CHRYSEMYS CORNICULATA SP. NOV. (FIGS 2–3, 5–7) Zoobank registration: u r n: l s i d: z o o b a n k. org:act: A1118AAC-7E71-4AD9-A130-23CB40A339B4. Holotype specimen: ETMNH-12491, complete carapace and nearly complete plastron, missing only small portions near the left and right bridges and the posterior-most rim of the plastron; set of incomplete lower jaws including the left and right dentaries; proximal left coracoid fragment; humerus fragment; complete right fibula; and numerous other indeterminate shell and bone fragments. Paratypes: ETMNH-503, complete, but partially crushed, carapace and plastron; ETMNH-3559, nearly complete carapace (missing only portions of right peripherals 2–5 and 8), incomplete plastron including both epiplastra, entoplastron, fragment of right hyoplastron, nearly complete right xiphiplastron, and nearly complete hyo, hypo and xiphiplastra, nearly complete left and right quadrates, complete right radius, two carpals, three complete to nearly complete manual phalanges, three manual unguals, a proximal left femur fragment, and numerous other indeterminate and unidentified shell and bone fragments; ETMNH-3561, incomplete carapace and nearly complete plastron, with the latter missing portions of the right hyoplastron and both bridges, humerus fragment, and numerous other indeterminate and unidentified shell and bone fragments; ETMNH-20544, nearly complete carapace and plastron missing only a few small fragments including those from right peripheral 2, left costal 3 and the left hyoplastron near the bridge, nearly complete left maxilla, nearly complete cervical vertebra 7, incomplete right and fragmentary left humeri; complete? manual ungual, fragmentary right ischium, and numerous other indeterminate and unidentified shell and bone fragments; ETMNH-20609, nearly complete shell (ontogenetically younger individual) missing portions of neural 1, left costals 1–2 and 4–5, right costals 1 and 5, all of left peripheral 8, and portions of the left hyoand right xiphiplastron, there are other places on the shell where small fragments are missing as the shell was pieced back together, and it includes numerous other indeterminate shell fragments not attached to the major preserved portion of shell. Referred specimens: ETMNH-4, right peripheral? 9 fragment; ETMNH-293, nearly complete right hyoplastron; ETMNH-297, nearly complete nuchal; ETMNH-301, left peripherals 7–8; ETMNH-3467, complete left epi- and hyoplastron; ETMNH-3527, left costal 6; ETMNH-3535, left peripheral 3; ETMNH-3544, nearly complete left epiplastron; ETMNH-3553, right peripheral 10; ETMNH-3567, incomplete left hypoplastron; ETMNH-3780, complete right xiphiplastron; ETMNH-7624, left peripheral 7; ETMNH-7631, nearly complete right costal 1 and indeterminate fragments; ETMNH-7632, articulated and nearly complete right hypoplastron and complete right xiphiplastron; ETMNH-7635, nearly complete right hypoplastron; ETMNH-7640, right peripheral 10; ETMNH-7651, nearly complete left and right hyoplastra; ETMNH-7652, complete right costal 3; ETMNH-7670, left peripheral 10 and indeterminate limb bone fragment; ETMNH-7671, neural 2; ETMNH-7673, left costal 6; ETMNH-7677, complete left epiplastron; ETMNH-7678, right costal 1 and right peripherals 8–9; ETMNH-11744, left peripherals 9–10 and complete phalanx; ETMNH-12846, right peripheral 3; ETMNH-12978, portions or right costals 2–5; ETMNH-12989, complete right costal 3; ETMNH-12990, right peripheral 5; ETMNH-12992, right peripheral 7; ETMNH-13271, right peripheral 7; ETMNH-13281, left peripheral 5; ETMNH-13282, incomplete pygal; ETMNH-13283, left peripheral 10; ETMNH-14165, incomplete peripheral 10; ETMNH-14399, costal fragment; ETMNH-14556, nearly complete right costal 4; ETMNH-15948, nearly complete right hyoplastron; ETMNH-15949, right peripheral 8; ETMNH-17273, left peripheral 8; ETMNH-17274, costal fragment; ETMNH-17257, incomplete right costal 6; ETMNH-17349, right peripheral 9. Type locality: Gray Fossil Site, Washington County, Tennessee, USA (Fig. 1). Type horizon and age: Latest Miocene-Early Pliocene (latest Hemphillian-Early Blancan NALMA). This estimate means the fossil locality, and C. corniculata, lies somewhere between Hh3 and Hh4, with recent age estimates based on biostratigraphy placing the site between 4.9–4.5 Mya. Etymology: From the Latin corniculum, deminutive of cornu, horn, for example, with small horns, referring to the anterior projections of the carapace, specifically on the nuchal, and sometimes referred to as the nuchal horns. The common name of this species is: horned painted turtle. Diagnosis: Chrysemys corniculata is placed in Emydidae due to the absence of musk ducts (would be notches on peripherals 3 and 4 if present), inframarginals reduced to two, normal hexagonal neurals 