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Borings in early Eocene turtle shell from the Wasatch Formation, South Pass, Wyoming

Published online by Cambridge University Press:  29 March 2016

John-Paul Zonneveld
University of Alberta, Edmonton, Alberta T2G 2E3, Canada 〈〉
William S. Bartels
Department of Geological Sciences, Albion College, Albion, Michigan 49224, USA 〈〉
Gregg F. Gunnell
Division of Fossil Primates, Duke Lemur Center, Durham, North Carolina, 27705, USA 〈〉
Luke P. McHugh
Canadian Natural Resources Limited, Calgary, Alberta T2P 2Z2, Canada


Borings in fossil turtle shells collected from the lowermost beds of the early Eocene Cathedral Bluffs Tongue of the Wasatch Formation in the northwestern part of the Green River Basin near South Pass, Wyoming, are herein described. Individual turtle shells in the study area are characterized by as few as one or two and as many as >100 borings. The borings include both non-penetrative forms (those which do not pass fully though the shell) as well as penetrative forms (those which pass fully from the exterior to the interior surface of the shell). All non-penetrative forms occur on external surfaces of the carapace and plastron (i.e. those that would have been accessible while the host taxon was alive). Two new ichnogenera and four new ichnospecies are established to describe these borings. Karethraichnus (new ichnogenus) includes three ichnospecies: K. lakkos (new ichnospecies), K. kulindros (new ichnospecies), and K. fiale (new ichnospecies). Karethraichnus lakkos are shallow (non-penetrating), hemispherical pits with rounded, to flattened bases. Karethraichnus kulindros are deep, non-penetrative traces with a cylindrical profile, an axis approximately perpendicular to the substrate surface and with rounded to flattened, hemispherical termini. Karethraichnus fiale are penetrative traces with a cylindrical to bi-convex or flask-shaped profile, and an axis approximately perpendicular to the substrate surface. Thatchtelithichnus (new ichnogenus) Thatchtelithichnus holmani (new ichnospecies) consist of non-penetrative borings into a bone substrate. They consist of a ring-shaped trace, with a central pedestal or platform. The position of the borings on the shells, and evidence of syn-emplacement healing of the borings in several of the turtles, indicates that these borings were emplacement by ectoparasites/mesoparasites while the animals were living. Similar traces in modern emydid turtles are attributed to ticks, leeches, or spirorchid liver flukes.

Copyright © 2016, The Paleontological Society 

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Achrai, B., and Wagner, H.D., 2013, Micro-structure and mechanical properties of the turtle carapace as a biological composite shield: Acta Biomaterialia, v. 9, p. 58905902.CrossRefGoogle ScholarPubMed
Alibardi, L., and Thompson, M., 2006, Cytochemical, biochemical and molecular aspects of the process of keratinization in the epidermis of reptilian scales: Progress in Histochemistry and Cytochemistry, v. 40, p. 73134.CrossRefGoogle ScholarPubMed
Allen, R., and Neill, W.T., 1952, Know your reptiles: the diamondback terrapin: Florida Wildlife, v. 6, p. 8, 42 p.Google Scholar
Arndt, R.G., 1975, The occurrence of barnacles and algae on the red-bellied turtle Chrysemys r. rubiventris (le Conte): Journal of Herpetology, v. 9, p. 357359.CrossRefGoogle Scholar
