taxonID	type	description	language	source
523F87B8FF91DC5BFC40FE6AE308FE74.taxon	materials_examined	TYPE SPECIES. — Ptychogaster emydoides Pomel, 1847.	en	Luján, Àngel H., Čerňanský, Andrej, Bonilla-Salomón, Isaac, Březina, Jakub, Ivanov, Martin (2021): Fossil turtles from the early Miocene localities of Mokrá-Quarry (Burdigalian, MN 4), South Moravian Region, Czech Republic. Geodiversitas 43 (20): 691-707, DOI: 10.5252/geodiversitas2021v43a20
523F87B8FF91DC57FC75FDCAE539F911.taxon	description	(Figs 2; 3) LOCALITIES. — MWQ 1 / 2001, MWQ 2 / 2003, MWQTC / 2001, MWQ 4 / 2018 and MCQ 3 / 2005. STUDIED MATERIAL. — Czech Republic. South Moravia Region, Mokrá-Quarry, carapace remains (Fig. 2 A-R’): Pal. 1300, nuchal and left peripherals 1 - 3; Pal. 1301, nuchal; Pal. 1302, neural 4; Pal. 1303, neural 5; Pal. 1304, neural 7; Pal. 1305, suprapygal 1; Pal. 1306, suprapygal 2; Pal. 1307, left costal 1; Pal. 1308, right costal 6; Pal. 1309, right costal 6; Pal. 1310, left costal 8; Pal. 1311, right costal 8; Pal. 1312, right peripheral 1; Pal. 1313, left peripheral 7; Pal. 1314, left peripheral 8. Plastral remains (Fig. 3 A-R): Pal. 1315, left hypoplastron; Pal. 1316, right hypoplastron; Pal. 1317, right hypoplastron; Pal. 1318, left xiphiplastron. DESCRIPTION The ptychogasterid material (Pal. 1300 - 1318) from Mokrá- Quarry consists of numerous disarticulated plates, which present preserved parts of the almost the entire shell. However, some plates are missing from the carapace (i. e., pygal, costals 2 - 4 and 7, neurals 1 - 3 and both medial and posterior peripherals). The plastron is represented only by the hypoplastron and the xiphiplastron. The following description is based on all available material, however, not all preserved plates are depicted in Figures 2 and 3. According to the dimensions of the nuchal plate (Pal. 1300 - 1301), costals (Pal. 1307 - 1311) and peripherals (Pal. 1312 - 1314), the shell would have been relatively large (> 20 cm: Figs 2; 3). Ornamentation of both carapace and plastron external surfaces is absent. As it is typical for Ptychogaster plates, dermal grooves are well developed. Unlike Testudo, neither growth lines nor sutures (i. e., completely fused to each other: see e. g., Fig. 2 A-D) are preserved, so it is impossible to evaluate them. The nuchal plate is hexagonal in outline, wider than long (Pal. 1301: Fig. 2 E-H). It contacts the first pair of peripherals, the first pair of costals and the neural 1. The anterior border possesses a shallow nuchal notch, affecting the nuchal border and the first peripherals (Pal. 1300: Fig. 2 A, B). The anteroposterolateral sides of the nuchal are rather equal in length, whereas the posterior border is narrow and slightly convex anteriorly (Fig. 2 E, F). In lateral view, the nuchal is vaguely curved (Fig. 2 I, J). Two transversal thickenings are developed on the internal surface of this bone (Fig. 2 C, D, 2 G-L). The cervical scute is present anteriorly, both dorsally and viscerally. It is a relatively large and trapezoidal element that is longer than wide (Fig. 2 E, F). The lateral edges of the cervical are slightly curved medially both in dorsal and visceral sides (Fig. 2 A, B, E-H). The overlap of this scute is less developed on the ventral surface (Fig. 2 G, H). According to the preserved neural plates, an alternating between octagonal and hexagonal plates forms the neural series: neural 4 octagonal (Pal. 1302: Fig. 2 M, N) and neural 5 and 7 hexagonal with short sides behind (Pal. 1303 - 1304: Fig. 2 O-S). A weak medial keel develops on the neural 7 (Pal. 1304: Fig. 2 R, S) and the suprapygals 1 - 2 (Pal. 1305 - 1306: Fig. 2 T, U). In visceral view, remains of the thoracic vertebrae attachments are present in all neural plates (Fig. 2 N, Q, S). The suprapygal plate 1 is trapezoidal with a wider posterior part (Pal. 1305: Fig. 2 T). The anterior border is concave, whereas the posterior one is slightly convex. It contacts the neural 8 anteriorly, costals 8 laterally and suprapygal 2 posteriorly. Suprapygal 2 is hexagonal, wider than long, and much wider than suprapygal 1 (Fig. 2 T, U). It contacts the suprapygal 1 and the posteromedial sides of costals 8 anteriorly, peripherals 11 laterally and pygal posteriorly. The anterolateral sides of the suprapygal 2 are slightly longer anteroposteriorly, compared to the posterolateral ones. The posterior side is vaguely convex anteriorly (Fig. 2 T, U). The vertebral scute series is partially preserved, which is quadrangular and slightly narrower than the costal series. Vertebral 1 contacts the cervical and marginals 1 - 2 anteriorly (Fig. 2 B, F). It seems to be lyre-shaped and covers the lateral corners of the nuchal and costals 1 (Pal. 1300 - 1301: Fig. 2 A, B, E, F). According to preserved portion of the vertebral 3, it expanded at least on costals 5 and neural 5 (Fig. 2 X, Y). The sulcus between the vertebrals 3 - 4 is wavy in its medial part, and more specifically in the part that is crossing the neural 5 (Fig. 2 O, P). Vertebral 4 likely contacts the vertebral 3 anteriorly, pleurals 3 - 4 laterally and vertebral 5 posteriorly (Fig. 2 P, A’, B’, D’, E’). Vertebral 5 is the widest vertebral scute, contacting with the vertebral 4 anteriorly, pleural 4 anterolaterally and marginals 11 - 12 posteriorly. It expands on costals 8, neural 8, peripheral 12 and pygal, and therefore covers the entire surface of the suprapygals 1 - 2. Although not fully preserved costals, the costal plate 1 is much longer than the rest of costal (Pal. 1307: Fig. 2 V, W). It is trapezoidal and always contacts the peripheral plates 1 - 3 anterolaterally, nuchal anteromedially and neurals 1 - 2 