2–8 (also occurs in a few geoemydids; e.g. Mauremys Gray, 1869), and costal-inguinal buttress confined to costal 5. It is placed in the Deirochelyinae due to the lack of pectoral overlap of the entoplastron and lack of a hingeable plastral lobe with a ligamentous bridge connection (also present in some emydines). Diagnosed as a member of the genus Chrysemys on the basis of a low domed shell; anal notch absent or very weak; notching between posterior peripherals and marginals absent; dorsal carapacial keel absent; vertebral 1 with relatively slight nuchal overlap and with anterolateral flaring/projections. Distinguished from all other Chrysemys by extreme anterior projections of nuchal under marginals 1; anterior-most point of nuchal in middle of anterolateral projections (or anterolateral nuchal horns); concavity of the posterior edge of the cervical scute on the visceral surface; posterior inflation of vertebral 1; shorter and posteriorly shifted bridge with the axillary buttress barely contacting the posterior of peripheral 3 and the inguinal buttress contacting the posterior of peripheral 7; relatively shorter femoral scute along the sagittal midline; thick overlap of plastral scutes (i.e. where the scute wraps onto the visceral surface of the plastron), particularly of the humeral and femoral scutes; anterior projection of the epiplastron beneath the gular resulting in a mid-length indent along the anterior plastral lobe rim; strongly scooped, or ventrally dipping medial portion of the epiplastra; inflation of posterior plastral lobe under femoral scutes; indent at lateral plastral edge of femoral scute-anal scute sulcus contact; flattened posterior edge of xiphiplastra; shortened fossa orbitalis; anteroposteriorly shortened and curved cavum tympani; pronounced depression immediately dorsal to processus articularis; wider angled lower jaws (86° in C. corniculata vs. 74–78° in modern Chrysemys); more pronounced sulcus cartilaginis meckelii; and more pronounced median ridge of the lower triturating surfaces. Description Methods: Terminology used throughout this study follows several well-known previous studies, including Thomson (1932), Zangerl (1969), Gaffney (1972), Ernst & Barbour (1989), Joyce & Bell (2004), Joyce (2007) and Jasinski (2018a). Although some previous studies have termed the keratinous, soft-tissue covering of the turtle shell as scales (e.g. Gaffney et al., 1998; Gilbert et al., 2001), we now know that the shell is covered with modified scales called scutes (e.g. Moustakas-Verho et al., 2014, 2017; Moustakas-Verho & Cherepanov, 2015) and therefore this latter term is used in the present study. Measurements are all maximum lengths and/ or widths unless otherwise stated. Orientations are in proper anatomical position unless otherwise stated. The electronic version of this article in Portable Document Format (PDF) will represent a published work according to the International Commission on Zoological Nomenclature (ICZN), and hence the new names contained in the electronic version are effectively published under that Code from the electronic edition alone. This published work and the nomenclatural acts it contains have been registered in ZooBank, the online registration system for the ICZN. The ZooBank LSIDs (Life Science Identifiers) can be resolved and the associated information viewed through any standard web browser by appending the LSID to the prefix http://zoobank.org/. Shell (Figs 2–3, 5): Chrysemys corniculata from the GFS is represented by multiple well preserved and mostly three-dimensional shells (including both carapaces and plastra). Although the specimens are sometimes somewhat crushed or ‘deformed’ while in situ, careful preparation often allows them to be re-assembled in their three-dimensional forms. The largest shells show maximum lengths (straight carapace length = SCL) of between 17.6 (ETMNH-12491) and 19.0 cm (ETMNH-20544), compared to C. picta, whose upper size (for females) is around 25.0 cm, although males achieve maximum size at around 15.3 cm (Ernst & Lovich, 2009). The shell is more oval than in modern Chrysemys (Fig. 4), which can be more rounded, although some individuals can be more oval, particularly in older individuals of modern Chrysemys. Carapace (Figs 2–3): There are multiple wellpreserved carapaces, either partial or nearly complete. These give a good indication of the general size, shape and characteristics of C. corniculata. ETMNH-12491 is a medium-sized individual, but one that has reached adulthood based on fusion of the shell (particularly carapacial) elements (e.g. Legler, 1960; Zug, 1991), measuring 17.6 cm sagittally (SCL). ETMNH-20544 is the largest individual recovered, represented by a nearly complete shell with a few cranial and postcranial fragments, and