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818


Journal of Paleontology 89(5):802–820


turtles (Terrapene carolina) in Oklahoma and Missouri (Schmidt, 1946; Carpenter, 1956; Schwartz and Schwartz, 1974; Ernst and Ernst, 1977). These pits were observed solely on the carapace, primarily on, or immediately posterior of, the suture between the second and third costal carapace epidermal scutes (Carpenter, 1956). The size of the pits is variable, from slight indentations to deep holes in the dermal bone. The pits were observed on 56% (174 out of 375) box turtles examined (Carpenter, 1956). Although ticks were not observed in association with these holes, it seems likely that they were tick-generated. They are similar in size and position to those observed from the neotropics, and significantly, occur solely on terrestrial box turtles and are absent on aquatic taxa (Chelydra serpentina, Kinosternon flavescens, Pseudemys scripta) collected with the same area (Carpenter, 1956).


Significance of the South Pass borings.—Turtle fossils are exceptionally abundant in the Eocene Wasatch, Bridger and Green River formations in the Green River Basin, Wyoming. Representatives of many chelonian groups including baenids, dermatemydids, trionychids, carettocheylids, and emydids, occur throughout the basin. Several papers have discussed the taxonomy and taphonomy of these taxa (e.g. Gunnell and Bartels, 1994; Zonneveld et al., 2000; Brand et al., 2000); however, pre-mortem bone damage was not reported in most of these manuscripts. Hutchison and Frye (2001) summarized the various types of pre-mortem bone damage they had observed on Cenozoic turtle bone, which includes predation damage (commonly from crocodilians), and damage likely attributable to fungal or bacterial infections such as USD and SCUD. Several of the forms they illustrate are similar to the borings discussed herein. These borings occur on Cenozoic specimens from the Bighorn, Wind River, and Green River Basins but are not generally very abundant (Hutchison and Frye, 2001). Over the past two decades, we have observed several


thousand turtle specimens, collected from the primarily fluvial Wasatch and Bridger formations and primarily lacustrine and marginal lacustrine Green River Formation in the Bridger Basin (western Green River Basin), Big Island/Blue Rim area (central Green River Basin), and South Pass area (northeastern Green River Basin). Karethraichnus occurs on rare geoemydid specimens and, even less commonly, on dermatemydid and baenid specimens throughout the basin. Thatchtelithichnus has, to date, only been observed on emydid specimens within the study area. Only in the study area have we observed Karethraichnus to be common. Indeed, essentially every single cf. Echmatemys sp. specimen collected or observed at South Pass is characterized by these traces. The paucity of these pre-mortem bone borings in most areas and their comparative abundance in the study area indicates some local condition that facilitated the short-term, localized spread of the parasite taxon/ taxa must have been present.


Summary


Borings, emplaced during the lifetime of the host organism, are common on the plastron and carapace of geoemydid turtles from the lowermost beds of the early Eocene Cathedral Bluffs Tongue of the Wasatch Formation in the northeastern part of the


References


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. 5890–5902.


Green River Basin near South Pass, Wyoming. Individual turtle shells are characterized by a single hole to over 100 borings. These borings are herein separated into two new ichno-


genera and four new ichnospecies. Karethraichnus lakkos are shallow, hemispherical pits with rounded, to flattened bases. They range in size from ~1mm to >5mm across. Karethraichnus kulindros are deep, non- penetrative traces with a cylindrical profile, an axis approximately perpendicular to


slightly oblique to the substrate surface and with rounded to flattened, hemispherical termini. They are commonly deeper than they are wide. They range in width from 0.5 to 5.5mm and in depth from 0.5 to 6.0mm. Karethraichnus fiale penetrate completely through the turtle shell. They are cylindrical to bi-convex or flask-shaped in profile, with an axis that runs approximately perpendicular to the substrate surfaces. Thatchtelithichnus holmani consist of shallow, non-penetrative ring-shaped borings, with a central pedestal or platform. Commonly the central platform consists of unmodified bone, although examples of rounded pillars or pedestals also occur. The position of the borings on the shells, and evidence of syn-


emplacement healing of the borings in several of the turtles, indi- cates that these borings were emplacement by ectoparasites/ mesoparasites while the animals were living. The identity of the parasite remains unknown. Ticks have been known to bore into the bones of modern terrestrial turtles leaving pits that are similar to K. lakkos described herein. Leeches have been observed attached to blood sinuses in aquatic turtle carapaces and plastrons. They are capable of bone decalcification, although the morphol- ogy of the bone damage has not been illustrated. Similarly, some vermiform parasites, such as spirorchid liver flukes, may cause lesions on the carapace and plastron of aquatic turtles. Regardless of parasite identity, the paucity of these bone borings in other areas, and their comparative abundance in the study area indicate some local condition that facilitated the short-term, localized spread of the parasite taxon/taxa.


Acknowledgments


We thank Richard Bromley, George Pemberton, and Leif Tapanila for their advice on borings in shells. We thank Adolf Seilacher for his discussions on turtle parasites, particularly barnacles.Wealso thank Leif Tapanila and Richard Bromley for their advice on the taxonomy of borings. We thank collections personnel at the University of Wyoming, University of Michigan, Michigan State University, and University of Alberta for help with both fossil and modern chelonian collections. We are also grateful to Murray Gingras, Jon Bloch, and Michael Frick for discussions on ichnology, paleontology, aquatic turtle ecology, and the distribution of parasites in modern turtle populations. Finally, we would like to acknowledge the assistance and helpful criticism of Journal of Paleontology Editor Steve Hageman, Associate Editor Gabriella Mangano, reviewer Mark Wilson, and one anonymous reviewer.


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