medially. The anterior border is sinuous to articulate with the corresponding peripherals. Costal 6 is similar in regards of its shape, being much wider than long (Pal. 1308 - 9: Fig. 2 X-B’). The medial side of Pal. 1308 shows a short anteromedial and long posteromedial sides (Fig. 2 X-Z). However, the medial side of Pal. 1308 is most likely rounded (Fig. 2 A’ - C’). Costal 8 is narrow, slightly wider than long, contacting the costal 7 anteriorly, neural 8 and suprapygal 1 medially, and suprapygal 2 and peripherals 11 - 12 posteriorly (Pal. 1310 - 11: Fig. 2 D’ - F’). The pleural scutes are not preserved with the exception of the first one. The marginopleural sulcus is entirely situated on peripheral plates, at least in both anterior and posterior part of the carapace (Fig. 2 A, B, E, F, V, A’, B’, D’, E’). The peripheral plates 1 - 3 are longer than wide and slightly trapezoidal (Fig. 2 A-D, G’ - J’). They are completely fused together (i. e., sutures are not visible) and crossed not only by the intermarginal sulcus, but also by the pleuro-marginal sulcus, unlike in Testudo. Peripherals 1 - 2 are prominent in anterior direction in Pal. 1300. Peripheral 7, which is partially preserved, is rectangular (Pal. 1313: Fig. 2 K’ - N’). In internal view, it displays a rough elongated area for the cartilaginous union of the inguinal process (Fig. 2 M’, N’). Pal. 1313 is the last peripheral involved in the shell bridge and also displays a weak lateral ridge on its external side. Peripheral 8 is rectangular and hosts both the pleuromarginal sulcus and intermarginal sulci (i. e., between marginals 8 - 9), which are situated far from the costoperipheral suture (Pal. 1314: Fig. 2 O’ - R’). The marginal scute 1 is rectangular, slightly wider than long, whereas marginal 2 is trapezoidal. Marginal 3 is approximately as wide as long. Marginal 8, the only complete scute from the bridge area, is rectangular and higher than wide. The ventral overlap of all preserved marginals is well developed. The preserved portion of the hyo-hypoplastral hinge is straight and approximately transversal (Pal. 1377: Fig. 3 M-P), which is located at the mid of the peripheral 6. The hypoplastron contact anteriorly with the hypoplastron through a hinge and laterally with the peripherals 6 - 7 through a completely ligamentous union in the inguinal process (Pal. 1315 - 17: Fig. 3 A-R). The visceral overlap of both abdominal and femoral scutes on hypoplastra is well developed (Fig. 3 M, N). The xiphiplastron is trapezoidal and its posterior tip is rather rounded. The abdominofemoral sulcus, developed on the ventral surface of each hypoplastron, is concave (Fig. 2 N, O). Moreover, the latter does not reach the inguinal process laterally, but is located slightly below it. The femoroanal sulcus is oblique, whereas the anal scute is triangular with rounded lateral borders (Fig. 2 S-V). The dorsal overlap of the anal scute is moderately developed (Fig. 2 S, T).	en	Luján, Àngel H., Čerňanský, Andrej, Bonilla-Salomón, Isaac, Březina, Jakub, Ivanov, Martin (2021): Fossil turtles from the early Miocene localities of Mokrá-Quarry (Burdigalian, MN 4), South Moravian Region, Czech Republic. Geodiversitas 43 (20): 691-707, DOI: 10.5252/geodiversitas2021v43a20
523F87B8FF91DC57FC75FDCAE539F911.taxon	discussion	REMARKS	en	Luján, Àngel H., Čerňanský, Andrej, Bonilla-Salomón, Isaac, Březina, Jakub, Ivanov, Martin (2021): Fossil turtles from the early Miocene localities of Mokrá-Quarry (Burdigalian, MN 4), South Moravian Region, Czech Republic. Geodiversitas 43 (20): 691-707, DOI: 10.5252/geodiversitas2021v43a20
523F87B8FF9DDC54FE95F8EEE74CFAB2.taxon	description	LOCALITIES. — MWQ 4 / 2018. STUDIED MATERIAL. — Czech Republic. South Moravia Region, Mokrá-Quarry, shell and postcranial remains (Fig. 4 A-M): Pal. 1363, shell fragment; Pal. 1364, scapula (i. e., anterodorsal process fragment); Pal. 1365, right fibula. DESCRIPTION Only a shell fragment formed by two portions of plates is known (Fig. 4 A-D). Pal. 1363 corresponds most likely to a carapace portion, but it is poorly preserved and is not possible to assess this confidently. The length and maximum width of the preserved plate fragment is 6 cm. The external part is completely smooth and is not crossed by any sulcus (Fig. 4 C, D). A suture is recognized on top of plate, which is concave and approximately 2 cm wide. A partial bone, belonging to the shoulder girdle, has been identified (Pal. 1364: Fig. 4 E-H). Only the anterodorsal process fragment is preserved. Pal. 1364 is subcylindrical in cross-section and distally rounded. The distal surface ends in a rough rounded area to join with the visceral part of the carapace (Fig. 4 H). The hind limb skeleton is restricted to one partial fibula (Fig. 4 I-M) that is elliptical in cross-section. Its distal articular surface is slightly small, oval and convex (Fig. 4 M). Both postcranial bones are poorly preserved and no significant details can be discerned.	en	Luján, Àngel H., Čerňanský, Andrej, Bonilla-Salomón, Isaac, Březina, Jakub, Ivanov, Martin (2021): Fossil turtles from the early Miocene localities of Mokrá-Quarry (Burdigalian, MN 4), South Moravian Region, Czech Republic. Geodiversitas 43 (20): 691-707, DOI: 10.5252/geodiversitas2021v43a20
