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Bourque—Middle–Late Miocene Kinosternon


the N1–C1 suture was more commonly observed. V3 is particularly long and overlaps the anterior of the proximal end of the C6 pair at their medial contact and does not overlap a neural (N5). The V3–4 sulcus typically intersects the C5 pair and N5 in other kinosternines. This character has been included as an autapomorphy in the diagnosis for Kinosternon pannekollops, but perhaps more specimens will elucidate how variable this feature is. The posterior shape of V4 is pointed posteriorly in FAM 12778, which was atypical in modern Kinosternon examined; however, some Kinosternon flavescens exhibit a similar V4 shape. The lateral sulci of V5 greatly overlap the suprapygal as seen in the holotype of K. rincon and to a lesser degree in specimens of K. flavescens, Kinosternon herrerai, Kinosternon integrum, and Kinosternon cruentatum. A specimen of Kinosternon leucostomum, UF/H 165987, exhibits a high degree of V5 to suprapygal overlap similar to that observed in FAM 12778. Marginals 1–3 broadly underlap the anterior


carapacial margin as with many species of Kinosternon. Conversely, underlap is narrow in Staurotypus, Sternotherus, and some Kinosternon such as Kinosternon steindachneri. Marginal 9 is as high as M8 and both are dorso-ventrally short as is the state with most kinosternines. Marginal 9 is not heightened to the extent of M10. Marginal 10 is notably heightened in FAM 12778, much more so than M8–9 and M11.


Plastron: The plastron of FAM 12778 is completely


preserved, with only some minor damage along anterior margins of the forelobe and anterior suture of the left hyoplastron. The length of the plastron is 125.72mm and 83% of the carapace length. It is comprised of paired epi-, hyo, hypo-, and xiphiplastra (Figs. 4.2 and 4.4; 5.2 and 5.6; and 6). While no substantial trace of an entoplastron is present in


FAM 12778, a minute anomalous bony structure exists poster- omedially on the ventral side of the right epiplastron at its contact with the hyoplastron. It is unclear if the ovate structure represents an entoplastral vestige, a minor pathology, or a small area of remodeled bone (e.g., a bony plug within a pockmark) along the forelobe hinge. This structure is situated in a small pockmark, and a symmetrical pock is present on the left epiplastron. Bramble et al. (1984) and I have observed atavistic entoplastra in the form of tiny rudimentary bones in modern specimens of Sternotherus odoratus (e.g., KU 7849 and UF/H 165594), but these appear to be dissimilar to what is present in FAM 12778. Character 31 in the phylogenetic analysis presented here was scored as lacking a substantial entoplastron for FAM 12778. The plastral forelobe is ~43.16mm long and 68.45mm


wide. It is expansive and would have likely been capable of encapsulating the anterior portion of the turtle in life. The forelobe was highly kinetic, evident by the stepped scute margins at the contact between the anterior humeral and posterior humeral scutes (Hutchison and Bramble, 1981), and rounded over sutural teeth along the epi–hyoplastral suture. The intergular scute is 16.68mm long (11% carapace length) and 24.06mm wide (16% carapace length). The length of the anterior humeral scute is 6.49mm at the midline and 4% of the carapace length. The axillary notch of the hyoplastron is moderate in depth, and typical for species with broad forelobes.


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It is not as deep as taxa with more narrow forelobes (e.g., Kinosternon steindachneri), nor is it diminished to the degree seen in Kinosternon cruentatum and Kinosternon leucostomum. The plastral hindlobe is ~43.76mm long and 61.46mm


wide. The anal scute is 29.45mmlong at the midline and ~20% of the carapace length. The femoral scute is moderately long at the midline and ~37% of the hindlobe length. The hindlobe is less expansive than the forelobe in relation to the width of the carapace, and would unlikely have been capable of fully encapsulating the hind portions of the turtle. Like the forelobe, the hindlobe was also highly kinetic made evident by rounded over sutural teeth along the hypo–xiphiplastral suture. A high degree of bone remodeling on the ventral hypoplastron just posterior to the posterior humeral–femoral scute sulcus likewise suggests this. Additionally, there are indications of connective tissue or skin scarring along the hypo–xiphiplastral suture, which would have facilitated kinesis. FAM 12778 almost certainly represents a male in possessing a distinctly concave hindlobe with the distal margins of the xiphiplastra curved ventrally (Figs. 5.6 and 6). A distinct caudal notch is present inFAM12778 and it is unknown if this feature is sexually dimorphic for Kinosternon pannekollops. Along the midline of the plastron, e.g., from the posterior of the epiplastra to the anal scutes, sulci that likely delineate interplastral scute skin are present. This interplastral scute skin was moderate (not as developed as that in many Sternotherus) and potentially gender informative as plastral skin is more typically seen in males than females of extant Kinosternon. The combined length of the axillary and inguinal bridge


buttresses is 34.3mm and ~23% of the carapace length. The axillary and inguinal inframarginal scutes contact one another on the distal hyoplastron, and the axillary–inguinal sulcus is wholly contained on the hyoplastron as with other species of Kinosternon. Conversely, in Sternotherus and Staurotypus, the axillary–inguinal sulcus overlaps onto the peripherals. In FAM 12778, the M5–6 sulcus broadly underlaps onto the hyoplastron and contacts the inguinal scute. Marginal 5 is extensive on the hyoplastron. A distinct inguinal groove is present, just proximal to the inguinal scute on the hypoplastron. This groove terminates anteriorly on the posterior-most hyoplastron. The inguinal bridge buttress is inclined as in Kinosternon wakeeniense, the subrubrum group, Kinosternon flavescens, and Kinosternon arizonense.


Etymology.—Species name is the combination of the Anglo- Saxon ‘panne’ for ‘pan’ and Greek ‘kollops’ for ‘handle,’ in reference to the panhandle region of northern Texas where the holotype was collected.


Occurrence.—Type locality: Lewis Quarry 7, Donley County, Texas. Age and formation: Late Miocene, Clarendonian NALMA, 12–9.5 Ma, Ogallala Formation (Schultz, 1977:fig. 6; Schultz, 1990:fig. 3; Tedford et al., 2004).


Remarks.—Kinosternon pannekollops is the largest known kinosternid from the Miocene of North America. Its large body size suggests that dwarfing has occurred over time


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