Ascenzi, A., and Silvestrini, G., 1984, Bone-boring marine micro-organisms: an experimental investigation: Journal of Human Evolution, v. 13, p. 531536.CrossRefGoogle Scholar
Baalbergen, E., Helwerda, R., Schelfhorst, R., Castillo Cajas, R.F., van Moorsel, C.H.M., Welter–Schulkes, F.W., Giokas, S., and Schilhuizen, M., 2014, Predator-prey interactions between shell-boring beetle larvae and rock-dwelling land snails: PLOS One, v. 6, p. e0100366.Google Scholar
Bader, K.S., Hasiotis, S.T., and Martin, L.D., 2009, Application of forensic science techniques to trace fossils on dinosaur bones from a quarry in the Upper Jurassic Morrison Formation, northeastern Wyoming: PALAIOS, v. 24, p. 140158.CrossRefGoogle Scholar
Behrensmeyer, A.K., 1978, Taphononomic and ecologic information from bone weathering: Paleobiology, v. 4, p. 150162.CrossRefGoogle Scholar
Belaústegui, Z., Gibert, J.M., de., Domenech, R., Muñiz, D., and Martinell, J., 2012, Clavate borings in a Miocene cetacean skeleton from Tarragona (NE Spain) and the fossil record of marine bone bioerosion: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 323–325, p. 6874.CrossRefGoogle Scholar
Bertling, M., Braddy, S.J., Bromley, R.G., Demathieu, G.R., Genise, J., Mikulas, R., Nielsen, J.K., Nielsen, K.S.S., Rindsberg, A.K., Schlirf, M., and Uchman, A., 2006, Names for trace fossils: a uniform approach: Lethaia, v. 39, p. 265286.CrossRefGoogle Scholar
Bielecki, A., Cichocka, J.M., Jabloński, A., Jeleń, I, Ropelewska, E., Biedunkiewicz, A., Terlecki, J., Nowakowski, J.J., Pakulnick, J., and Szalchciak, J., 2012, Coexistence of Placobdella costata (Fr, Müller, 1846) (Hirudinida: Glossiphoniidae) and mud turtle Emys orbicularis: Biologia, v. 67, p. 731738.CrossRefGoogle Scholar
Bishop, B.E., Savitsky, B.A., and Abdel-Fattah, T., 2007, Ulcerative shell disease (USD) and its possible relationship to bioaccumulation of lead (Pb) in aquatic turtles in an urban environment: Virginia /West Virginia Water Research Symposium, p. 182190.Google Scholar
Boycott, A.E., 1921, The land Mollusca of the parish of Aldenham: Transactions of the Hertfordshire Natural History Society and Field Club, v. 17, p. 220245.Google Scholar
Boycott, A.E., 1934, The habits of land Mollusca in Britain: Journal of Ecology, v. 22, p. 138.CrossRefGoogle Scholar
Brand, L.R., Goodwin, H.T., Ambrose, P.D., and Buchheim, H.P., 2000, Taphonomy of turtles in the Middle Eocene Bridger Formation, SW Wyoming: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 162, p. 171189.CrossRefGoogle Scholar
Brett, C.E., 1985, Tremichnus: a new ichnogenus of circular–parabolic pits in fossil echinoderms: Journal of Paleontology, v. 59, p. 625635.Google Scholar
Britt, B.B., Scheetz, R.D., and Dangerfield, A., 2006, A suite of dermestid beetle traces on dinosaur bone from the Upper Jurassic Morrison Formation, Wyoming, USA, Ichnos, v. 15, p. 5971.Google Scholar
Bromley, R.G., 1981, Concepts in ichnology illustrated by small round holes in shells: Acta Geològica Hispànica, v. 16, p. 5564.Google Scholar
Bromley, R.G., 1993, Predation habits of octopus past and present and a new ichnospecies, Oichnus ovalis: Bulletin of the Geological Society of Denmark, v. 40, p. 167173.Google Scholar
Bromley, R.G., 2004, A stratigraphy of marine bioerosion, in McIlroy, D., ed., The application of ichnology to palaeoenvironmental and stratigraphic analysis: Geological Society of London Special Publication, v. 228, p. 455479.Google Scholar