523F87B8FF9DDC54FE95F8EEE74CFAB2.taxon	discussion	REMARKS Fossil remains of giant tortoises are not very common in Miocene assemblages of Central Europe; their record being limited to few localities from Austria, Germany, Hungary and Switzerland (Alba et al. 2010, 2011; Carmona et al. 2011; Luján 2015). Loveridge & Williams (1957) proposed that all European giant tortoises should be transferred into the extant genus Geochelone. This proposal was adopted for some time, and consequently, large tortoise remains in Europe are still frequently referred to in the literature as Geochelone sp. (e. g., Auffenberg 1974; Młynarski 1976). However, current phylogenies do not support a close relationship between Mio- Pleistocene large tortoises and Geochelone. More recently, Bourgat & Bour (1983) referred all giant fossil tortoises to the genus Cheirogaster. Most subsequent works accepted this genus attribution (e. g., Luján et al. 2010, 2014), until recently when Pérez-García & Vlachos (2014) proposed that European Neogene giant tortoises constitute a clade that is more derived than the type species of Cheirogaster. To allocate these taxa, Pérez-García & Vlachos (2014) erected the genus Titanochelon, with Ti. bolivari (Hernández- Pacheco, 1917) as its type species. This genus is characterized by a shell reaching over 100 cm and the fusion of marginal scutes 12 (i. e., constituting a supracaudal scute). However, the evolution of gigantism amongst fossil tortoises is clearly a homoplastic phenomenon, mainly related to insular conditions, or adaptation to either global or local environmental changes (Kear 2010; Luján et al. 2010, 2017 b; Itescu et al. 2014). Similarly, the fusion of marginal scutes 12 occurs in many extant and extinct genera and cannot be considered autapomorphic for the genus Titanochelon. In summary, the taxonomy of the Miocene giant tortoises of Europe is still a subject of debate and will require improvement of existing data matrix (e. g., including more skull characters) in order to decipher the phylogenetic relationships of Titanochelon (Luján et al. 2017 b).	en	Luján, Àngel H., Čerňanský, Andrej, Bonilla-Salomón, Isaac, Březina, Jakub, Ivanov, Martin (2021): Fossil turtles from the early Miocene localities of Mokrá-Quarry (Burdigalian, MN 4), South Moravian Region, Czech Republic. Geodiversitas 43 (20): 691-707, DOI: 10.5252/geodiversitas2021v43a20
523F87B8FF9EDC55FF09F9A4E31CF8D0.taxon	description	(Figs 5; 6) LOCALITIES. — MWQ 1 / 2001, MWQ 2 / 2003, MWQTC / 2001, MWQ 4 / 2018 and MCQ 3 / 2005. STUDIED MATERIAL. — Czech Republic. South Moravia Region, Mokrá-Quarry, carapace remains (Fig. 5 A-V’): Pal. 1319, nuchal; Pal. 1320, nuchal; Pal. 1321, nuchal; Pal. 1322, neural 4; Pal. 1323, neural 6; Pal. 1350, neural 7; Pal. 1322, neural 4; Pal. 1324, pygal; Pal. 1325, pygal; Pal. 1328, right costal 1; Pal. 1326, left costal 1; Pal. 1327, left costal 1; Pal. 1329, right costal 2; Pal. 1330, left costal 3; Pal. 1331, left costal 6; Pal. 1332, left peripheral 1; Pal. 1333, right peripheral 1; Pal. 1334, left peripheral 2; Pal. 1335, left peripheral 2; Pal. 1336, right periph- eral 2; Pal. 1337, right peripheral 2; Pal. 1338, right peripheral 5; Pal. 1339, left peripheral 7; Pal. 1340, right peripheral 7; Pal. 1341, right peripheral 8; Pal. 1342, left peripheral 9; Pal. 1343, right peripheral 9; Pal. 1344, left peripheral 10; Pal. 1345, left peripheral 10; Pal. 1346, left peripheral 10; Pal. 1347, left peripheral 11; Pal. 1348, left peripheral 11. Pal. 1349, left peripheral 11. Plastral remains (Fig. 6 A-H’): Pal. 1351 A, left epiplastron; Pal. 1352, entoplastron; Pal. 1353, entoplastron; Pal. 1354, entoplastron; Pal. 1351 B, right hyoplastron; Pal. 1355, right hyoplastron; Pal. 1356, left hypoplastron; Pal. 1357, right hypoplastron; Pal. 1358, right hypoplastron; Pal. 1359, right hypoplastron; Pal. 1360, left xiphiplastron; Pal. 1361, left xiphiplastron; Pal. 1362, left xiphiplastron. DESCRIPTION Testudo material recovered from Mokrá-Quarry consists mainly of disarticulated plates (Pal. 1319 - 1362). However, some costals, peripherals and neurals are missing. They belong to a medium-sized Testudo, which reached approximately 20 - 25 cm. The following description is based on the entire available material, but not all the plates are depicted (Figs 5; 6). Most of the dermal grooves and sutures are visible. There is no ornamentation on the carapace or plastron. Growth lines are discernible in some carapace plates, and more specifically in both peripheral and costal plates (e. g., Fig. 5 K, A’). The nuchal plate is hexagonal, slightly wider than long, and its anterior edge is pointed (Fig. 5 A-D). The posterolateral edges are straight, whereas the anterior ones are curved in medial direction. A transverse thickening is recognizable on the visceral surface of the nuchal (Fig. 5 C, D). The cervical scute is longer than wide and its total length constitutes less than a half of the nuchal plate. It is well developed both dorsally and viscerally according to the three preserved plates (Pal. 1319 - 21). The anterior edge is narrower than the posterior ones. The lateral sulci are almost straight and parallel to each other (Fig. 5 A, B). Only three neurals plates are preserved, which vary in shape from subsquare (i. e., neural 4) to hexagonal (i. e., neurals 6 to 7: Fig. 5 E-G). It is noteworthy that none of them is more than twice as wide as long, and that they are encroached transversally by the intervertebral sulci (Fig. 5 E-G). The pygal plate is trapezoidal with slightly concave anterolateral margins. Its external surface is moderately convex, whereas its internal one is rather concave (Fig. 5 H-J). The marginal scutes 12 are missing, and therefore the supracaudal scute is not divided by a sagittal groove (Pal. 1324 - 25: Fig. 5 H-J). The shape of the vertebral scutes cannot be ascertained with a confidence because they are incomplete. Only five of the eight costal plates are present (i. e., costals 1 - 3 and 5 - 6), which are trapezoidal. Costals 1 and 3 host the intervertebral sulci (Fig. 5 K, L, O, P, W’), whereas the costals 2 and 6 host the interpleural sulci (Fig. 5 M, N, Q, R, W’). The pleural scutes are poorly preserved, so no significant details can be discerned. Despite this, the pleuromarginal sulcus coincides with the costoperipheral suture all along the preserved peripheral and costal plates (e. g., Fig. 5 K, L, W’). Peripheral plates 1 - 3, together with the nuchal, make up the anterior opening of the shell. Peripheral 1 is heptagonal, while the peripheral 2 is subtriangular. Peripheral 1 displays a moderately developed spike at about the middle of the anterior edge. The presence of protrusions on the remaining peripherals (i. e., 2 and 3) cannot be evaluated. Peripheral 5 (Pal. 1338: Fig. 5 A’, B’), which is the only preserved plate involved in the shell bridge, is rectangular and rather flat, unlike posterior peripherals. Pal. 1338 displays a very weak longitudinal lateral ridge that is placed slightly above of the marginoabdominal sulcus. Peripherals 7 - 11 (Fig. 5 C’ - W’), together with the pygal plate (Fig. 5 H-J), form the posterior opening of the shell. An elongated and subvertical scar of the dorsal projection of the hypoplastron is discernible in peripheral 7 internally (Fig. 5 E’, F’). As a rule, the peripherals 8 - 10 in Testudo are rectangular with dorsal surfaces slightly concave, whereas in Ptychogaster, they are rather subrectangular and the dorsal concavity is well-developed. Peripheral 11 hosts the lateral edge of the supracaudal scute (Fig. 5 S’ - W’). The marginal scute 1 is trapezoidal, whereas the remaining posterior marginals are either subsquare or subrectangular (Fig. 5 A-D, S-V, W’). Based on two nuchal plates (Pal. 1319 - 21), two peripherals 1 (Pal. 1332 - 33) and three costals 1 (Pal. 1326 - 28), the triple junction amongst the pleural 1, vertebral 1, and marginal 1 is located outside the nuchal plate. Noticeably, the posterior border of the marginal 5 (Pal. 1338; Fig. 5 A’, B’) is parallel (i. e., instead of oblique) relative to those anteroposterior edges of the peripheral 5. A partial epiplastron, together with the hyoplastron (Fig. 6 A-F, O-R), forms the anterior plastral lobe, which is rather trapezoidal (Fig. 6 I’). Despite not being entirely preserved, the epiplastral dorsal pad seems rectangular and longer than wide. It is moderately developed posteroventrally and overhangs it slightly. Consequently, a small gular pocket is present (Fig. 6 A-C). The gular scutes are triangular, with slightly sinuous lateral margins, and form an angle of less than 45 ° relative to the sagittal axis (Fig. 6 B-D). The three available entoplastra (Pal. 1352 - 54: Fig. 6 G-N) are hexagonal both ventrally and viscerally. They are partially covered by the gular scutes, which generally extend up to the middle of the entoplastron. The ventral surface of the epiplastra, covered by the gular scutes, is not in relief. In all specimens, the gular scutes are not crossed by the humeropectoral sulcus transversally (Fig. 6 H, I, K, L). Pal. 1351 B is the best preserved hyoplastron (Fig. 6 O-R), which hosts entirely both the humeropectoral and pectoroabdominal scutes: the former is nearly straight and obliquely oriented relative to the sagittal plane (i. e., only slightly sinuous: Fig. 6 R), while the latter is curved and transversally oriented relative to the sagittal plane (Fig. 6 R, V). The preserved portion of the humeral scute indicates that this was trapezoidal. The abdominal seems to be the largest scute of the plastron. Moreover, its medial sector is slightly oblique (Fig. 6 Y-Z). The hypo-xiphiplastral suture is roughly straight and well developed, which means that a plastral hinge is absent. The xiphiplastron is trapezoidal and its ventral side is very flat and crossed by the femoroanal sulcus (Fig. 6 A’ - C’). It is noteworthy that the distinct notch in their lateral margins, between the anal and femoral scutes, is missing (Fig. 6 A’, C’). The femoral is trapezoidal and much longer medially than the anal scutes, which are subrectangular and wider than long. The femoroanal sulcus is slightly sinuous and obliquely oriented relative to the sagittal plane. The anal notch is wider than long, and its visceral area covered by the anal scutes is variable, from moderately (Pal. 1361: Fig. 6 A’, B’) to well developed (Pal. 1362: Fig. 6 E’, F’).	en	Luján, Àngel H., Čerňanský, Andrej, Bonilla-Salomón, Isaac, Březina, Jakub, Ivanov, Martin (2021): Fossil turtles from the early Miocene localities of Mokrá-Quarry (Burdigalian, MN 4), South Moravian Region, Czech Republic. Geodiversitas 43 (20): 691-707, DOI: 10.5252/geodiversitas2021v43a20
523F87B8FF9EDC55FF09F9A4E31CF8D0.taxon	discussion	REMARKS The genus Testudo s. l. is a clade with five extant and multiple extinct species of terrestrial tortoises of western Palearctic distribution (Lapparent de Broin et al. 2006 a, b; Fritz & Bininda-Emonds 2007; Corsini et al. 2014; Delfino et al. 2012; Luján et al. 2016). Despite that, in the past, three extant genus-group taxa were distinguished as genera (see e. g., Turtle Taxonomy Working Group 2014), currently, the use of a single genus (Testudo s. l.) and three subgenera (Testudo, Agrionemys and Chersine) is being better accepted (see Luján et al. 2016; Graciá et al. 2017; Turtle Taxonomy Working Group 2017). Among extinct taxa, the taxonomy of Testudo s. l. also included the distinction of the extinct genus Paleotestudo Lapparent de Broin, 2000. Although the latter has not been the object of an exhaustive review, results presented by Luján et al. (2016) clearly pointed out that Paleotestudo is a junior subjective synonym of subgenus Chersine. Regarding