measuring 19.03 cm (SCL). ETMNH-20609 represents the smallest complete (to nearly complete) specimen, with a carapace length of 14.03 cm (SCL). The shell has fine, inconspicuous texturing, although this cannot be physically felt and can normally only been seen when held to low angles with light (Fig. 3 C-D). This inconspicuous texturing is present in modern Chrysemys, albeit even more subtle (Fig. 3 E-F), and is not present in some other fossil Chrysemys taxa such as Chrysemys antiqua Clark, 1937 and Chrysemys timida Hay, 1908. No median dorsal carapacial keel is present, and the posterior of the shell tends to have a sharp, downturned bend to it, making the posterior portion of the carapace nearly vertical (Fig. 3B). This downturn can be present in modern Chrysemys, although it is always at a shallower angle than in C. corniculata. As noted above, the carapace of C. corniculata is more oval (regardless of size) than modern C. picta, which tend to exhibit a relatively rounded or circular carapace in dorsal view (length/width ratios of 1.40–1.48 in C. corniculata compared to 1.29–1.36 in C. picta). In C. corniculata, the carapace also has a characteristic doming or raised portion at the anteromedial portion of the shell (Fig. 3A), meaning the nuchal is noticeably higher than the surrounding elements, mainly the peripherals and corresponding marginals lateral to it. This is most easily seen in anterior view, where modern Chrysemys are usually gently rounded, while there is an open angle between the bridge and the sagittal anterior midline. Simply put, it is analogous to ‘circular vs. triangular’ in anterior view. Additionally, the entire bridge is shifted posteriorly in C. corniculata. This is evidenced by the axillary buttress barely contacting the posterior of peripheral 3 in C. corniculata (Fig. 3B), while contacting the anterior of peripheral 3 in modern C. picta. For the inguinal buttress, both taxa contact the posterior of peripheral 7, indicating the bridge is a bit longer, larger and more robust in modern C. picta. Sutures of the carapace (Fig. 2A): The nuchal of C. corniculata represents a more pronounced and extreme morphology than that of modern Chrysemys. It maintains the roughly hexagonal shape of most emydid nuchals. The nuchal is nearly equal in length and width, although it is slightly longer than wide in ETMNH-12491 (42.4 mm long vs. 39.3 mm wide). Its anterior portion (anterior to the nuchal-peripheral 1-costal 1 intersection, 75% of nuchal length) is definitively longer than its posterior portion (25%). It is transversely constricted near the middle of the marginals 1, and flares anteriorly in C. corniculata, more so than in modern C. picta. The portion under the cervical scute does not reach as far anteriorly as that under the marginals 1, and these three projections are separated by two prevalent notches. The lateral edges of the anterolateral projections of the nuchal (nuchal horns) in C. corniculata are posterolaterally angled, making the anterior-most portion of the nuchal in the middle of these anterolateral projections, while there is no posterolateral-angling in other Chrysemys, including Chrysemys isoni Weems & George, 2013. As stated above, the doming or raised region at the anteromedial portion of the shell means the nuchal is noticeably higher than the rest of the surrounding elements, mainly the marginals lateral to it (Fig. 3A). This is most clearly seen in anterior view, where modern C. picta tend to be gently rounded (Fig. 4C), while the anterolateral edges of the carapace are angled dorsally to the nuchal (from the bridge). The majority of the eight neurals agree morphologically with those in modern Chrysemys. Neural 2 is squat in C. corniculata with a significantly rounded posterior border, while the posterior border in modern C. picta is straight. Neural 8 is more squared-off than modern C. picta, which tend to have a more ‘triangular’ neural 8, although the neurals in turtles tend to have minor quantitative variations (e.g. Cherepanov, 1994). The posterior suprapygal, immediately anterior to the pygal, is concave on its anterior border, with a wider than long hexagonal shape. The anterior border of the posterior suprapygal in modern C. picta tends to be straight (mediolaterally or transversely oriented), but can be slightly concave, although not to the same extent as C. corniculata. While in modern C. picta the pygal routinely has parallel lateral edges, in C. corniculata it is contracted posteriorly, being narrower posteriorly than anteriorly. The pygal also possesses a small but conspicuous notch posteromedially with sharp edges. While an inconspicuous notch can be found in some modern Chrysemys, the notch in the latter has gently sloping sides and is relatively smaller. The anterior edge