, 2, 3, 4, 88, 127, 135, 179, 275, 279, 287, vertebrae, and fin spines (described as “I. hamatus Cope, 1872” in Hussakof, 314, and 353, jaw elements (Stewart, 1898b); KUVP 103, composite skel- 1908, p. 76); AMNH FF 1910 (holotype), maxillaries, quadrates, and ver- eton; KUVP 124, 158, 160, 165, 166, 168, 183, 262, 320, 477, 565, 735, tebrae (Hussakof, 1908, fig. 76; Bardack, 1965a); AMNH FF 1939, frag- 929, 930, 991, 9820, 9888, 11661, 11834, and 12011, cranial elements, mentary skeleton (Nelson, 1973, fig. 3B; see also Hussakof, 1908, p. 75); vertebrae, and/or pectoral girdle (Stewart, 1900; Bardack, 1965a; Everhart, AMNH FF 1997 (paratype of I. anaides), mandibles, fin rays, and verte- 2005c, p. 81); KUVP 155, cranial elements (holotype of Xiphactinus brae (Hussakof, 1908, p. 76); AMNH FF 2005, skull (Bardack, 1965a); brachygnathus Stewart, 1899c); KUVP 245, partial skeletal elements as AMNH FF 2091 (holotype of I. acanthicus Cope, 1877b), parasphenoid, gastric residues of Cretoxyrhina mantelli (KUVP 247: Shimada, 1997e); fragmentary dentary, and vertebrae (Hussakof, 1908, p. 75, fig. 39); AMNH NAMAL 2000-0925-099, complete skeleton (Shimada and Everhart, 2004; FF 2186 (holotype of I. multidentatus), premaxillary and part of maxillary Everhart, 2005c, p. 73); SDNHM 63.01 and 63.02, nearly complete skel- (Hussakof, 1908, p. 76); AMNH FF 7356, maxilla (described as “I. anaides” eton; SDSMT 2510 (now at Science Museum of Minnesota, St. Paul), in Hussakof, 1908, p. 76); AMNH FF 8572, partial neurocranium; AMNH partial skeleton; USNM 52 (holotype), partial first pectoral fin ray; USNM FF 8573, premaxilla; AMNH FF 8584, mandible; FHSM VP-8567, nearly 1495, nearly complete skeleton; USNM 1646, 3782, 4207, 11554, and complete skeleton (Bardack, 1965a; as “Ichthyodectes sp.” in Bardack, 11653, cranial elements; USNM 11178, vertebra; USNM 11650 and 21375, 1965b); FHSM VP-644, teeth (the catalogue number is assigned to an nearly complete skeleton (Bardack, 1965a; Everhart, 2005c, p. 76, fig. entire fossiliferous rock specimen which yielded the teeth: Beeson and 2.2); YPM 2177, nearly complete skeleton (Thorpe, 1934). Shimada, 2004); KUVP 78, disarticulated skull and vertebrae; KUVP 88, Notes–Synonyms of Xiphactinus audax are: Saurocephalus audax mandibles and right maxilla; KUVP 97, hyomandibula and jaw fragments; Cope, 1870, S. thaumas Cope, 1870, Portheus molossus Cope, 1871 (“P. KUVP 102, mandibles and pectoral and pelvic fin rays (Stewart, 1900); mollosus,” a probable misspelling, in Sternberg, 1905, p. 127; “P. colos- KUVP 104, composite of two individuals (Bardack, 1965a); KUVP 117, sus,” a probable misspelling, in Sternberg, 1907b, p. 122), P. thaumas Cope, neurocranium and vertebrae; KUVP 118, mandibles and right maxilla 1872, P. lestrio Cope, 1873, P. mudgei Cope, 1874, P. lowii Stewart, 1898a, (syntype of I. parvis Stewart, 1900); KUVP 125, skull parts and vertebrae and X. brachygnathus Stewart, 1899c. Hussakof (1908, p. 90) listed the (Stewart, 1900); KUVP 132, partial neurocranium; KUVP 357, skull; following specimens as “Portheus sp.”: AMNH FF 1683 (fragmentary KUVP 934, pectoral girdle; KUVP 944, fin ray fragments; KUVP 11662, pectoral fin rays), 1684 (scapula and partial pectoral fin), 7342 (fragmen- skull and pectoral girdle; KUVP 12012, skull and jaws; KUVP 12023, tary mandible), 7360 (pectoral girdles), and 7388 (three neural arches). pelvic girdle; USNM 12358, nearly complete skeleton (Bardack, 1965a). Bardack (1965a, table 3) reported another composite specimen of this spe- Notes–Leidy (1857, 1858, 1873) referred this species to cies housed in Oakley, Kansas Public School numbered 13-25. Other pa- ?Cladocyclus occidentalis, which was referred to as Icthyodectes pers that describe the occurrence of this taxon from the Niobrara Chalk of occidentalis by Loomis (1900). Hussakof (1908, p. 77) listed AMNH FF Kansas include papers by Crook (1892), Hay (1898a), Stewart (1899a), 1912 (fragmentary skeleton), the holotype of I. prognathus (Cope, 1870), Osborn (1904), Sternberg (1905, 1907a, 1907b), and McClung (1908). which is a probable synonym of I. ctenodon. Other synonyms of I. ctenodon Beamon (2001) discussed a X. audax specimen in FHSM without refer- are: I. anaides Cope, 1872, I. hamatus Cope, 1872, I. multidentatus Cope, ring to a specific catalogue number, but his description suggests that it is 1872, I. cruentus Hay, 1898b, I. acanthicus Cope, 1877b, and I. parvis FHSM VP-333 (Fig. 1). Everhart (2004b) reported some vertebra of X. 200 audax (unnumbered) associated with a plesiosaur remain (USNM 9468) 34; Hussakof, 1908, p. 62; Nelson, 1973, fig. 6D; Taverne, 2001c). from the Smoky Hill Chalk of Kansas. Additional remarks (at genus level)–Species of Bananogmius have been referred to the genera Beryx Stewart, 1898d, Anogmius Cope, Family Saurodontidae Cope, 1871 1877a (spelled Agnomius by Loomis, 1900), Osmeroides Agassiz, 1843, Genus Prosaurodon Stewart, 1999a and Ananogmius White and Moy-Thomas, 1940 (for review, see Schultze Prosaurodon pygmaeus (Loomis, 1900) et al., 1982; Taverne, 2001c). Nelson (1973, figs. 6C, 7) illustrated AMNH Referred material–BSPHG 1893x43, mandible; FMNH PF3337, FF 3072, a partial skeleton of “Bananogmius sp.” partial skeleton; KUVP 56637, mandible; KUVP 56638, mandibles; KUVP Genus Luxilites Jordan, 1924 56639, mandible; LACM 128316, jaw elements; LACM 141422, partial Luxilites striolatus Jordan, 1924 skull (Stewart, 1999a). Referred material–KUVP 295 (holotype), skull and trunk (Taverne, Genus Saurocephalus Harlan, 1824 2002a; Everhart, 2005c, p. 97). Notes–The holotype is the only known specimen of this taxon. Saurocephalus lanciformis Harlan, 1824 Referred material–AMNH FF 2073, fragmentary maxilla Genus Syntegmodus Loomis, 1900 (Hussakof, 1908, p. 97); AMNH FF 7355 (holotype of Saurocephalus Syntegmodus altus Loomis, 1900 pamphagus Hay, 1899), skull parts; AMNH FF 7373, jaw parts; FMNH Referred material– AMNH FF 2112 (neotype), partial skull PF902, neurocranium and jaws; KUVP 109, mandibles; KUVP 154, up- (Taverne, 2001b; Everhart, 2005c, p. 96). per jaw and predentary (Bardack and Sprinkle, 1969). Notes–Whereas Hussakof (1908, p. 61) identified AMNH FF 2112 Notes–Synonyms of Saurocephalus lanciformis are: S. arapahovius as “Anogmius altus?,” Hay (1903, p. 41–46) placed Syntegmodus in Cope, 1872, S. dentatus Stewart, 1898a, and S. pamphagus Hay, 1899. Anogmius Cope, 1877a, a junior homonym replaced by Bananogmius The species was also discussed by Loomis (1900). Whitley, 1940. Recently, Taverne (2001b) suggested that Syntegmodus is Genus Saurodon Hays, 1830 a distinct genus and is not a synonym of Bananogmius. The holotype was destroyed during the Second World War in Germany, and the neotype is the Saurodon leanus Hays, 1830 only known specimen of this taxon. Referred material–AMNH FF 1614, mandible (Hussakof, 1908, p. 98; Nelson, 1973, fig. 3A); AMNH FF 1648, skull (Bardack and Sprinkle, Genus Niobrara Jordan, 1924 1969); AMNH FF 1906 (holotype of Saurocephalus phlebotomus Cope, Niobrara encarsia Jordan, 1924 1870), partial skull (Hussakof, 1908, p. 98); AMNH FF 1907, jaws and Referred material–KUVP 179 (holotype), complete skeleton palatine; AMNH FF 2110 (holotype of Saurocephalus goodeanus (Cope, (Taverne, 2001a; Everhart, 2005c, p. 97). 