the species Testudo kalksburgensis Toula, 1896, it was originally described by Toula (1896) based on one specimen from Kalksburg, Vienna: a partial shell currently housed at the IGUW. At the beginning of the twentieth century, new fragmentary material, coming from the late Miocene locality of Au am Leithaberge, was referred to T. kalksburgensis by Siebenrock (1914). Shortly after, Staesche (1931) also erected Testudo kalksburgensis var. steinheimensis based on various specimens from the middle Miocene (MN 7) locality of Steinheim (Germany). Although T. kalksburgensis was considered a junior subjective synonym of T. antiqua Bronn, 1831 by Glaessner (1933), subsequently Młynarski (1976) resurrected the species once again. Indeed, the validity of T. kalksburgensis was also confirmed by latter studies (see Młynarski 1980; Bachmayer & Młynarski 1981; Schleich 1981; Gemel & Rauscher 2000; Gemel 2002; Danilov et al. 2012; Luján et al. 2016; Březina et al. 2021).	en	Luján, Àngel H., Čerňanský, Andrej, Bonilla-Salomón, Isaac, Březina, Jakub, Ivanov, Martin (2021): Fossil turtles from the early Miocene localities of Mokrá-Quarry (Burdigalian, MN 4), South Moravian Region, Czech Republic. Geodiversitas 43 (20): 691-707, DOI: 10.5252/geodiversitas2021v43a20
523F87B8FF9EDC54FEAFFA4AE4CEF995.taxon	materials_examined	TYPE SPECIES. — Testudo (Chersine) hermanni Gmelin, 1789. Testudo (Chersine) cf. kalksburgensis Toula, 1896 (Figs 5; 6) LOCALITIES. — MWQ 1 / 2001, MWQ 2 / 2003, MWQTC / 2001, MWQ 4 / 2018 and MCQ 3 / 2005. STUDIED MATERIAL. — Czech Republic. South Moravia Region, Mokrá-Quarry, carapace remains (Fig. 5 A-V’): Pal. 1319, nuchal; Pal. 1320, nuchal; Pal. 1321, nuchal; Pal. 1322, neural 4; Pal. 1323, neural 6; Pal. 1350, neural 7; Pal. 1322, neural 4; Pal. 1324, pygal; Pal. 1325, pygal; Pal. 1328, right costal 1; Pal. 1326, left costal 1; Pal. 1327, left costal 1; Pal. 1329, right costal 2; Pal. 1330, left costal 3; Pal. 1331, left costal 6; Pal. 1332, left peripheral 1; Pal. 1333, right peripheral 1; Pal. 1334, left peripheral 2; Pal. 1335, left peripheral 2; Pal. 1336, right periph- eral 2; Pal. 1337, right peripheral 2; Pal. 1338, right peripheral 5; Pal. 1339, left peripheral 7; Pal. 1340, right peripheral 7; Pal. 1341, right peripheral 8; Pal. 1342, left peripheral 9; Pal. 1343, right peripheral 9; Pal. 1344, left peripheral 10; Pal. 1345, left peripheral 10; Pal. 1346, left peripheral 10; Pal. 1347, left peripheral 11; Pal. 1348, left peripheral 11. Pal. 1349, left peripheral 11. Plastral remains (Fig. 6 A-H’): Pal. 1351 A, left epiplastron; Pal. 1352, entoplastron; Pal. 1353, entoplastron; Pal. 1354, entoplastron; Pal. 1351 B, right hyoplastron; Pal. 1355, right hyoplastron; Pal. 1356, left hypoplastron; Pal. 1357, right hypoplastron; Pal. 1358, right hypoplastron; Pal. 1359, right hypoplastron; Pal. 1360, left xiphiplastron; Pal. 1361, left xiphiplastron; Pal. 1362, left xiphiplastron. DESCRIPTION Testudo material recovered from Mokrá-Quarry consists mainly of disarticulated plates (Pal. 1319 - 1362). However, some costals, peripherals and neurals are missing. They belong to a medium-sized Testudo, which reached approximately 20 - 25 cm. The following description is based on the entire available material, but not all the plates are depicted (Figs 5; 6). Most of the dermal grooves and sutures are visible. There is no ornamentation on the carapace or plastron. Growth lines are discernible in some carapace plates, and more specifically in both peripheral and costal plates (e. g., Fig. 5 K, A’). The nuchal plate is hexagonal, slightly wider than long, and its anterior edge is pointed (Fig. 5 A-D). The posterolateral edges are straight, whereas the anterior ones are curved in medial direction. A transverse thickening is recognizable on the visceral surface of the nuchal (Fig. 5 C, D). The cervical scute is longer than wide and its total length constitutes less than a half of the nuchal plate. It is well developed both dorsally and viscerally according to the three preserved plates (Pal. 1319 - 21). The anterior edge is narrower than the posterior ones. The lateral sulci are almost straight and parallel to each other (Fig. 5 A, B). Only three neurals plates are preserved, which vary in shape from subsquare (i. e., neural 4) to hexagonal (i. e., neurals 6 to 7: Fig. 5 E-G). It is noteworthy that none of them is more than twice as wide as long, and that they are encroached transversally by the intervertebral sulci (Fig. 5 E-G). The pygal plate is trapezoidal with slightly concave anterolateral margins. Its external surface is moderately convex, whereas its internal one is rather concave (Fig. 5 H-J). The marginal scutes 12 are missing, and therefore the supracaudal scute is not divided by a sagittal groove (Pal. 1324 - 25: Fig. 5 H-J). The shape of the vertebral scutes cannot be ascertained with a confidence because they are incomplete. Only five of the eight costal plates are present (i. e., costals 1 - 3 and 5 - 6), which are trapezoidal. Costals 1 and 3 host the intervertebral sulci (Fig. 5 K, L, O, P, W’), whereas the costals 2 and 6 host the interpleural sulci (Fig. 5 M, N, Q, R, W’). The pleural scutes are poorly preserved, so no significant details can be discerned. Despite this, the pleuromarginal sulcus coincides with the costoperipheral suture all along the preserved peripheral and costal plates (e. g., Fig. 5 K, L, W’). Peripheral plates 1 - 3, together with the nuchal, make up the anterior opening of the shell. Peripheral 1 is heptagonal, while the peripheral 2 is subtriangular. Peripheral 1 displays a moderately developed spike at about the middle of the anterior