of the pygal exhibits a gentle and inconspicuous concave curvature. The bridge peripherals of C. corniculata are vertical, making them appear narrow when viewed dorsally. This mostly agrees with Recent Chrysemys, although those in the modern taxa are not as extreme, and therefore not as inconspicuous when viewed dorsally. The other peripherals agree morphologically with other Chrysemys. The edge of the carapace is smooth, with only inconspicuous notches between the posterior peripherals of some individuals (e.g. ETMNH-3561, -20609), not including those notches on and immediately around the nuchal. The outer rim is also relatively gracile, especially in comparison to the other deirochelyines known from the GFS (Trachemys haugrudi; Jasinski, 2018a). The axillary buttress barely contacts the posterior of marginal 3 in C. corniculata (Fig. 3B), while it contacts the anterior of peripheral 3 in modern C. picta. The inguinal buttresses of both taxa usually contact the posterior of peripheral 7. This suggests a relatively shorter and potentially less robust bridge for C. corniculata. As noted above, the bridge peripherals are oriented mainly dorsoventrally, making them more visible laterally and less significant when viewed dorsally or ventrally. This leads to the posterior portion of the carapace flaring out laterally beginning midway through peripheral 7, as in modern Chrysemys. Sulci of the carapace (Fig. 2C): The bony shell of a turtle is covered by keratinous scales or plates called scutes. Scutes are modified scales and develop differently than reptilian scales, namely with the former developing from local epithelial thickenings called placodes through radial growth while scales (along with feathers, teeth and hair) develop as ectodermal appendages through epithelial-mesenchymal interaction (e.g. Cherepanov, 2006; Moustakas-Verho et al., 2014, 2017; Moustakas-Verho & Cherepanov, 2015). These scutes are separated into various different sections by sulci (or seams), which can be seen on the surface of the bones and give an indication of their morphology and appearance even when they are not present or preserved, such as in fossils. The cervical scute is long and narrow, often coming to an anterior point sagittally (medioanteriorly). The cervical underlap is shorter than the overlap in C. corniculata (~74–85%), although less so than in modern C. picta (~67–80%), but these values tend to vary, as seen by the range. The posterior width of the overlap of the cervical scute is narrower (compared to its length) in C. corniculata than either C. isoni or C. picta. The lateral edges of the cervical are parallel to slightly angled anteriorly as the cervical comes to an anterior point. As stated above, it is separated from marginals 1 by a pronounced lateral notch. The anterior-most point(s) of the carapace lie medially on the anterior projecting portion of marginals 1 (anterolateral projections of the nuchal under marginals 1). Although vertebrals 2–5 agree morphologically with those in C. picta, vertebral 1 is more morphologically unique (Fig. 3C). In C. corniculata, vertebral 1 is straight anteriorly and well-rounded (concave) posteriorly. There is lateral inflation posteriorly while it is constricted anteriorly. At its anterolateral edges vertebral 1 flares laterally and becomes significantly wider, giving it a slightly hourglass-like shape. However, in modern C. picta, the anterior and posterior borders of vertebral 1 tend to be relatively straight to slightly curved, while the anterior half is normally wider (the sides can also sometimes be approximately parallel) (Fig. 3E). Regardless, the only unique curvature present is at the anterolateral points, which curve slightly laterally. Vertebral 4 also exhibits more curvature and has more sinusoidal sulci, although this is mainly visible posterolaterally where it contacts pleural 4. As in other Chrysemys, and emydids in general, C. corniculata has four pairs of pleurals that mostly agree morphologically with those of modern Chrysemys. However, as for the morphology of the vertebrals, the medial sulci of pleurals 1 and 4 in C. corniculata are more sinuous and curved where they contact vertebrals 1 and 4, respectively. As noted above, marginal 1 has a pronounced medial projection, clearly distinguishing its anteromedial and anterolateral halves (Figs 2, 7 A-D). The anterior e
Published as part of Jasinski, Steven E., 2023, A new species of Chrysemys (Emydidae: Deirochelyinae) from the latest Miocene-Early Pliocene of Tennessee, USA and its implications for the evolution of painted turtles, pp. 149-183 in Zoological Journal of the Linnean Society 198 on pages 152-166, DOI: 10.1093/zoolinnean/zlac084, http://zenodo.org/record/7924771
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