1877b)), partial skull, AMNH FF 7354, neurocranium and jaws; AMNH Notes–The holotype is the only known specimen of this taxon. FF 8323, pectoral girdle; AMNH FF 8544, partial skull and vertebrae; FHSM VP-11322, skull and vertebral column; FMNH PF1333, skull; FMNH Genus Martinichthys McClung, 1926 PF3337, skull, vertebrae, and fins; FMNH PF3741, complete fish; KUVP Martinichthys brevis McClung, 1926 142, mandibles and quadrate; KUVP 153, maxilla and predentary; KUVP Referred material–FHSM VP-15567, rostrum (Everhart, 2005c, 155, maxilla and dentary; KUVP 161, skull (holotype of Saurodon fig. 5.7); KUVP 497 (holotype), nearly complete skull; KUVP 40015, par- xiphirostris Stewart, 1898c); KUVP 212 (holotype of Daptinus broadheadi tial skull (Taverne, 2000b; Everhart, 2005c, p. 93). Stewart, 1898a); KUVP 343, skull; KUVP 28713, mandibles and right maxilla (Bardack and Sprinkle, 1969); KUVP 180, skull (Everhart, 2005c, Martinichthys ziphioides (Cope, 1877c) fig. 5.2); MCZ 5342, jaws; YPM 3942, neurocranium (Bardack and Referred material–AMNH FF 2131 (holotype), rostrum (Hay, Sprinkle, 1969). 1903, p. 22–24; Hussakof, 1908, p. 95); KUVP 498, partial skull; KUVP Notes–Synonyms of Saurodon leanus are: Saurocephalus phle- 499 (holotype of Martinichthys latus McClung, 1926), 500 (holotype of botomus Cope, 1870, Ichthyodectes goodeanus Cope, 1877b, S. M. alternatus McClung, 1926), 501 (holotype of M. intermedius McClung, goodeanus (Cope, 1877b), Daptinus broadheadi Stewart, 1898a (genus 1926), 502 (holotype of M. acutus McClung, 1926), 503 (paratype of M. Daptinus was erected by Cope, 1873), Saurodon xiphirostris Stewart, acutus McClung, 1926), 504 (holotype of M. gracilis McClung, 1926), 1898c, S. broadheadi (Stewart, 1898a) (e.g., see Loomis, 1900), S. ferox 506, and 507, rostral specimens (Taverne, 2000b; Everhart, 2005c, p. 93). Stewart, 1898c. Bardack (1965b, p. 9) noted FHSM VP-1650 as “Saurodon Notes–This species was initially assigned to the genus Erisichthe sp.” Cope, 1873, by Cope (1877c), and it was subsequently assigned to the Additional remarks (at family level)–Hussakof (1908, p. 98) re- genus Protosphyraena Leidy, 1857 (e.g., Loomis, 1900; Hay, 1903) until ferred the following three specimens to “Saurocephalid indet.”: AMNH FF McClung (1926) established the genus Martinichthys. Taverne (2000b) 1606, partial skull; AMNH FF 1607, partial skull; AMNH FF 1911, verte- synonymized the following species with Martinichthys ziphioides (Cope, brae. 1877c): M. acutus McClung, 1926, M. alternatus McClung, 1926, M. Order Tselfatiiformes Nelson, 1994 gracilis McClung, 1926, M. intermedius McClung, 1926, and M. latus McClung, 1926. Family Plethodidae Loomis, 1900 Additional remarks (at genus level)–Everhart and Everhart (1993) Genus Bananogmius Whitley, 1940 noted that the occurrences of Martinichthys specimens are restricted to the Bananogmius aratus (Cope, 1877a) lower one third of the Smoky Hill Chalk. Everhart and Everhart (1994) Referred material– AMNH FF 2403 (holotype), nearly complete reported a rostral specimen of Martinichthys from the Smoky Hill Chalk skeleton (Hay, 1903, p. 34–41; Hussakof, 1908, p. 61, fig. 30; Nelson, (unnumbered specimen; species unspecified) that shows bite marks of an 1973, fig. 8B; Taverne, 2000a, 2000d, 2001c). undetermined vertebrate. Besides FHSM VP-15567 (rostrum of M. brevis: see above), Everhart (2005c, p. 95, 96) noted additional 18 Martinichthys Bananogmius favirostris (Cope, 1877b) specimens in FHSM (VP-15549 – VP-15566, and VP-15568) of which Referred material–AMNH FF 2109 (paratype), partial skull; one of them (VP-15568) may belong to an undescribed taxon. AMNH FF 2111 (holotype), partial skull and vertebrae (Hay, 1903, p. 30– 201 Genus Pseudothryptodus Loomis, 1900 Pentanogmius. Bananogmius polymicrodus (e.g., see Fielitz and Shimada, Pseudothryptodus intermedius Loomis, 1900 1999), described as Beryx polymicrodus by Stewart (1898d), as Anogmius polymicrodus by Stewart (1899d) and Jordan (1924), and as Osmeroides Referred material–None (see “Notes” below). polymicrodus (Stewart, 1898d) by Loomis (1900), is considered to be a Notes–The only specimen of this taxon, the holotype, was destroyed junior synonym of this species (“B. evolutus” in Schultze et al., 1982). during the Second World War in Germany (Taverne, 2003). KUVP 308 represents the holotype of Stewart’s (1898d) Beryx maltidentatus, which was once referred to Osmeroides evolutus (Cope, Genus Thryptodus Loomis, 1900 1877b) by Loomis (1900). AMNH FF 8319 was described as a possible Thryptodus zitteli Loomis, 1900 specimen of Moorevillia hardi Applegate, 1970, by Applegate (1970, p. Referred material–AMNH FF 19557, rostrum; KUVP 456, 457, 416); however, the assimilation of this specimen into Pentanogmius and 459, rostral specimens (Taverne, 2000c); KUVP 25899, rostrum evolutus eliminates the Niobrara record of M. hardi. (Taverne, 2003; Everhart, 2005c, p. 96). Genus Zanclites Jordan, 1924 Notes–The holotype was destroyed during the Second World War in Germany (Shimada and Schumacher, 2003; Everhart, 2005c, p. 96, fig. Zanclites xenurus Jordan, 1924 5.8). Stewart (1900, p. 391, 392) and Hay (1903, p. 40, 41) suggested that Referred material–KUVP 52 (holotype), complete skeleton Thryptodus is a synonym of Anogmius Cope, 1877a, a junior homonym (Taverne, 1999; Everhart, 2005c, p. 97). replaced by Bananogmius Whitley, 1940. Recently, Taverne (2001c, 2003) Notes–The holotype is the only known specimen of this taxon. suggested that Thryptodus is a distinct genus and is not synonymous with Bananogmius. Everhart (2005c, p. 97) noted the occurrence of a Genus Dixonanogmius Taverne, 2000c Thryptodus specimen (FHSM VP-15571) that consists of “most of the Dixonanogmius sp. skull and several vertebrae.” Referred material–KUVP 299 and 463, fin fragments; KUVP 