edge. The presence of protrusions on the remaining peripherals (i. e., 2 and 3) cannot be evaluated. Peripheral 5 (Pal. 1338: Fig. 5 A’, B’), which is the only preserved plate involved in the shell bridge, is rectangular and rather flat, unlike posterior peripherals. Pal. 1338 displays a very weak longitudinal lateral ridge that is placed slightly above of the marginoabdominal sulcus. Peripherals 7 - 11 (Fig. 5 C’ - W’), together with the pygal plate (Fig. 5 H-J), form the posterior opening of the shell. An elongated and subvertical scar of the dorsal projection of the hypoplastron is discernible in peripheral 7 internally (Fig. 5 E’, F’). As a rule, the peripherals 8 - 10 in Testudo are rectangular with dorsal surfaces slightly concave, whereas in Ptychogaster, they are rather subrectangular and the dorsal concavity is well-developed. Peripheral 11 hosts the lateral edge of the supracaudal scute (Fig. 5 S’ - W’). The marginal scute 1 is trapezoidal, whereas the remaining posterior marginals are either subsquare or subrectangular (Fig. 5 A-D, S-V, W’). Based on two nuchal plates (Pal. 1319 - 21), two peripherals 1 (Pal. 1332 - 33) and three costals 1 (Pal. 1326 - 28), the triple junction amongst the pleural 1, vertebral 1, and marginal 1 is located outside the nuchal plate. Noticeably, the posterior border of the marginal 5 (Pal. 1338; Fig. 5 A’, B’) is parallel (i. e., instead of oblique) relative to those anteroposterior edges of the peripheral 5. A partial epiplastron, together with the hyoplastron (Fig. 6 A-F, O-R), forms the anterior plastral lobe, which is rather trapezoidal (Fig. 6 I’). Despite not being entirely preserved, the epiplastral dorsal pad seems rectangular and longer than wide. It is moderately developed posteroventrally and overhangs it slightly. Consequently, a small gular pocket is present (Fig. 6 A-C). The gular scutes are triangular, with slightly sinuous lateral margins, and form an angle of less than 45 ° relative to the sagittal axis (Fig. 6 B-D). The three available entoplastra (Pal. 1352 - 54: Fig. 6 G-N) are hexagonal both ventrally and viscerally. They are partially covered by the gular scutes, which generally extend up to the middle of the entoplastron. The ventral surface of the epiplastra, covered by the gular scutes, is not in relief. In all specimens, the gular scutes are not crossed by the humeropectoral sulcus transversally (Fig. 6 H, I, K, L). Pal. 1351 B is the best preserved hyoplastron (Fig. 6 O-R), which hosts entirely both the humeropectoral and pectoroabdominal scutes: the former is nearly straight and obliquely oriented relative to the sagittal plane (i. e., only slightly sinuous: Fig. 6 R), while the latter is curved and transversally oriented relative to the sagittal plane (Fig. 6 R, V). The preserved portion of the humeral scute indicates that this was trapezoidal. The abdominal seems to be the largest scute of the plastron. Moreover, its medial sector is slightly oblique (Fig. 6 Y-Z). The hypo-xiphiplastral suture is roughly straight and well developed, which means that a plastral hinge is absent. The xiphiplastron is trapezoidal and its ventral side is very flat and crossed by the femoroanal sulcus (Fig. 6 A’ - C’). It is noteworthy that the distinct notch in their lateral margins, between the anal and femoral scutes, is missing (Fig. 6 A’, C’). The femoral is trapezoidal and much longer medially than the anal scutes, which are subrectangular and wider than long. The femoroanal sulcus is slightly sinuous and obliquely oriented relative to the sagittal plane. The anal notch is wider than long, and its visceral area covered by the anal scutes is variable, from moderately (Pal. 1361: Fig. 6 A’, B’) to well developed (Pal. 1362: Fig. 6 E’, F’). REMARKS The genus Testudo s. l. is a clade with five extant and multiple extinct species of terrestrial tortoises of western Palearctic distribution (Lapparent de Broin et al. 2006 a, b; Fritz & Bininda-Emonds 2007; Corsini et al. 2014; Delfino et al. 2012; Luján et al. 2016). Despite that, in the past, three extant genus-group taxa were distinguished as genera (see e. g., Turtle Taxonomy Working Group 2014), currently, the use of a single genus (Testudo s. l.) and three subgenera (Testudo, Agrionemys and Chersine) is being better accepted (see Luján et al. 2016; Graciá et al. 2017; Turtle Taxonomy Working Group 2017). Among extinct taxa, the taxonomy of Testudo s. l. also included the distinction of the extinct genus Paleotestudo Lapparent de Broin, 2000. Although the latter has not been the object of an exhaustive review, results presented by Luján et al. (2016) clearly pointed out that Paleotestudo is a junior subjective synonym of subgenus Chersine. Regarding the species Testudo kalksburgensis Toula, 1896, it was originally described by Toula (1896) based on one specimen from Kalksburg, Vienna: a partial shell currently housed at the IGUW. At the beginning of the twentieth century, new fragmentary material, coming from the late Miocene locality of Au am Leithaberge, was referred to T. kalksburgensis by Siebenrock (1914). Shortly after, Staesche (1931) also erected Testudo kalksburgensis var. steinheimensis based on various specimens from the middle Miocene (MN 7) locality of Steinheim (Germany). Although T. kalksburgensis was considered a junior subjective synonym of T. antiqua Bronn, 1831 by Glaessner (1933), subsequently Młynarski (1976) resurrected the species once again. Indeed, the validity of T. kalksburgensis was also confirmed by latter studies (see Młynarski 1980; Bachmayer & Młynarski 1981; Schleich 1981; Gemel & Rauscher 2000; Gemel 2002; Danilov et al. 2012; Luján et al. 2016; Březina et al. 2021).	en	Luján, Àngel H., Čerňanský, Andrej, Bonilla-Salomón, Isaac, Březina, Jakub, Ivanov, Martin (2021): Fossil turtles from the early Miocene localities of Mokrá-Quarry (Burdigalian, MN 4), South Moravian Region, Czech Republic. Geodiversitas 43 (20): 691-707, DOI: 10.5252/geodiversitas2021v43a20