Bromley, R.G., and Frey, R.W., 1974, Redescription of the trace fossil Gyrolithes and taxonomic evaluation of Thalassinoides, Ophiomorpha and Spongeliomorpha: Bulletin of the Geological Society of Denmark, v. 23, p. 311335.Google Scholar
Brooks, R.J., Galbraith, D.A., and Layfield, J.A., 1990, Occurrence of Placobdella parasitica (Hirudinea) on snapping turtles, Chelydra serpentina, in southeastern Ontario: The journal of Parasitology, v. 76, p. 190195.CrossRefGoogle Scholar
Cabral., U.G., Riff, D., Kellner, A.W.A., and Henriques, D.D.R., 2011, Pathological features and insect boring marks in a crocodyliform from the Bauru Basin, Cretaceous of Brazil: Zoological Journal of the Linnean Society, v. 163, p. S140S151.CrossRefGoogle Scholar
Caine, E.A., 1986, Carapace epibionts on nesting loggerhead sea turtles, p. Atlantic coast of U.S.A.: Journal of Experimental Marine Biology and Ecology, v. 5, p. 1526.CrossRefGoogle Scholar
Carpenter, C.C., 1956, Carapace pits in the three-toed box turtle, Terrapene carolina triunguis (Chelonia-Emydidae): The Southwest Naturalist, v. 1, p. 8386.CrossRefGoogle Scholar
Donovan, S.K., 2014, When is a fossil not a fossil? When it is a trace fossil: Lethaia. doi, p. 10.1111/let.12089Google Scholar
Donovan, S.K., and Jagt, J.W.M., 2002, Oichnus Bromley borings in the irregular echinoid Hemipneustes Agassiz from the type Maastrichtian (Upper Cretaceous, The Netherlands and Belgium): Ichnos, v. 9, p. 6774.CrossRefGoogle Scholar
Ellis, A.E., 1969, British Snails, Oxford, Clarendon Press, 298 p.Google Scholar
Ernst, C.H., and Ernst, E.M., 1977, Ectoparasites associated with neotropical turtles of the genus Callopsis (Testudines, Emydidae, Batagurinae): Biotropica, v. 9, p. 139142.CrossRefGoogle Scholar
Esch, G.W., and Gibbons, J.W., 1967, Seasonal incidence of parasitism in the Painted Turtle Chrysemys picta marginata Agassiz: The Journal of Parasitology, v. 53, p. 818821.CrossRefGoogle ScholarPubMed
Fairchild, G.B., 1943, An annotated list of the bloodsucking insects, ticks and mites known from Panama: American Journal of Tropical Medicine, v. 23, p. 569591.CrossRefGoogle Scholar
Fiorillo, A.R., 1987, Trample marks: caution from the Cretaceous: Current Research in the Pleistocene, v. 4, p. 7375.Google Scholar
Fiorillo, A.R., 1990, Utilization of prey bones by predatory dinosaurs: Journal of Vertbrate Paleontology, Abstracts of papers 10 (supplement to 3), p. 22A.Google Scholar
Flynn, R.J., 1973, Parasites of laboratory animals, Ames, Iowa, Iowa State University Press, 840 p.Google Scholar
Frick, M.G., Williams, K.L., and Robinson, M., 1998, Epibionts associated with nesting loggerhead sea turtles (Caretta caretta) in Georgia, USA: Herpetological Review, v. 29, p. 211214.Google Scholar
Frick, M.G., Williams, K.L., Veljacic, D.X., Jackson, J.A., and Knight, S.E., 2000, Epibiont community succession on nesting loggerhead sea turtles, Caretta caretta, from Georgia, USA: Proceedings of the 20th Annual Sea Turtle Symposium, Orlando, Florida, p. 280282.Google Scholar
Fürsich, F.T., 1973, A revision of the trace fossils Spongeliomorpha, Ophiomorpha and Thalassinoides: Neus Jahrbuch fur Geologie und Paläontologie Monatshefte, v. 1973, p. 719735.Google Scholar
Garner, M.M., Herrington, R., Howerth, E.W., Homer, B.L., Nettles, V.F., Izara, R., Shotts, E.B. jr, and Jacobsen, E.R., 1997, Shell disease in river cooters (Pseudemys concinna) and yellow-bellied turtles (Trachemys scripta) in a Georgia (USA) lake: Journal of Wildlife Diseases, v. 33, p. 7886.CrossRefGoogle Scholar