12008; dentary; KUVP 69373, fin fragment (Taverne, 2000c). Genus Plethodus Dixon, 1850 Notes–Taverne (2000c) established the genus Dixonanogmius by Plethodus rotundus (Loomis, 1900) designating Plethodus oblongus (Dixon, 1850) from England as its type Referred material–None (see “Notes” below). species (see also Cavin and Forey, 2001). Notes–The holotype (partial skull) described under the name Additional remarks (at family level)–Hay (1903, p. 44, figs. 33, Thryptodus rotundus by (Loomis, 1900) is the only known specimen of 34) and Hussakof (1908, p. 62) described AMNH FF 994 (lower dental this taxon, and it was destroyed during the Second World War in Germany. plate) and AMNH FF 2055 (caudal fin) as “Anogmius sp.” Part of the Taverne (2000c) placed this species in the genus Plethodus; however, the holotype of Ferrifrons rugosus Jordan, 1924 (KUVP 296) was found to taxonomic status of the genus Plethodus remains unresolved (see Cavin be a plethodid material (KUVP 68902; see Arratia and Chorn, 1998, p. and Forey, 2001). 301). Taverne (2000d) referred KUVP 57273 to as a tail of an undeter- mined plethodid. As noted by Taverne (2003, p. 170) and Taverne and Genus Pseudanogmius Taverne, 2002b Gayet (2005, p. 67, 83), there is an undescribed plethodid taxon in the Pseudanogmius maiseyi Taverne, 2002b Smoky Hill Chalk (FHSM VP-3248: see also Everhart, 2005c, p. 95, 96). Referred material– AMNH FF 1116 (paratype), partial skeleton Beeson and Shimada (2004) reported the occurrence of “Plethodidae” in a (as “AMNH 1616” in Hay, 1903, fig.15, and in Hussakof, 1908, p. 62); fossiliferous rock specimen catalogued as FHSM VP-644. AMNH FF 1977 (paratype), caudal skeleton; AMNH FF 8129 (holotype), Subdivision Elopomorpha partial skeleton (Taverne, 2002b; Everhart, 2005c, p. 97). Notes–The taxon was referred to “Anogmius sp.” by Hay (1903) Order Elopiformes Greenwood, Rosen, Weitzman, and Myers, 1966 and Hussakof (1908), and “Bananogmius sp.” by Nelson (1973, fig. 8C). Family Elopidae Jordan, 1923 Genus Palelops Applegate, 1970 Genus Pentanogmius Taverne, 2000c Palelops eutawensis Applegate, 1970 Pentanogmius evolutus (Cope, 1877b) Referred material–FMNH PF3336, scale (Applegate, 1970, p. Referred material–AMNH FF 2101 (holotype), jaw fragment 406). (Hussakof, 1908, p. 62; Hay, 1903, p. 46, 47; Nelson, 1973, figs. 2D, 5B); AMNH FF 2137, cranial elements and scapula; AMNH FF 3073, cranial Order Albuliformes Greenwood, 1977 elements; AMNH FF 3074, cranial elements and vertebrae; AMNH FF Family Albulidae Bleeker, 1859 7358, cranial elements; AMNH FF 8319, complete caudal fin (Nelson, 1973, figs. 5C, 5D, 6B; Taverne, 2004); BMNH P.10159, partial skeleton Genus and species undetermined (Taverne, 2000a); KUVP 2, complete skeleton; KUVP 6, ceratohyal; KUVP Referred material–FHSM VP-644, vomers and teeth (the cata- 11, glossohyal and parasphenoid; KUVP 14, mandibles and ceratohyal; logue number is assigned to an entire fossiliferous rock specimen which KUVP 16 (cf. Taverne, 2000a, 2000d), tail; KUVP 273, tail; KUVP 308, yielded the teeth: Beeson and Shimada, 2004; Fig. 2G). partial skull; KUVP 12083, partial skull and vertebrae (for review, see Notes–Applegate (1970, p. 413) noted that KUVP “has a specimen Schultze et al., 1982); KUVP 4, partial skull (as “B. polymicrodus”: Fielitz (F 985) from the Niobrara labeled Leptichthys which is an unquestionable and Shimada, 1999, p. 511); KUVP 283, mandible (Jordan, 1924, p. 221); albulid”; however, the catalogue number “F 985” does not exist in KUVP. KUVP 69016 and 69372, partial skull; BMNH P.9202, partial skeleton, BMNH P.9645, P.10158, and P.11147, cranial elements (Taverne, 2004); Order Anguilliformes Berg, 1940 BMNH P.9646, caudal fin; BMNH P.10610, complete skeleton (Taverne, Family Urenchelidae Woodward, 1900 2000d). Genus Urenchelys Woodward, 1900 Notes–This taxon was previously referred to Anogmius evolutus Urenchelys abditus Wiley and Stewart, 1981 and Bananogmius evolutus (for review, see Taverne, 2004). Taverne (2001c) suggested that species does not belong to the genus Bananogmius Referred material–KUVP 47241 (holotype), partial skeleton (Wiley and subsequently (Taverne, 2004) placed the species into the genus and Stewart, 1981). Notes–The holotype is on a valve of the inoceramid bivalve, 202 Platyceramus platinus (Stewart, 1990b, 1990c, 1996). scribed P. sheareri. Cope (1877b) later placed Oricandinus sheareri Cope, 1872, within Pachyrhizodus (spelled “Pachyrhizodus Sheari” in Loomis, Order Crossognathiformes Taverne, 1989 1900). Pachyrhizodus sheareri (Cope, 1877b) and O. tortus Cope, 1877b, Family Crossognathidae Woodward, 1901 are likely synonyms of P. minimus, but Applegate (1970) stated that the Genus Apsopelix Cope, 1871 preservation of specimens of these two species is too poor to be conclusive. Additional remarks (at genus level)–Jordan’s (1924) Kansanus Apsopelix anglicus (Dixon, 1850) martini (holotype: KUVP 128) is considered to be Pachyrhizodus sp. Referred material–FHSM 7958, complete skeleton; KUVP 309, (Schultze et al., 1982, p. 37). complete skeleton; KUVP 310, nearly complete skeleton lacking caudal fin; KUVP 318, head and pectoral girdle; KUVP 519, complete skeleton Order Aulopiformes Rosen, 1973 (Teller-Marshall and Bardack, 1978). Suborder Enchodontoidei sensu Nelson, 1994 Notes–Synonyms of Apsopelix anglicus are: A. sauriformis Cope, Superfamily Cimolichthyoidea Nelson, 1994 1871, Syllaemus latifrons Cope, 1875, Pelycorapis berycinus Cope, 1877a, Leptichthys agilis Stewart, 1899e, Helmintholepis vermiculatus Cockerell, Family Cimolichthyidae Goody, 1969 1919, Palaeoclupea dakotensis Dante, 1942, and S. hanifii Green, 1957. Genus Cimolichthys Leidy, 1857 Family Pachyrhizodontidae Cope, 1872 Cimolichthys nepaholica (Cope, 1872) Genus Pachyrhizodus Dixon, 1850 Referred material–AMNH FF 1736, fragmentary skeleton (Hussakof, 1908, p. 66); AMNH FF 1737 (holotype of Empo merrillii Pachyrhizodus caninus Cope, 1872 Cope, 1874), fragmentary palatine, ectopterygoid, and vomer; AMNH FF Referred material–AMNH FF 1658, fragmentary caudal fin 1738 (holotype of E. contracta Cope, 1874), palatines, vomer, and man- (Hussakof, 1908, p. 85); AMNH FF 1662, mandible (Hay, 1903; Nelson, dibles; AMNH FF 1739, partial skull; AMNH FF 1740, partial vertebral 1973, fig. 2C); AMNH FF 1758 (holotype of Pachyrhizodus latimentum column with pectoral and caudal fins; AMNH FF 1741, skull fragments; Cope, 1872), mandibles, hyomandibula, and interoperculum; AMNH FF AMNH FF 1742, partial skull (Hussakof, 1908, p. 67); AMNH FF 1751, 