523F87B8FF9EDC54FEAFFA4AE4CEF995.taxon	type_taxon	TYPE SPECIES. — Testudo (Chersine) hermanni Gmelin, 1789.	en	Luján, Àngel H., Čerňanský, Andrej, Bonilla-Salomón, Isaac, Březina, Jakub, Ivanov, Martin (2021): Fossil turtles from the early Miocene localities of Mokrá-Quarry (Burdigalian, MN 4), South Moravian Region, Czech Republic. Geodiversitas 43 (20): 691-707, DOI: 10.5252/geodiversitas2021v43a20
523F87B8FF9EDC54FEAFFA4AE4CEF995.taxon	description	(Figs 5; 6) LOCALITIES. — MWQ 1 / 2001, MWQ 2 / 2003, MWQTC / 2001, MWQ 4 / 2018 and MCQ 3 / 2005. STUDIED MATERIAL. — Czech Republic. South Moravia Region, Mokrá-Quarry, carapace remains (Fig. 5 A-V’): Pal. 1319, nuchal; Pal. 1320, nuchal; Pal. 1321, nuchal; Pal. 1322, neural 4; Pal. 1323, neural 6; Pal. 1350, neural 7; Pal. 1322, neural 4; Pal. 1324, pygal; Pal. 1325, pygal; Pal. 1328, right costal 1; Pal. 1326, left costal 1; Pal. 1327, left costal 1; Pal. 1329, right costal 2; Pal. 1330, left costal 3; Pal. 1331, left costal 6; Pal. 1332, left peripheral 1; Pal. 1333, right peripheral 1; Pal. 1334, left peripheral 2; Pal. 1335, left peripheral 2; Pal. 1336, right periph- eral 2; Pal. 1337, right peripheral 2; Pal. 1338, right peripheral 5; Pal. 1339, left peripheral 7; Pal. 1340, right peripheral 7; Pal. 1341, right peripheral 8; Pal. 1342, left peripheral 9; Pal. 1343, right peripheral 9; Pal. 1344, left peripheral 10; Pal. 1345, left peripheral 10; Pal. 1346, left peripheral 10; Pal. 1347, left peripheral 11; Pal. 1348, left peripheral 11. Pal. 1349, left peripheral 11. Plastral remains (Fig. 6 A-H’): Pal. 1351 A, left epiplastron; Pal. 1352, entoplastron; Pal. 1353, entoplastron; Pal. 1354, entoplastron; Pal. 1351 B, right hyoplastron; Pal. 1355, right hyoplastron; Pal. 1356, left hypoplastron; Pal. 1357, right hypoplastron; Pal. 1358, right hypoplastron; Pal. 1359, right hypoplastron; Pal. 1360, left xiphiplastron; Pal. 1361, left xiphiplastron; Pal. 1362, left xiphiplastron. DESCRIPTION Testudo material recovered from Mokrá-Quarry consists mainly of disarticulated plates (Pal. 1319 - 1362). However, some costals, peripherals and neurals are missing. They belong to a medium-sized Testudo, which reached approximately 20 - 25 cm. The following description is based on the entire available material, but not all the plates are depicted (Figs 5; 6). Most of the dermal grooves and sutures are visible. There is no ornamentation on the carapace or plastron. Growth lines are discernible in some carapace plates, and more specifically in both peripheral and costal plates (e. g., Fig. 5 K, A’). The nuchal plate is hexagonal, slightly wider than long, and its anterior edge is pointed (Fig. 5 A-D). The posterolateral edges are straight, whereas the anterior ones are curved in medial direction. A transverse thickening is recognizable on the visceral surface of the nuchal (Fig. 5 C, D). The cervical scute is longer than wide and its total length constitutes less than a half of the nuchal plate. It is well developed both dorsally and viscerally according to the three preserved plates (Pal. 1319 - 21). The anterior edge is narrower than the posterior ones. The lateral sulci are almost straight and parallel to each other (Fig. 5 A, B). Only three neurals plates are preserved, which vary in shape from subsquare (i. e., neural 4) to hexagonal (i. e., neurals 6 to 7: Fig. 5 E-G). It is noteworthy that none of them is more than twice as wide as long, and that they are encroached transversally by the intervertebral sulci (Fig. 5 E-G). The pygal plate is trapezoidal with slightly concave anterolateral margins. Its external surface is moderately convex, whereas its internal one is rather concave (Fig. 5 H-J). The marginal scutes 12 are missing, and therefore the supracaudal scute is not divided by a sagittal groove (Pal. 1324 - 25: Fig. 5 H-J). The shape of the vertebral scutes cannot be ascertained with a confidence because they are incomplete. Only five of the eight costal plates are present (i. e., costals 1 - 3 and 5 - 6), which are trapezoidal. Costals 1 and 3 host the intervertebral sulci (Fig. 5 K, L, O, P, W’), whereas the costals 2 and 6 host the interpleural sulci (Fig. 5 M, N, Q, R, W’). The pleural scutes are poorly preserved, so no significant details can be discerned. Despite this, the pleuromarginal sulcus coincides with the costoperipheral suture all along the preserved peripheral and costal plates (e. g., Fig. 5 K, L, W’). Peripheral plates 1 - 3, together with the nuchal, make up the anterior opening of the shell. Peripheral 1 is heptagonal, while the peripheral 2 is subtriangular. Peripheral 1 displays a moderately developed spike at about the middle of the anterior edge. The presence of protrusions on the remaining peripherals (i. e., 2 and 3) cannot be evaluated. Peripheral 5 (Pal. 1338: Fig. 5 A’, B’), which is the only preserved plate involved in the shell bridge, is rectangular and rather flat, unlike posterior peripherals. Pal. 1338 displays a very weak longitudinal lateral ridge that is placed slightly above of the marginoabdominal sulcus. Peripherals 7 - 11 (Fig. 5 C’ - W’), together with the pygal plate (Fig. 5 H-J), form the posterior opening of the shell. An elongated and subvertical scar of the dorsal projection of the hypoplastron is discernible in peripheral 7 internally (Fig. 5 E’, F’). As a rule, the