Thin Section Microscopy of the Fossil Fish Cylindracanthus
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130 Proceedings of the South Dakota Academy of Science, Vol. 96 (2017) THIN SECTION MICROSCOPY OF THE FOSSIL FISH CYLINDRACANTHUS Barbara S. Grandstaff1, Rodrigo A. Pellegrini2, David C. Parris2*, and Donald Clements2 1School of Veterinary Medicine University of Pennsylvania 3800 Spruce Street Philadelphia, PA 19104-6045 2New Jersey State Museum 205 West State Street PO Box 530 Trenton, NJ 08625-0530 *corresponding author email: [email protected] ABSTRACT Since discovery of the definitive specimen of Cylindracanthus in the Verendrye Formation of Hyde County, South Dakota, new interest has been focused this genus, known only from its distinctive rostral spines. Hypothetically linked to ascip- enseriform (sturgeon-like) fishes, the spines, found in circum-Atlantic Cretaceous to Eocene marine formations, have been examined in thin sections for more than a century. We have expanded this range of microscopy to include detailed descrip- tion of the tooth pedicels, examination of possible lesions and healed bony tissue, and petrography of the fossil, and probable functional anatomy. We have repeated the historical comparative studies with modern billfish specimens. Billfish rostral structure has no real resemblance (structural or mineralogical) to Cylindracanthus. The tooth base attachments in billfish are subdermal. In contrast, the tooth base structures and the lesions and damaged surfaces in Cylindracanthus give no evidence of having been subdermal. We have also expanded our comparative histological research to include Polyodon (paddlefishes) and Acipenser (sturgeons). Although fish teeth are typically acrodont (i. e., fused to the oral surfaces of the jaw and palatal elements), the teeth of Cylindracanthus are functionally pleur- odont, attached along (in fact, within) a groove, from which they protrude only slightly. The teeth point backwards toward the mouth (away from the tapered end of the rostrum). Hypothetically, the teeth could function in predation by causing damage during a stabbing motion of the rostral spine; they would be useless in a striking and/or slapping motion. The teeth would be highly effective in head-on attacks of shelled cephalopods, such as ammonites. This would have been advan- tageous during the Cretaceous Period but of no value later during the Eocene, after the ammonites had become extinct. We conjecture that the teeth became vestigial after the K-Pg event, as they are unknown in Cenozoic specimens. Keywords Cylindracanthus, Cretaceous, Eocene, histology Proceedings of the South Dakota Academy of Science, Vol. 96 (2017) 131 INTRODUCTION Agassiz (1833-1843) used the name Coelorhynchus for fossils which he identi- fied as probable rostral elements. Leidy (1857a) independently gave the name Cylindracanthus to fossils from New Jersey and Alabama which he interpreted as dorsal fin spines. He later (1857b) noted that his Cylindracanthus fossils resem- bled figures of Coelorhynchus published by Dixon (1850). Leriche (1905) pro- posed the name Glyptorhynchus to replace Coelorhynchus because Coelorhynchus was preoccupied by a chimaeroid, and noted that Glyptorhynchus is similar to the rostrum of Blochius. Glyptorhynchus is now considered a junior synonym of Cylindracanthus (Purdy et al. 2001). Smith-Woodward (1888, 1891) believed that Cylindracanthus (Coelorhynchus) was a fin spine. Histologic studies by Williamson (1849) and Carter (1927) did not clearly identify these fossils as either fin spines or rostra. Thus both anatomic and taxonomic interpretations of Cylindracanthus remain subject to question (Friedman 2012). Our understanding of Cylindracanthus has advanced greatly in recent years. A specimen from South Dakota (SDSM 30638) was instrumental in confirming that this enigmatic fossil is a rostrum: this specimen preserves both teeth and ring-shaped tooth pedicels (Figure 1A). The tooth pedicels form elevations on the bottoms of two wider grooves that extend over the entire preserved length of the specimen. Teeth occupy the tops of many of the pedicels (Parris et al. 2001). Tooth crowns and pedicels are also preserved on a specimen from the Bluffport Marl Member of the Demopolis Chalk in Marengo County, Alabama, ASM-PV 994.2.111 (Parris et al. 2001). Both tooth-bearing specimens are Late Cretaceous in age. Other Cretaceous specimens, including Leidy’s type material from New Jersey (ANSP 5186-5188) and the Cretaceous specimen that was sectioned for this study (Figure 1C) preserve tooth pedicels but do not preserve any teeth. Some Eocene specimens from North Carolina (Figure 1B, D) and Alabama (Parris et al. 2001) preserve tooth pedicels; these specimens also lack associated teeth. No tooth crowns were preserved with any of the Eocene