1881 (holotype), jaw elements; AMNH FF 1900, caudal fin; AMNH FF skull fragments and caudal fin (Goody, 1970); AMNH FF 1776, partial 1903, vertebrae (Hussakof, 1908, p. 85); AMNH FF 2041, vertebrae and skull with jaws (Hay, 1903); AMNH FF 1882 (holotype of Cimolichthys unpaired fin (Nelson, 1973, fig. 8A; Hay, 1903); FHSM VP-2189, skull sulcaus Cope, 1872), palatine, mandibles, and vertebrae; AMNH FF 1904 (Everhart, 2005c, p. 83); KUVP 164, premaxillae; KUVP 278, mandible, (holotype), fragmentary palatine (Hussakof, 1908, p. 66); AMNH FF 1967, maxilla, and glossohyal (Stewart, 1900); LACM 132705, partial maxilla skull fragments; AMNH FF 1969, neurocranium (Goody, 1970); AMNH (Schwimmer et al., 1997, p. 77). FF 1989 (holotype of C. seminaceps Cope, 1872), palatine, mandibles, Notes–Loomis (1900) discussed Pachyrhizodus caninus from the and vertebrae; AMNH FF 2032, caudal fin and vertebrae (Hussakof, 1908, Niobrara of Kansas. Pachyrhizodus latimentum Cope, 1872, and P. p. 67); AMNH FF 2369, caudal fin and vertebrae; AMNH FF 2522, neuro- curvatus Loomis, 1900, are synonyms of P. caninus (Hay, 1903). Hakel cranium and upper jaws; AMNH FF 3889, premaxilla; AMNH FF 6412, and Stewart (2003) reported a nearly complete skeleton of P. caninus from partial neurocranium; AMNH FF 8348, skull fragments and vertebrae; the Smoky Hill Chalk of Kansas. AMNH FF 8363, mandible (Goody, 1970); FHSM VP-644, teeth (the cata- logue number is assigned to an entire fossiliferous rock specimen which Pachyrhizodus kingi Cope, 1872 yielded the teeth: Beeson and Shimada, 2004); KUVP 62, disarticulated Referred material–AMNH FF 8386 (holotype), fragmentary max- skull and vertebrae; KUVP 70, jaws, ceratohyal, and ethmoid; KUVP 71, illa (Applegate, 1970, p. 411); KUVP 75 (holotype of Pachyrhizodus jaws and ceratohyal (Stewart, 1900); KUVP 59032, pectoral fin, verte- leptognathus Stewart, 1898d), cranial elements; KUVP 316 (holotype of brae, and intestinal cast (Moodie, 1911). P. velox Stewart, 1898d), cranial elements (Schultze et al., 1982). Notes–Synonyms of Cimolichthys nepaholica are: C. seminaceps Notes ¯ Pachyrhizodus leptognathus from the Niobrara Chalk of Cope, 1872, C. sulcatus Cope, 1872, Empo nepaholica Cope, 1872, E. Kansas was discussed by Loomis (1900). Applegate (1970) considered sulcata Cope, 1874, E. semianceps Cope, 1874, E. contracta Cope, 1874, Stewart’s (1898d) P. leptognathus and P. velox to be synonyms of P. kingi. E. merrillii Cope, 1874, and E. lisbonensis Stewart, 1899c. Loomis’s (1900) Pachyrhizodus leptopsis Cope, 1874 “C. nepaeolica (Cope, 1872)” is a probable misspelling of C. nepaholica. Cope (1872), Stewart (1900), and Hay (1903) recognized Empo as a sepa- Referred material–AMNH FF 1756 (holotype), fragmentary rate genus. Goody (1970) stated that the genus Empo is synonymous with dentary (Hay, 1903, p. 64; Hussakof, 1908, p. 85); KUVP 151, partial Cimolichthys. Bardack (1965b, p. 9) noted FHSM VP-1649 and VP-5014 mandible (Stewart, 1900). as “Cimolichthys sp.” Everhart (2005c, p. 87) noted a “Cimolichthys speci- Notes–Loomis (1900) and Stewart (1990a) discussed men” (FHSM VP-15065) that contained “a large Enchodus” (FHSM VP- Pachyrhizodus leptopsis from the Niobrara of Kansas. Applegate (1970) 15066) and an unidentified fish (FHSM VP-15067). Everhart (2005c, p. considered P. leptopsis to be a synonym of P. caninus; however, it is still 87) also noted another Cimolichthys specimen (FHSM VP-14024) occurred regarded as a separate species (Stewart, 1990a; Stewart and Bell, 1994; with an E. petrosus specimen (FHSM VP-14025). Everhart, 2005c). Family Dercetidae Pictet, 1850 Pachyrhizodus minimus Stewart, 1899b Genus Anguillavus Hay, 1903 Referred material–AMNH FF 1998 (holotype of Oricardinus sheareri Cope, 1872), partial maxilla; AMNH FF 2114 (holotype of O. Anguillavus hackberryensis Martin, 1922 tortus Cope, 1877b), partial dentary and vertebrae; CMC VP-7552, com- Referred material–KUVP 927 (holotype), complete skeleton (Mar- plete skeleton (Everhart, 2005c, p. 83, 84); FHSM VP-326 (Miller, 1957; tin, 1922). Everhart, 2005c, p. 83); FHSM VP-644, teeth (the catalogue number is Notes–This taxon was initially described as an anguilliform fish, assigned to an entire fossiliferous rock specimen which yielded the teeth: but Wiley and Stewart (1981) placed it within Dercetidae. Beeson and Shimada, 2004); KUVP 249 (holotype of Eurychir lindleyi Genus Leptecodon Williston, 1899 Jordan, 1924), skull, pectoral girdle, and vertebrae; KUVP 327 (holotype), mandibles (Stewart, 1899b; Everhart, 2005c, p. 83). Leptecodon rectus Williston, 1899 Notes–Eurychir lindleyi Jordan, 1924, is considered to be Referred material–KUVP 35 (holotype), two individuals (Bardack, Pachyrhizodus minimus (Schultze et al., 1982, p. 36). Cope (1872) de- 203 1976, p. 371; Stewart, 1990b, fig. 5). amicrodus Stewart, 1898d), dentary (Goody, 1976); KUVP 379, disar- Notes–Stewart (1990b, p. 36) noted that only two specimens of ticulated skull; KUVP 800, palatine; KUVP 802, palatine; KUVP 803, Leptecodon are known (holotype and one unspecified specimen) and both jaws; KUVP 804, disarticulated skull; KUVP 805, disarticulated cranium; are on valves of the inoceramid bivalve, Platyceramus platinus (see also KUVP 806, caudal fin; KUVP 807, complete skeleton; KUVP 833, pa- Schultze et al., 1982). At least one of them contains Omosoma bones as latine tooth; KUVP 836, cranium; KUVP 837, palatine; KUVP 839, man- stomach contents (Stewart, 1990b, p. 36, 1990c, p. 57, 1996, p. 390). dible (Green, 1913). Notes–Synonyms of this species include: Enchodus amicrodus Genus Stratodus Cope, 1872 Stewart, 1898d, E. saevus Hay, 1903 (according to Goody, 1976), and Stratodus apicalis Cope, 1872 Tetheodus pephredo Cope, 1874 (according to Hay, 1903). Everhart Referred material–AMNH FF 1753 (holotype), partial skull (2005c, fig. 5.3) illustrated three jaw fragments of E. petrosus (unnum- (Hussakof, 1908, p. 102); KUVP 23, skull with jaws (Stewart, 1900); bered specimens). KUVP 21 is attributed to E. petrosus by Green (1913). KUVP 333, partial skeleton (Wiley and Stewart, 1981). Enchodus shumardi Leidy, 1856 Notes–Churcher (1991) noted the occurrence of this species from the Niobrara Chalk of Kansas (specimens unspecified). Beeson and Shimada Referred material–FHSM VP-644, teeth (the catalogue number is (2004) reported this species in a fossiliferous rock specimen curated as assigned to an entire fossiliferous rock specimen which yielded the teeth: FHSM VP-644, but subsequent examination suggests that the material iden- Beeson and Shimada, 2004); KUVP 321 (neotype of Enchodus parvis tified as Stratodus apicalis is not of the taxon (K. Shimada, unpubl. 