peripherals 8 - 10 in Testudo are rectangular with dorsal surfaces slightly concave, whereas in Ptychogaster, they are rather subrectangular and the dorsal concavity is well-developed. Peripheral 11 hosts the lateral edge of the supracaudal scute (Fig. 5 S’ - W’). The marginal scute 1 is trapezoidal, whereas the remaining posterior marginals are either subsquare or subrectangular (Fig. 5 A-D, S-V, W’). Based on two nuchal plates (Pal. 1319 - 21), two peripherals 1 (Pal. 1332 - 33) and three costals 1 (Pal. 1326 - 28), the triple junction amongst the pleural 1, vertebral 1, and marginal 1 is located outside the nuchal plate. Noticeably, the posterior border of the marginal 5 (Pal. 1338; Fig. 5 A’, B’) is parallel (i. e., instead of oblique) relative to those anteroposterior edges of the peripheral 5. A partial epiplastron, together with the hyoplastron (Fig. 6 A-F, O-R), forms the anterior plastral lobe, which is rather trapezoidal (Fig. 6 I’). Despite not being entirely preserved, the epiplastral dorsal pad seems rectangular and longer than wide. It is moderately developed posteroventrally and overhangs it slightly. Consequently, a small gular pocket is present (Fig. 6 A-C). The gular scutes are triangular, with slightly sinuous lateral margins, and form an angle of less than 45 ° relative to the sagittal axis (Fig. 6 B-D). The three available entoplastra (Pal. 1352 - 54: Fig. 6 G-N) are hexagonal both ventrally and viscerally. They are partially covered by the gular scutes, which generally extend up to the middle of the entoplastron. The ventral surface of the epiplastra, covered by the gular scutes, is not in relief. In all specimens, the gular scutes are not crossed by the humeropectoral sulcus transversally (Fig. 6 H, I, K, L). Pal. 1351 B is the best preserved hyoplastron (Fig. 6 O-R), which hosts entirely both the humeropectoral and pectoroabdominal scutes: the former is nearly straight and obliquely oriented relative to the sagittal plane (i. e., only slightly sinuous: Fig. 6 R), while the latter is curved and transversally oriented relative to the sagittal plane (Fig. 6 R, V). The preserved portion of the humeral scute indicates that this was trapezoidal. The abdominal seems to be the largest scute of the plastron. Moreover, its medial sector is slightly oblique (Fig. 6 Y-Z). The hypo-xiphiplastral suture is roughly straight and well developed, which means that a plastral hinge is absent. The xiphiplastron is trapezoidal and its ventral side is very flat and crossed by the femoroanal sulcus (Fig. 6 A’ - C’). It is noteworthy that the distinct notch in their lateral margins, between the anal and femoral scutes, is missing (Fig. 6 A’, C’). The femoral is trapezoidal and much longer medially than the anal scutes, which are subrectangular and wider than long. The femoroanal sulcus is slightly sinuous and obliquely oriented relative to the sagittal plane. The anal notch is wider than long, and its visceral area covered by the anal scutes is variable, from moderately (Pal. 1361: Fig. 6 A’, B’) to well developed (Pal. 1362: Fig. 6 E’, F’).	en	Luján, Àngel H., Čerňanský, Andrej, Bonilla-Salomón, Isaac, Březina, Jakub, Ivanov, Martin (2021): Fossil turtles from the early Miocene localities of Mokrá-Quarry (Burdigalian, MN 4), South Moravian Region, Czech Republic. Geodiversitas 43 (20): 691-707, DOI: 10.5252/geodiversitas2021v43a20
523F87B8FF9EDC54FEAFFA4AE4CEF995.taxon	discussion	REMARKS The genus Testudo s. l. is a clade with five extant and multiple extinct species of terrestrial tortoises of western Palearctic distribution (Lapparent de Broin et al. 2006 a, b; Fritz & Bininda-Emonds 2007; Corsini et al. 2014; Delfino et al. 2012; Luján et al. 2016). Despite that, in the past, three extant genus-group taxa were distinguished as genera (see e. g., Turtle Taxonomy Working Group 2014), currently, the use of a single genus (Testudo s. l.) and three subgenera (Testudo, Agrionemys and Chersine) is being better accepted (see Luján et al. 2016; Graciá et al. 2017; Turtle Taxonomy Working Group 2017). Among extinct taxa, the taxonomy of Testudo s. l. also included the distinction of the extinct genus Paleotestudo Lapparent de Broin, 2000. Although the latter has not been the object of an exhaustive review, results presented by Luján et al. (2016) clearly pointed out that Paleotestudo is a junior subjective synonym of subgenus Chersine. Regarding the species Testudo kalksburgensis Toula, 1896, it was originally described by Toula (1896) based on one specimen from Kalksburg, Vienna: a partial shell currently housed at the IGUW. At the beginning of the twentieth century, new fragmentary material, coming from the late Miocene locality of Au am Leithaberge, was referred to T. kalksburgensis by Siebenrock (1914). Shortly after, Staesche (1931) also erected Testudo kalksburgensis var. steinheimensis based on various specimens from the middle Miocene (MN 7) locality of Steinheim (Germany). Although T. kalksburgensis was considered a junior subjective synonym of T. antiqua Bronn, 1831 by Glaessner (1933), subsequently Młynarski (1976) resurrected the species once again. Indeed, the validity of T. kalksburgensis was also confirmed by latter studies (see Młynarski 1980; Bachmayer & Młynarski 1981; Schleich 1981; Gemel & Rauscher 2000; Gemel 2002; Danilov et al. 2012; Luján et al. 2016; Březina et al. 2021).	en	Luján, Àngel H., Čerňanský, Andrej, Bonilla-Salomón, Isaac, Březina, Jakub, Ivanov, Martin (2021): Fossil turtles from the early Miocene localities of Mokrá-Quarry (Burdigalian, MN 4), South Moravian Region, Czech Republic. Geodiversitas 43 (20): 691-707, DOI: 10.5252/geodiversitas2021v43a20