Graham, T.E., Saumure, R.A., and Ericson, B., 1997, Map turtle leech loads: The Journal of Parasitology, v. 83, p. 11851186.CrossRefGoogle ScholarPubMed
Gunnell, G.F., and Bartels, W.S., 1994, Early Bridgerian (Middle Eocene) vertebrate paleontology and paleoecology of the southern Green River Basin, Wyoming: Contributions to Geology, University of Wyoming, v. 30, p. 5770.Google Scholar
Häntzschel, W., 1962, Trace fossils and problematica. In. Treatise on Invertebrate Paleontology, part W, Moore, R.C., ed). Geological Society of America, and University of Kansas, New York and Lawrence, p. 177245.Google Scholar
Häntzschel, W., 1965, Vestigia invertebratorum et problematica, in Westphal, F., ed., Fossilium catalogus, v. 108, p. 1140.Google Scholar
Hasiotis, S.T., 2004, Reconnaissance of Upper Jurassic Morrison Formation Ichnofossils, Rocky Mountain Region, USA: paleoenvironmental, stratigraphic, and Paleoclimatic significance of terrestrial and freshwater ichnocoenoses: Sedimentary Geology, v. 167, p. 177268.CrossRefGoogle Scholar
Hefti, E., Trechsel, U., Rufenacht, H., and Fleisch, H., 1980, Use of dermestid beetles for cleaning bones: Calcified Tissue International, v. 31, p. 4547.CrossRefGoogle ScholarPubMed
Higgs, N.D., Glover, A.G., Dahlgren, T.G., and Little, C.T.S., 2011, Bone–boring worms, p. characterizing the morphology, rate, and method of bioerosion by Osedax mucofloris (Annelida, Siboglinidae): Biological Bulletin, v. 221, p. 307316.CrossRefGoogle Scholar
Hulse, A.C., 1976, Carapacial and plastral flora and fauna of the Sonora Mud Turtle, Kinosternon sonoriense Conte (Reptilia, Testudines, Kinosternidae): Journal of Herpetology, v. 10, p. 4548.CrossRefGoogle Scholar
Hutchison, H., and Frye, F.L., 2001, Evidence of pathology in early Cenozoic turtles: Paleobios, v. 21, p. 1219.Google Scholar
Jackson, C.G., and Ross, A., 1971a, Molluscan fouling of the Ornate Diamondback Terrapin Malaclemys terrapin macrospilota Hay: Herpetologica, v. 27, p. 341344.Google Scholar
Jackson, C.G., and Ross, A., 1971b, The occurrence of barnacles on the alligator snapping turtle Macroclemys temmincki (Troost): Journal of Herpetology, v. 5, p. 188189.CrossRefGoogle Scholar
Jackson, C.G. Jr., and Ross, A., 1972, Balanomorph barnacles on Chrysemys alabamensis: Quarterly Journal of the Florida Academy of Science, v. 35, p. 173176.Google Scholar
Jacobson, E.R., Cheatwood, J.L., and Maxwell, L.K., 2000, Mycotic diseases of reptiles: Seminars in Avian and Exotic Pet Medicine, v. 9, p. 94101.CrossRefGoogle Scholar
Jans, M.M.E., 2008, Microbial bioerosion of bone – a review, in Wisshak, M., and Tapanila, L., eds., Current developments in bioerosion: Erlangen Earth Conference Series, Berlin, Springer Verlag, p. 397413.CrossRefGoogle Scholar
Johnson, C.A., Griffith, J.W., Tenorio, P., Hytrek, S., and Lang, C.M., 1998, Fatal trematodiasis in research turtles: Comparative Medicine, v. 48, p. 340343.Google ScholarPubMed
Johnson, J.H., 2004, Husbandry and medicine of aquatic reptiles: Seminars in Avian and Exotic Pet Medicine, v. 13, p. 223228.CrossRefGoogle Scholar
Kaiser, T.M., 2000, Proposed modification to fossil mammalian bone from Plio–Pleistocene hominid-bearing deposits of Laetoli (northern Tanzania): Annals of the Entomological Society of America, v. 93, p. 693700.CrossRefGoogle Scholar
Kaplan, H.M., 1957, Septicemic cutaneous ulcerative disease of turtles: Proceedings of the Animal Care Panel, v. 7, p. 273277.Google Scholar