2006). Stewart, 1898d), dentary; KUVP 323, dentary; KUVP 377, dentary, pala- tal bones, and preopercle; KUVP 823, opercular bones; KUVP 824, dentary; Stratodus oxypogen Cope, 1877b KUVP 825, dentary; KUVP 826, dentary; KUVP 827, skull with jaws; Referred material– AMNH FF 2113 (holotype), jaws, palatine, KUVP 828, palatine tooth; KUVP 829, skull; KUVP 838, palatine tooth and vertebrae (Cope, 1877b; see also Hay, 1903). (Green, 1913). Notes–Enchodus parvis Stewart, 1898d, is a synonym of E. Superfamily Enchodontoidea Nelson, 1994 shumardi (Loomis, 1900; Green, 1913). Family Enchodontidae Woodward, 1901 Additional remarks (at genus level)–Fishman, Shimada et al. (1995) and Fishman, Fielitz et al. (1995) referred to FHSM VP-11062 and Genus Enchodus Agassiz, 1835 KUVP 809, respectively, as Enchodus from the Smoky Hill Chalk of Kan- Enchodus dirus (Leidy, 1857) sas, but they did not specify the species. Everhart (2005c, p. 87) discov- Referred material–KUVP 322, dentary; KUVP 516, palatine; ered “a large Enchodus” (FHSM VP-15066) inside a “Cimolichthys speci- KUVP 817, skull (Green, 1913). men” (FHSM VP-15065) along with an unidentified fish (FHSM VP- Notes–Leidy (1857) described this species as Phasganodus dirus, 15067). but Cope (1875) showed that it belongs to Enchodus. Additional remarks (at order level)–Everhart (2005c, p. 99, tables 5.1, 13.1) noted the occurrence of Apateodus Woodward, 1901 (super- Enchodus gladiolus (Cope, 1872) family Ichthyotingoidea Nelson, 1994: family incertae sedis) in the Smoky Referred material–AMNH FF 1685, partial skull; AMNH FF 1818, Hill Chalk of Kansas (specimens unspecified; see also Stewart, 1990a, p. partial skull (Hay, 1903); AMNH FF 1820 (holotype of Enchodus dolichus 25, 30). Cope, 1875), palatine; AMNH FF 1837, skull; AMNH FF 1865, partial Order Polymixiiformes Rosen and Patterson, 1969 skull (Hussakof, 1908, p. 71, 72); FHSM VP-644, teeth (the catalogue number is assigned to an entire fossiliferous rock specimen which yielded Family Polymixiidae Gill, 1862 the teeth: Beeson and Shimada, 2004); KUVP 174, disarticulated skull; Genus Omosoma Costa, 1857 KUVP 808, partial skull; KUVP 812, hyopalatines (Fielitz, 2002); KUVP Omosoma garretti Bardack, 1976 814, skull and pectoral fins; KUVP 816, upper jaws and mandible; KUVP 819, disarticulated skull; KUVP 820, disarticulated skull (Green, 1913); Referred material–AMNH FF 9837, 9838, and 9840, multiple in- KUVP 821, dentary (Stewart, 1900); KUVP 831, disarticulated skull; dividuals (Bardack, 1976; Stewart, 1990c); FHSM VP-5115, multiple in- KUVP 832, mandible, palatines, and premaxilla (Green, 1913). dividuals (Stewart, 1990b); FMNH PF3040 (holotype; as “FMNH Notes–Cope (1872) designated Enchodus gladiolus as a new spe- PF43040” in Stewart, 1990b, p. 37), individual (Bardack, 1976); KUVP cies of Cimolichthys. Cimolichthys gladiolus Cope, 1872, was based on 32398, 47 individuals; KUVP 59016, 59017, and 65738, multiple indi- an unnumbered specimen at the AMNH that was lost and has never been viduals (Stewart, 1990b, 1990c). recovered (Hay, 1903). Enchodus dolichus Cope, 1875 (e.g., see Loomis, Notes–Almost all Omosoma garretti specimens are found on valves 1900), is a synonym of E. gladiolus (Green, 1913). The name E. dolichus of the inoceramid bivalve, Platyceramus platinus (Bardack, 1976; Stewart, continued to be used (Hay, 1903; McNulty and Kienzlen, 1970) until Goody 1996). Partial individuals of this species have been found associated with (1976) reaffirmed as a synonym. and possibly inside the holotype of Leptocodon rectus as stomach contents (Stewart 1990b, p. 36, fig. 5, 1990c, p. 57, 1996, p. 390). Enchodus petrosus Cope, 1874 Superorder Acanthopterygii Rosen and Patterson, 1969 Referred material–AMNH FF 198 (holotype of Enchodus saevus Hay, 1903), disarticulated skull; AMNH FF 1605 (holotype of Tetheodus Order Beryciformes Berg, 1940 pephredo Cope, 1874), palatines and part of dentary; AMNH FF 1608 Family Holocentridae Richardson, 1846 (holotype), neurocranium and palatine; AMNH FF 1668, partial skull; Genus Kansius Hussakof, 1929 AMNH FF 2062, skull fragments (Hay, 1903; Hussakof, 1908, p. 73); Kansius sternbergi Hussakof, 1929 AMNH FF 2098, palatines, jaws, and suspensorium; AMNH FF 2098, partial skull and vertebrae; AMNH FF 8139, palatines; BMNH P9647, Referred material–FHSM VP-25 (holotype), impression of two skull (Goody, 1976); FHSM VP-644, teeth (the catalogue number is as- individuals (Hussakof, 1929; Bardack, 1965b); FHSM VP-8123, ca. 50 signed to an entire fossiliferous rock specimen which yielded the teeth: individuals; KUVP 345, multiple individuals (Bardack, 1976). Beeson and Shimada, 2004); FHSM VP-2939, skull; FHSM VP-14025, Notes–Almost all Kansius sternbergi specimens are found on valves partial skeleton (Everhart, 2005c, p. 85, 87); KUVP 51, mandible and of the inoceramid bivalve, Platyceramus platinus (Bardack, 1976; Stewart, opercle; KUVP 126, palatines (Green, 1913); KUVP 324 (holotype of E. 1996). 204 Genera and species undetermined catalogue numbers (e.g., many specimens discussed in “Notes” and “Ad- Referred material–AMNH FF 9840, multiple individuals (Stewart, ditional remarks”) nor does it include specimens that are catalogued in 1990b, 1990c); FHSM VP-2161, individual (Stewart, 1996, p.390; prob- museums but have never been documented in published papers. It does, ably the same material as “SMM 11543” in Bardack, 1976, figs. 7–8); however, include the only known specimens (holotypes) of KUVP 27241, multiple individuals?; KUVP 47245, 28 individuals; KUVP Pseudothryptodus intermedius and Plethodus rotundus that were destroyed 47247, multiple individuals?; KUVP 49403, 104 individuals, KUVP 57251, during the Second World War. Of the 643 catalogue numbers, 133 belong 65697, 65700, 69467–69483, and 72413–72415, multiple individuals; to chondrichthyans and 510 to osteichthyans. KUVP 82576, individual; LACM 132681, individual; LACM 131596, The 643 described paleoichthyological specimens support the oc- individual; USNM 336382–336386, multiple individuals (Stewart, 1990b, currence of a minimum of 70 fish species from the Smoky Hill Chalk of 1990c, 1996, p. 390). Kansas, comprising at least 16 chondrichthyans and 54 osteichthyans (Table Notes–All holocentrid specimens are found on valves of the 1). Of these 70 species, 44 are represented by type specimens from the inoceramid bivalve, Platyceramus platinus (Stewart, 1996). Stewart (1996) Smoky Hill Chalk. These counts do not include undescribed taxa, such as noted that FHSM VP-2161 and KUVP 82576 may be conspecific. a number of Stewart’s (1982, 1996, p. 390) holocentrid taxa (including Additional remarks (at family level)–Although we cite only one Caproberyx sp. and Trachichthyoides sp.), Everhart’s (2005c, tables 5.1, taxon of undetermined genus and species of Holocentridae, Stewart (1990a, 13.1) Belonostomus sp. and Apateodus sp., and Shimada’s (unpublished 1990b, 1990c, 1996) noted the presence of four undescribed genera of the data referred to in Taverne, 2003, p. 170) plethodid taxon. Once these taxa family Holocentridae. Stewart (1996, p. 390) noted the occurrence of are formally described, the total taxonomic count of non-tetrapod verte- Caproberyx Regan, 1911, and Trachichthyoides Woodward, 1902 (un- brates would increase if there is no deletion of species from the Smoky Hill specified specimens), on valves of the inoceramid bivalve, Platyceramus Chalk and no synonymization of listed taxa. platinus, from the Smoky Hill Chalk of Kansas. In his list of Smoky Hill Chalk vertebrates, Russell (1988, 1993) listed 64 non-tetrapod vertebrate taxa, not counting species of Stewart’s Order incertae sedis (within Acanthomorpha Rosen, 1973) (1982) four undescribed holocentrid genera. Our list includes a number of Family Ferrifronsidae Arratia and Chorn, 1998 species not in Russell’s list. For example, the number of known Squalicorax species from the Smoky Hill Chalk is doubled by the addition of S. Genus Ferrifrons Jordan, 1924 pristodontus and S. volgensis. However, the increased number of taxa in Ferrifrons rugosus Jordan, 1924 this paper is not due to a simple addition of previously unreported species. Referred material–KUVP 296 (holotype), anterior half of skel- In some instances, the number of species decreased in certain groups. For eton (Schultze et al., 1982). example, Russell (1988) listed six species of Ptychodus, but we removed Notes–This taxon is considered to be an acanthomorph fish (Arratia two species (P. mammillaris and P. occidentalis) as not occurring in the and Chorn, 1998). Part of the holotype (KUVP 296) was found to be a Smoky Hill Chalk and added a species (P. cf. P. latissimus). Russell re- non-Ferrifrons element, but plethodid material (KUVP 68902; see Arratia corded 16 tselfatiiform species, including seven Martinichthys species. and Chorn, 1998, p. 301). Recent studies of this fish group by Taverne (2000b, 2000c, 2002b) have resulted in the reduction of Martinichthys species to two; however, some Order incertae sedis previously unreported and new taxa are added to our list (e.g., Plethodus, Family incertae sedis Pseudanogmius, and Dixonanogmius), which makes the total count of Genus Laminospondylus Springer, 1957 Tselfatiiformes from the Smoky Hill Chalk of Kansas 13 (for summary, see Taverne and Gayet, 2005). Thus, it should be emphasized that the taxo- Laminospondylus transversus Springer, 1957 nomic composition of certain groups listed by Russell (1988) is quite dif- Referred material–KUVP 312, complete fish (Jordan, 1924, pl. ferent from that in our list. 21; for review, see Schultze et al., 1982). It is noteworthy that the occurrences of 12 of the 16 chondrichthyan Notes–KUVP 312 was initially described as Leptichthys agilis by taxa and 32 of the 54 osteichthyan taxa in the Smoky Hill Chalk are sup- Stewart, 1900, and also followed by Jordan (1924). ported by fewer than five specimens, and 5 chondrichthyan taxa and 21 Order incertae sedis osteichthyan taxa are represented by a single specimen each (Table 1). Sev- eral of these ‘single-representative taxa’ (by number of catalogued speci- Family incertae sedis mens) are newly recognized taxa from the Smoky Hill Chalk, such as Genus Aethocephalichthys Fielitz, Stewart, and Wiffen, 1999 Johnlongia sp., Squalicorax volgensis, Rhinobatos sp., and Aethocephalichthys hyainarhinos Fielitz, Stewart, and Wiffen, 1999 Palaeobalistum sp. It should be noted that these recently added taxa are represented by small (>10 mm) specimens (Fig. 2) collected as a result of Referred material–KUVP 84901, neurocranium (Fielitz et al., paying particular attention to minimizing the collecting bias in the Niobrara 1999). fossil record (e.g., see Hamm and Shimada, 2002b). Subclass Sarcopterygii Bottjer (2002) characterized the Smoky Hill Chalk of Kansas as a Order Coelacanthiformes Lund and Lund, 1985 fossil Lagerstätte. However, our data suggest that the Niobrara fossil record is manifested with complex multiple effects of preservation, collecting, and Family, genus, and species undetermined research biases. Our personal experiences in the field and in museum col- Referred material–LACM 131958, partial cranium (Stewart et al., lections suggest that, relative to remains of Cretoxyrhina, Squalicorax 1991; Everhart, Everhart, and Stewart, 1995; Everhart, 2005c, p. 101; falcatus, Cimolichthys, and Enchodus petrosus for example, the occur- Fig. 2H). rences of the following fish taxa are simply rare in the Smoky Hill Chalk: Edaphodon, Hadrodus, Lepisosteus, Asarotus, Plethodus, DISCUSSION Pseudothryptodus, Anguillavus, Leptecodon, and coelacanth. Some other Our study shows that the sum of paleoichthyological specimens rep- fishes, such as Omosoma and holocentrids, are also not so common; how- resenting whole or parts of fishes from the Smoky Hill Chalk documented ever, when they are found, these fishes often come in groups, as many as in literature is 643. This represents the sum of all the numbers in parenthe- tens of complete or nearly complete, conspecific individuals preserved to- ses in Table 1 with FHSM VP-644 included separately for each taxon. This gether on inoceramid bivalve shells. Their unique mode of preservation figure includes neither specimens discussed in literature without referable and the well-preserved nature of the specimens have attracted some de- tailed examinations of these fishes (e.g., Bardack, 1976; Stewart, 1990b, 205

TABLE 1. List of fish taxa from Smoky Hill Chalk of Kansas reported to date. Number in parentheses indicates total specimen count by referable catalogue numbers documented in literature (i.e., sum of specimens listed in “Referred material” for each taxon). Single asterisk (*) denotes count that includes FHSM VP-644, catalogue number assigned to fossiliferous rock specimen that contained the taxon (Beeson and Shimada, 2004). Double asterisk (**) indicates taxon with no representative specimen (i.e., the only known specimen for the taxon is lost).