specimens we have examined to date. Tooth pedicels in Eocene specimens are smaller than those in Cretaceous specimens (Parris et al. 2001). Tooth pedicels on the New Jersey Cretaceous specimen are about 1mm across. Tooth pedicels on the North Carolina Eocene specimens are about 0.4mm across. Absence of tooth crowns in most of the specimens that preserve tooth pedicels suggests the teeth are easily lost post mortem. Both of the Cylindracanthus speci- mens that preserve teeth (SDSM 30638 and ASM-PV 994.2.111) were recovered from sediments which record low energy depositional environments (shale and chalk respectively). Tooth pedicels are not preserved in all Eocene specimens. Their absence might be taphonomic given that some Cylindracanthus specimens show significant loss of surface detail due to transport abrasion. The pedicels in Cylindracanthus occupy a relatively protected position at the bottoms of the surface grooves, however, and some relatively unworn Eocene specimens (such as ASM-PV 989.4.200) lack tooth pedicels. Fierstine (2001) reported that tooth pedicels are found on only some Cylindracanthus specimens. Teeth and tooth pedicels may have been entirely lacking in some Eocene members of this genus. 132 Proceedings of the South Dakota Academy of Science, Vol. 96 (2017) Distribution of Cylindracanthus. Cylindracanthus fossils have been found in North America, Europe, and Africa. Specimens of Cylindracanthus range from Cenomanian (Vullo et al. 2009) to Eocene (Leriche 1905; Fierstine 2001) in age, and Cylindracanthus has been reported from rocks as young as Miocene (Adnet et al. 2010). The South Dakota specimen (SDSM 30638) comes from the Verendrye Formation of the Pierre Shale Group (Campanian). Other speci- mens included in this study were recovered from the Cretaceous and Eocene of Alabama, the Eocene of North Carolina, and the Cretaceous of New Jersey. Figure 1. Cretaceous (A, C) and Eocene (B, D) specimens. Arrows indicate locations of tooth-bearing grooves. A. South Dakota Cretaceous specimen SDSM 30638 showing one of two grooves in which tooth pedicels and teeth are located. B. Eocene from North Carolina. The two wider grooves with tooth pedicels are visible running down the center Proceedings of the South Dakota Academy of Science, Vol. 96 (2017) 133 of the specimen. A and B are at the same scale. C. New Jersey Cretaceous specimen donated by Mr. Tony Fabian prior to sectioning. A line of tooth pedicels is visible running along the length of the specimen. Scale is in millimeters. The specimen as figured here was sacrificed; it is now preserved as histologic slides NJSM PH1 through NJSM PH7. D. Eocene specimen from North Carolina showing a closer view of the tooth pedicels in wider grooves near the middle and bottom of the image. C and D are at the same scale to facilitate comparison of tooth pedicel sizes in Cretaceous and Eocene specimens; scale is in millimeters. The specimen figured in B and D is in the private collection of Mr. Eric Sadorf. METHODS Transverse and longitudinal histologic thin sections were made through tooth pedicels on a Cretaceous Cylindracanthus specimen from New Jersey (now NJSM PH1 through PH7) and an Eocene Cylindracanthus specimen from North Carolina (NJSM 24275). Lesions on a second Eocene specimen from North Carolina (NJSM 24276) were sectioned. No Cylindracanthus teeth were sectioned due to the extreme rarity of tooth-bearing specimens. Transverse and longitudinal histologic sections were made on segments removed from the middle of the bill and predentary of an Atlantic blue marlin taxidermy mount (Makaira cf. nigricans, NJSM TC1061). One marlin bill longitudinal section was oriented horizontally to sample histology along the length of the tooth-bearing band that covers the lateral aspect of the bill. Polyodon (paddlefish) sections were cut from a dermal scale removed near the edge of the rostrum about a quarter of the way behind the tip of the rostrum and from the rostral portion of one of the nasal bones of a desiccated head collected from the Missouri River banks south of Yankton, South Dakota (NJSM B477). The Polyodon dermal scale was sectioned horizontally, parallel to its surface. Both a transverse section and a vertical (para- sagittal) longitudinal section were made of the paddlefish nasal bone. Histologic sections were prepared at the New Jersey State Museum by one of us (R.A.P). Fossil specimens were impregnated with Epo-tek 301 epoxy resin before thin sectioning; impregnation was done under vacuum to minimize bubbles. The modern Polyodon specimens were not impregnated prior to sectioning. Both transverse (mediolateral) and longitudinal (anteroposterior) histologic sections were made of Cylindracanthus, Makaira, and Polyodon. All thin sections were cut using a low-deformation wafer saw, ground flat on lap tables,