Kelly, S.R.A., and Bromley, R.G., 1984, Ichnological nomenclature of clavate borings: Palaeontology, v. 27, p. 793807.Google Scholar
Kennedy, W.J., 1967, Burrows and surface traces from the Lower Chalk of southern England: Bulletin of the British Museum, v. 15, p. 127167.Google Scholar
Kiel, S., Goedert, J.L., Kahl, W.-A., and Rouse, G.W., 2010, Fossil traces of the bone-eating worm Osedax in early Oligocene whale bones: Proceedings of the National Academy of Sciences, v. 107, p. 86568659.CrossRefGoogle ScholarPubMed
Koffler, B.R., Seigel, R.A., and Mendonca, M.T., 1978, The seasonal occurrence of leeches on the wood turtle, Clemmys insculpta (Reptilia, Testudines, Emydidae): Journal of Herpetology, v. 12, p. 571572.CrossRefGoogle Scholar
Lovich, J.E., Gotte, S.W., Ernst, C.H., Harshbarger, J.C., Laemmerzahl, A.F., and Gibbons, J.W., 1996, Prevalence and histopathology of shell disease in turtles from Lake Blackshear, Georgia: Journal of Wildlife Diseases, v. 32, p. 259265.CrossRefGoogle ScholarPubMed
MacCulloch, R.D., 1981, Leech parasitism on the Western Painted Turtle, Chrysemys picta belli in Saskatchewan: The Journal of Parasitology, v. 67, p. 128129.CrossRefGoogle Scholar
Martin, D.R., 1972, Distribution of helminth parasites in turtles native to southern Illinois: Transactions of the Illinois Academy of Science, v. 65, p. 6167.Google Scholar
McAucliffe, J.R., 1977, An hypothesis explaining variations of hemogregarine parasitemia in different aquatic turtle species: The Journal of Parasitology, v. 63, p. 580581.CrossRefGoogle Scholar
McCoy, J.C., Failey, E.L., Price, S.J., and Dorcas, M.E., 2007, An assessment of leech parasitism on semi-aquatic turtles in the western Piedmont of North Carolina: Southeastern Naturalist, v. 6, p. 191202.CrossRefGoogle Scholar
Mittermeier, R.A., 1971, Notes on the behavior and ecology of Rhinoclemys annulata Gray: Herpetologica, v. 27, p. 485488.Google Scholar
Mordan, P.B., 1977, Factors affecting the distribution and abundance of Aegopinella and Nesovitrea (Pulmonata: Zonitidae) at Monks Wood National Nature Reserve, Huntingdonshire: Biological Journal of the Linnean Society, v. 9, p. 5972.CrossRefGoogle Scholar
Müller, A.H., 1970, Über ichnia von typ Ophiomorpha und Thalassinoides (Vestigia invertebratorum Crustacea): Deutschland Akadamie Wiss. Berlin Mh, v. 12, p. 775787.Google Scholar
Müller, A.H., 1977, Zur ichnologie der subherzynen Oberkreide (Campan): Zeitschrift für geologische Wissenschaften, Berlin, v. 5, p. 881897.Google Scholar
Nielsen, K.S.S., and Nielsen, J.K., 2001, Bioerosion in Pliocene to late Holocene tests of benthic and planktonic foraminiferans, with a revision of the ichnogenera Oichnus and Tremichnus: Ichnos, v. 8, p. 99116.CrossRefGoogle Scholar
Oceguera-Figueroa, A., and Siddall, M.E., 2008, Placobdella lamothei n. sp. (Hirudinea: Glossiphoniidae), a new leech parasite of freshwater turtles from Estado de México, Mexico: Revista Mexicana de Biodiversidad, v. 79, p. 135139.Google Scholar
Paik, I.S., 2000, Bone chip–filled burrows associated with bored dinosaur bone in floodplain paleosols of the Cretaceous Hasandong Formation, Korea: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 157, p. 213225.CrossRefGoogle Scholar
Palmer, T.J., and Wilson, M.A., 1988, Parasitism of Ordovician bryozoans and the origin of pseudoborings: Palaeontology, v. 31, p. 939949.Google Scholar