Chondrichthyes Osteichthyes

Edaphodon sp. (1) Micropycnodon kansasensis (2) Pentanogmius evolutus (25) Ptychodus anonymus (2) Hadrodus marshi (1) Zanclites xenurus (1) Ptychodus cf. P. latissimus (1) Palaeobalistum sp. (1*) Dixonanogmius sp. (4) Ptychodus martini (2) Lepisosteus sp. (1) Palelops eutawensis (1) Ptychodus mortoni (4) Protosphyraena gladius (10) Albulidae gen. et sp. indet. (1*) Ptychodus polygyrus (4) Protosphyraena nitida (22) Urenchelys abditus (1) Scapanorhynchus raphiodon (2) Protosphyraena perniciosa (50) Apsopelix anglicus (5) Johnlongia sp. (1) Protosphyraena tenuis (24) Pachyrhizodus caninus (11) Cretalamna appendiculata (3) Paraliodesmus guadagnii (2) Pachyrhizodus kingi (3) Cretoxyrhina mantelli (47’) Asarotus arcanus (1) Pachyrhizodus leptopsis (2) Pseudocorax laevis (2) Gillicus arcuatus (30) Pachyrhizodus minimus (7*) Squalicorax falcatus (32*) Ichthyodectes ctenodon (35*) Cimolichthys nepaholica (26*) Squalicorax kaupi (28) Xiphactinus audax (64) Anguillavus hackberryensis (1) Squalicorax pristodontus (2) Prosaurodon pygmaeus (7) Leptecodon rectus (1) Squalicorax volgensis (1*) Saurocephalus lanciformis (6) Stratodus apicalis (3) Rhinobatos sp. (1*) Saurodon leanus (22) Stratodus oxypogen (1) Bananogmius aratus (1) Enchodus dirus (3) Bananogmius favirostris (2) Enchodus gladiolus (16*) Luxilites striolatus (1) Enchodus petrosus (26*) Syntegmodus altus (1) Enchodus shumardi (12*) Niobrara encarsia (1) Omosoma garretti (9) Martinichthys brevis (3) Kansius sternbergi (3) Martinichthys ziphioides (10) Holocentridae gen. et sp. indet. (37) Pseudothryptodus intermedius (1**) Ferrifrons rugosus (1) Thryptodus zitteli (5) Laminospondylus transverses (1) Plethodus rotundus (1**) Aethocephalichthys hyainarhinos (1) Pseudanogmius maiseyi (3) Coelacanthiformes (1)

1990c). In contrast, Protosphyraena and Martinichthys are generally rep- example, an assemblage of individuals of the same fish species may occur resented only by pectoral fins and rostra, respectively, possibly because in close proximity (e.g., holocentrid taxa on inoceramid valves: Stewart, other parts of their skeletons were poorly ossified in life. Even though these 1990b, 1990c), and the entire assemblage, or part of the assemblage, may remains are generally incomplete and occur sparsely isolated in the chalk be given a single catalogue number. In addition, Bardack (1965a) docu- beds, they are taxonomically easy to diagnose and show paleoecologic and mented the fact that a number of ichthyodectiform specimens are actually biostratigraphic significance (e.g., Stewart, 1988; Everhart and Everhart, composite skeletons (i.e., a reconstructed skeleton consisting of more than 1993, 1994). Another example of mixed effects of preservation, collecting, one individual fish). Nevertheless, our field experience suggests that fish and research biases is the case with Cretoxyrhina mantelli. Fourteen of the fossils are far more common than tetrapod fossils in the Smoky Hill Chalk 47 referred specimens in literature are represented by well-studied nearly of Kansas. Our intuition is that there is a collecting bias towards large tetra- complete skeletons, skeletal parts, or associated teeth (e.g., Shimada, 1997c, pods, such as mosasaurs, in the Niobrara fossil record, and that the propor- 1997d). However, our field experience suggests that isolated teeth of the tion of fish specimens catalogued in museum collections estimated by shark are far more common than associated teeth and skeletal remains. Russell (1988: i.e., 57%) appears far smaller than the actual proportion of Because various biases are apparently operating on each fish taxon at dif- fishes to tetrapods in the field. This is supported by Everhart’s (2005c) ferent levels, characterizing the Niobrara fauna as a fossil Lagerstätte may account of field collecting. Based on his field notes, fish specimens make be an overstatement. up 78% of all specimens collected in a six year period. Russell (1988) estimated that about 7,416 vertebrate specimens had The Smoky Hill Chalk was deposited over about five million years been recovered from the “Niobrara Chalk” of which approximately 57% (Obradovich and Cobban, 1975, p. 50), but it must be noted that the (4,222 specimens) are non-tetrapod vertebrates. Whereas our data do not “Niobrara fauna” is a chronologically mixed (= time averaged) faunal as- include tetrapod species, we note that counting the number of specimens semblage (Stewart, 1990a; for time-averaging, see Kidwell and for each species does not reflect the abundance of any given taxon. Some Behrensmeyer, 1993), because fossils have been collected from various chondrichthyans are represented by a complete skeleton, associated tooth horizons within the 180-m-thick stratigraphic member. In other words, not set, or a set of skeletal parts from an individual (e.g., Shimada, 1997c, all listed species coexisted at any given point of time during the deposition 1997d), but the majority of cataloged specimens are isolated teeth. In some of the Smoky Hill Chalk. For example, Bardack (1965b) noticed that the cases, one tooth is given a unique catalogue number, whereas other teeth occurrences of Ptychodus and Protosphyraena are restricted to the lower are curated in groups with ‘lot’ numbers corresponding to multiple samples part of the stratigraphic member, and Stewart (1988) demonstrated that of the same taxon. This situation is not restricted to chondrichthyans. For even different species of Protosphyraena have different stratigraphic ranges. 206 The first peak in research activity was the decade between 1870 and 1879. The published research at this time period were Cope’s (1870, 1871, 1872, 1873, 1874, 1875, 1877a, 1877b, 1877c, 1878) descriptions of the major- ity of osteichthyan taxa (Fig. 4). Curiously, no work on the Smoky Hill Chalk fauna seemed to take place during the 1880s with regard to fishes. The second major period of research activity was between the late 1890s and into the early 1900s (Fig. 4). This period was probably the most productive in the shortest amount of time. Cope’s interest in fishes largely vanished by 1880 based on his publication record (e.g., see “References” below), but new workers such as O. P. Hay, F. B. Loomis, A. Stewart, and S. W. Williston resumed taxonomic work on the fish fauna. In addition to the description of new species, much of this work was a re-examination of Cope’s descriptions of the 1870s. Taxonomic research continued from 1910 until early 1960s, although to