Pfaller, J.B., Frick, M.G., Reich, K.J., Williams, K.L., and Bjorndal, K.A., 2008, Carapace epibionts of loggerhead turtles (Caretta caretta) nesting at Canaveral National Seashore, Florida: Journal of Natural History, v. 42, p. 10951102.Google Scholar
Pickerill, R.K., and Donovan, S.K., 1998, Ichnology of the Pliocene Bowden shell bed, southeast Jamaica, in Donovan, S.K., ed., The Pliocene Bowden Shell Bed, southeast Jamaica: Contributions to Tertiary and Quaternary Geology, v. 35, p. 161175.Google Scholar
Pirrone, C.A., Buatois, L.A., and Bromley, R.G., 2014, Ichnotaxobases for bioerosion trace fossils in bones: Journal of Paleontology, v. 88, p. 195203.CrossRefGoogle Scholar
Quattier, D., 2002, Les perforations biogéniques ou ces escargots qui grignotent nos massives calcaires: Revue des Spéléologues du Grand Sud-Ouest, v. 96, p. 1415.Google Scholar
Quattier, D., 2011, Observations on the saxicavoud habits of Cepaea nemoralis (Linnaeus, 1758) (Pulmonata, Stylommatophora, Helicidae) in the Pyrenees (France): Biodiversity Journal, v. 2, p. 201206.Google Scholar
Raphael, B.L., 2003, Chelonians (turtles and tortoises), in Fowler, M.F., and Miller, R.E., eds., Zoo and Wild Animal Medicine, 5th ed., St. Louis, Missouri, Elsevier, p. 5558.Google Scholar
Readel, A.M., Phillips, C.A., and Wetzel, M.J., 2008, Leech parasitism in a turtle assemblage, p. effectsof host and environmental characteristics: Copeia, v. 2008, p. 227233.CrossRefGoogle Scholar
Roberts, E.M., Rogers, R.R., and Foreman, B.Z., 2007, Continental insect borings in dinosaur bone: examples from the late Cretaceous of Madagascar and Utah: Journal of Paleontology, v. 81, p. 201208.CrossRefGoogle Scholar
Rogers, R.R., 1992, Non-marine borings in dinosaur bones from the Upper Cretaceous Two Medicine Formation, northwestern Montana: Journal of Vertebrate Paleontology, v. 12, p. 528531.CrossRefGoogle Scholar
Roehler, H.W., 1991, Revised stratigraphic nomenclature for the Wasatch and green River Formations of Eocene age, Wyoming, Utah and Colorado: United States Geological Survey Professional Paper, v. 1506C, 17 p.Google Scholar
Roehler, H.W., 1992, Correlation, composition, areal distribution, and thickness of Eocene stratigraphic units, Greater Green River Basin, Wyoming, Utah and Colorado: United States Geological Survey Professional Paper, v. 1506E, 49 p.Google Scholar
Rose, F.L., Koke, J., Koehn, R., and Smith, D., 2001, Identification of the etiological agent for necrotizing scute disease in the Texas Tortoise: Journal of Wildlife Diseases, v. 37, p. 223228.CrossRefGoogle ScholarPubMed
Ross, A., and Jackson, C.G., 1972, Barnacle fouling of the ornate diamondback terrapin Malaclemys terrapin macrospilota: Crustaceana, v. 22, p. 203205.CrossRefGoogle Scholar
Ryan, T.J., and Lambert, A., 2005, Prevalence and colonization of Placobdella on two species of freshwater turtles (Graptemys geographica and Sternotherus odoratus): Journal of Herpetology, v. 39, p. 284287.CrossRefGoogle Scholar
Sawyer, R.T., 1986, Leech biology and behaviour, vol. 2, Feeding, biology, ecology, and systematics, New York, Oxford University Press, p. 419793.Google Scholar
Scaravelli, D., Affronte, M., and Costa, F., 2001, Analysis of epibiont presence on Caretta caretta from Adriatic Sea: Proceedings, First Mediterranean Conference on Marine Turtles, Rome, Italy, 3 p.Google Scholar