a lesser degree than in the past. There was a small peak in the 1920s reflecting an increase in new species descriptions from the Smoky Hill Chalk (e.g., by Jordan, 1924). This is also the time frame when G. F. Sternberg, a prominent fossil hunter (Fig. 1), began to collect fossils from the Niobrara Chalk of Kansas extensively (see Rogers, 1991). The third major peak in research activity began in about the mid- 1960s, but accelerated in the 1970s and is still on the increase today (Fig. 4). This recent resurgence in the study of fish taxa of the Smoky Hill Chalk is likely a reflection of the renewed interest in systematic ichthyology brought on by seminal papers such as Greenwood et al. (1966) and the first Interre- lationships of Fishes (Greenwood et al., eds., 1973), as well as the training of new ichthyologists from such institutions as the AMNH and BMNH. Much of the work on the osteichthyans has been centered on re-descrip- tion of the taxonomic groups and placing them in a phylogenetic context (e.g., Taverne, 1989, 1999, 2000a, 2000b, 2000c, 2001a, 2001b, 2001c, 2002a, 2002b, 2003). Additionally, there has been more work related to biostratigraphy and paleoecology of the Smoky Hill Chalk fishes (e.g., Stewart, 1990a, 1990b, 1999b; Everhart, Everhart, and Shimada, 1995; Everhart, Everhart, and Stewart, 1995; Shimada, 1997e; Everhart, 2005d). FIGURE 3. Graphs showing history of studies of fossil fishes from Smoky Hill A significant trend in the current explosion of taxonomic research in the Chalk of Kansas. A, number of new species described from 1870 to 2005; B, number Smoky Hill Chalk is an increase in the study of chondrichthyans, (e.g., of articles published from 1870 to 2005. Shimada, 1997c, 1997d; Hamm, 2001; Everhart and Caggiano, 2004; However, biostratigraphic studies of Niobrara vertebrates are difficult. Part Shimada and Cicimurri, 2005). Little work was done on this group after of the reason is due to the fact that the stratigraphy of the Smoky Hill Chalk Williston (1900a, 1900b). This renewed interest may be due to a rise in was poorly known until Hattin (1982) published on the Smoky Hill Chalk collecting in the Smoky Hill Chalk during the past several decades, and an geology. Therefore, before 1982, fossil collectors were unsure about the increasing appreciation of the diversity in the chondrichthyan taxa. It can stratigraphic level of fossils they recovered. To resolve this problem, Bennett also be attributed to the work of Hattin (1982) and Stewart (1990a) that (1990, 2000) showed that the stratigraphic level of any given specimen allowed specimens to be described in a stratigraphic and biostratigraphic may be inferred if adequate locality information accompanies the speci- context. men. Unfortunately, it is not uncommon for Niobrara vertebrate specimens CONCLUSION to retain the locality description only as “Kansas Chalk” or at best “western Kansas” (Bardack, 1965b; C. Fielitz and K. Shimada, personal observa- The Niobrara Chalk of Kansas is said to be “the most-diverse and tions), especially those collected in the late 1800s and early 1900s (e.g., see best-known Mesozoic fish assemblage in North America” (Wilson and a number of ichthyodectiform specimens cited in Bardack, 1965a). To com- Bruner, 2004, p. 583). Our study shows that, although the ichthyofauna plicate the situation, towns or counties used to describe fossil localities could constitutes a time-averaged assemblage, there are minimally 70 fish spe- have geographically shifted in the past (Bardack, 1965b). Furthermore, cies, comprising at least 16 chondrichthyans and 54 osteichthyans, cur- locality information may be omitted in literature depending on the focus of rently known from the Smoky Hill Chalk of Kansas. It is encouraging that, study, (e.g., Cretoxyrhina specimens in Shimada, 1997c), and not all local- in recent years, there apparently is renewed interest in systematic ichthyol- ity and/or stratigraphic data in literature are reliable or reproducible (e.g., ogy, biostratigraphy, and paleoecology, concerning Niobrara fishes. How- type locality of Micropycnodon kansasensis by Hibbard and Graffham, ever, much work on vertebrate biostratigraphy within the Niobrara Chalk 1941: see Stewart, 1990a). is still needed. It is our belief that progress in future studies of Niobrara A detailed discussion of the exploration and collecting history of fauna depends particularly on two factors. First, collecting bias must be Smoky Hill Chalk fossils is provided by Everhart (2005c). Our historical minimized, and second, a ‘good’ record of stratigraphic and locality data examination suggests that the number of new species descriptions from must be kept for each collected specimen. Whereas collecting every re- the Smoky Hill Chalk has substantially declined since the end of the nine- main of fossil fishes (including searching and recovering microscopic speci- teenth century and early twentieth century (Fig. 3A). It is intriguing, how- mens) and identifying the exact stratigraphic horizon and geographic posi- ever, that the amount of research in fish taxa and related biostratigraphy tion in the field may not be easy, such practices would contribute to eluci- has constantly increased for the last three decades (Fig. 3B). dating a more accurate picture of the paleoecology and evolutionary his- Our further historical review of literature (Fig. 4) suggests that there tory of the Niobrara fauna. were three major periods of Smoky Hill Chalk paleoichthyoloical research. 207

FIGURE 4. Graph showing number of times species were cited per decade of research.

ACKNOWLEDGMENTS It must be noted that the authors assume the responsibility of the content of this paper, not the reviewers. We thank the following individuals for providing us with specimen Both authors contributed in equal parts to this work. We did our data: J. Chorn (KUVP), I. Rutzky (AMNH), M. J. Everhart, R. J. best to provide an exhaustive entrée to the literature on Niobrara fishes; Zakrzewski (FHSM). We also thank G. D. Walters (FHSM) for providing however, if any relevant papers are inadvertently omitted, we would like to the image used for Figure 1, M. J. Everhart for Figures 2A and 2H, and S. learn about such publications. Compilation of publications cited in this work A. Hamm for Figure 2C. We are indebted to D. Bardack and M. J. Everhart was completed in December of 2005. for reviewing the manuscript and for providing us with useful information.

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