Schärer, M.T., 2003, A survey of the epibiota of Eretmochelys imbricata (Testudines, p. Cheloniidae) of Mona Island, Puerto Rico: Revista de Biología Tropical, v. 51, p. 8790.Google Scholar
Schmidt, K.P., 1946, Turtles collected by the Smithsonian biological survey of the Panama Canal Zone: Smithsonian Miscellaneous Collections, v. 106, p. 19.Google Scholar
Schwartz, C.W., and Schwartz, E.R., 1974, The three–toed box turtle in central Missouri; its population, home range, and movements: Missouri Department of Conservation Terr. Ser. 5.Google Scholar
Seigel, R.A., 1983, Occurrence and effects of barnacle infestations on Diamondback Terrapins (Malaclemys terrapin): American Midland Naturalist, v. 109, p. 3439.CrossRefGoogle Scholar
Siddall, M.E., and Gaffney, E.S., 2004, Observations on the leech Placobdella ornate feeding from bony tissues of turtles: The Journal of Parasitology, v. 90, p. 11861188.CrossRefGoogle Scholar
Shuster, M.W., and Steidtmann, J.R., 1988, Tectonic and sedimentary evolution of the northern Green River Basin, western Wyoming, in Schmidt, C.J., and Perry, W.J., eds., Interaction of the Rocky Mountain foreland and the Cordilleran thrust belt: Geological Society of America Memoir, v. 171, p. 515530.CrossRefGoogle Scholar
Stanton, W.I., 1986, Snail holes in Helixigenic cavities in hard limestone - an aid to the interpretation of karst landforms: Proceedings of the University of Bristol Speleaological Society, v. 17, p. 231232.Google Scholar
Steidtmann, J.R., and Middleton, L.T., 1991, Fault chronology and uplift history of the southern Wind River Range, Wyoming: Implications for Laramide and post-Laramide deformation in the Rocky Mountain foreland: Geological Society of America Bulletin, v. 103, p. 472485.2.3.CO;2>CrossRefGoogle Scholar
Tappan, M., 1994, Bone weathering in the tropical rain forest: Journal of Archaeological Science, v. 21, p. 667673.CrossRefGoogle Scholar
Tapanila, L., Roberts, E.M., Bouaré, M.L., Sissoko, F., and O’Leary, M.A., 2004, Bivalve borings in phosphatic coprolites and bone: PALAIOS, v. 19, p. 565573.2.0.CO;2>CrossRefGoogle Scholar
Timm, R.M., 1982, Dermestids: Field Museum of Natural History Bulletin, v. 53, p. 1418.Google Scholar
Thomas, R.D.K., 1976, Gastropod predation on sympatric Neogene species of Glycymeris (bivalvia) from the eastern United States: Journal of Paleontology, v. 50, p. 488489.Google Scholar
Wallach, J.D., 1975, The pathogenesis and etiology of ulcerative shell disease in turtles: Journal of Zoo Animal Medicine, v. 6, 1113.CrossRefGoogle Scholar
Weiner, S., and Wagner, H.D., 1998, The material bone: structure–mechanical function relations: Annual Reviews of Materials Science, v. 8, p. 271298.CrossRefGoogle Scholar
Zonneveld, J.-P., Bartels, W.S., and Clyde, W.C., 2003, Stratal architecture of an Early Eocene fluvial–lacustrine depositional system, Little Muddy Creek area, southwestern Green River Basin, Wyoming, in Raynolds, R.G., and Flores, R.M., eds., Cenozoic systems of the Rocky Mountain Region: Rocky Mountain SEPM, p. 253287.Google Scholar
Zonneveld, J.-P., and Gingras, M.K., 2014, Sedilichnus, Oichnus, Fossichnus, and Tremichnus: ‘small round holes in shells’ revisited: Journal of Paleontology, v. 88, p. 895905.Google Scholar
Zonneveld, J.-P., Gunnell, G.F., and Bartels, W.S., 2000, Early Eocene fossil vertebrates from the southwestern Green River Basin, Lincoln and Uinta counties, Wyoming: Journal of Vertebrate Paleontology, v. 20, p. 369386.CrossRefGoogle Scholar
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