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LUJÁN Masaryk University, Faculty of Science, Department of Geological Sciences, 611 37 Brno (Czechia) and Comenius University in Bratislava, Faculty of Natural Sciences, Department of Ecology, 842 15 Bratislava (Slovakia) and Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, Edifici ICTA, 08193 Cerdanyola del Vallès (Spain) angel.lujan@icp.cat (corresponding author) Andrej ČERŇANSKÝ Comenius University in Bratislava, Faculty of Natural Sciences, Department of Ecology, 842 15 Bratislava (Slovakia) cernansky.paleontology@gmail.com Isaac BONILLA-SALOMÓN Department of Geology and Paleontology, Faculty of Natural Sciences, Comenius University, Mlynská dolina, Ilkovičova, 842 15 Bratislava (Slovakia) salomon1@uniba.sk Jakub BŘEZINA Moravian Museum, Department of Geology and Paleontology, 659 37 Brno (Czechia) jbrezina@mzm.cz Martin IVANOV Masaryk University, Faculty of Science, Department of Geological Sciences, 611 37 Brno (Czechia) mivanov@sci.muni.cz Submitted on 19 September 2019 | accepted on 30 March 2020 | published on 26 October 2021 urn:lsid:zoobank.org:pub:04F82471-9F26-4935-97CE-1A286E08C958 Luján À. H., Čerňanský A., Bonilla-Salomón I., Březina J. & Ivanov M. 2021. — Fossil turtles from the early Miocene localities of Mokrá-Quarry (Burdigalian, MN4), South Moravian Region, Czech Republic, in Steyer J.-S., Augé M. L. & Métais G. (eds), Memorial Jean-Claude Rage: A life of paleo-herpetologist. Geodiversitas 43 (20): 691-707. https://doi.org/10.5252/geodiversitas2021v43a20. http://geodiversitas.com/43/20 ABSTRACT KEY WORDS Early Miocene, Fossil turtles from Mokrá-Quarry, South Moravia Region, Czech Republic, are described in Burdigalian, this paper. Remains come from two already known karstic fissures uncovered in Mokrá-Western Ptychogaster, Quarry (1/2001 Turtle Joint and 2/2003 Reptile Joint), as well as three new karstic fissures from Testudo kalksburgensis, Chersine, Mokrá-Western Quarry (TC/2001 and 4/2018) and Mokrá-Central Quarry (3/2005). All locali- Czech Republic, ties correspond to the early Miocene (Burdigalian: late Eggenburgian-Ottnangian MN4). The Central Paratethys, Carpathian Foredeep newly described material belongs to several turtle individuals, including over 100 shell elements, Basin. so the material studied here constitutes one of the largest samples in regards of the Czech Repub- GEODIVERSITAS • 2021 • 43 (20) © Publications scientifiques du Muséum national d’Histoire naturelle, Paris. www.geodiversitas.com 691 Luján À. H. et al. lic fossil record. Most of these remains have been identified as Ptychogaster (Ptychogaster) sp. and Testudo (Chersine) cf. kalksburgensis Toula, 1896. However, three specimens presented here (i.e., one shell fragment and two postcranial bones) have been identified as a large tortoise (Testudi- nidae indet.). Turtle fauna is congruent with two ecological environments, including a dry karst landscape with open to dense steppe vegetation inferred for the heliophile testudinids, as well as freshwater masses to the semi-terrestrial ptychogasterid. Finally, this paper expands our knowledge of fossil turtle assemblages in Central Europe during the early Miocene. RÉSUMÉ Tortues fossiles des localités du Miocène inférieur de Mokrá-Quarry (Burdigalien, MN4), Moravie du Sud, République tchèque. Les tortues fossiles de la carrière de Mokrá (Moravie du Sud, République tchèque) sont décrites dans cet article. Les restes proviennent de deux fissures karstiques déjà connues, découvertes dans la carrière de Mokrá-Western (1/2001 Turtle Joint et 2/2003 Reptile Joint), ainsi que de trois nouvelles fissures karstiques des carrières de Mokrá-Western (TC/2001 et 4/2018) et de Mokrá- Central (3/2005). Toutes les localités (MN4) correspondent au Miocène inférieur (Burdigalien: Eggenburgien supérieur-Ottnangien). Le matériel nouvellement décrit appartient à plusieurs individus de tortues et comprend plus de 100 éléments de carapace, ce qui en fait l’une des plus grandes collections de tortues fossiles de République tchèque. La plupart de ces restes ont été MOTS CLÉS identifiés comme Ptychogaster (Ptychogaster) sp. et Testudo (Chersine) cf. kalksburgensis Toula, Miocène inférieur, 1896. Cependant, trois spécimens présentés ici (un fragment de carapace et deux os postcrâniens) Burdigalien, ont été identifiés comme appartenant à une grande tortue (Testudinidae indet.). Cette faune est Ptychogaster, Testudo kalksburgensis, congruente avec deux environnements écologiques, dont un paysage karstique sec à végétation Chersine, steppique ouverte à dense d’après les testudinidés héliophiles, et des masses d’eau douce d’après Paratéthys centrale, bassin d’avant-chaîne des le Ptychogaster semi-terrestre. Enfin, cet article complète nos connaissances sur les assemblages de Carpates. tortues fossiles d’Europe centrale au cours du Miocène inférieur. INTRODUCTION An assemblage formed by five karstic localities from Mokrá- Quarry (South Moravia Region) is studied here in detail. It Miocene turtle assemblages, which are frequent in Europe, is more-or-less biostratigraphically comparable to the Dol- are mostly found in basin deposits and karst areas. Currently, nice localities. This large open cast mine currently consists over ten early Miocene sites are known in Czechia, all of of three quarries named as follows: The Western (MWQ); them Burdigalian in age (MN3-MN4: Fejfar 1974, 1990; Central (MCQ) and Eastern (MEQ) respectively (Fig. 1). Fejfar & Kvaček 1993; Böhme & Ilg 2003). However, major- Unlike squamates, amphibians and mammals, turtle remains ity of turtle reports are based on unpublished remains (e.g., have been recovered in two of the three quarries (MWQ and PhD’s or online databases), or are recovered from faunal list MCQ: Ivanov et al. 2006; Ivanov 2008). Palaeontological in publications (Rzehak 1912, 1919; Čtyroký et al. 1973; research into Neogene fillings of fossiliferous karst joints Böhme & Ilg 2003; Kvaček et al. 2004; Schäfer 2013; Ivanov from Western Quarry (1/2001 and the 2/2003) started at the 2015; Bonilla-Salomón et al. 2021). Therefore, these records beginning of the 21st century, coinciding with commercial should be treated with caution, as some of them cannot be extraction work (Ivanov & Musil 2004; Ivanov et al. 2006). verified based on available sources. According to mammal During that time, research of the early Badenian marine fauna, the oldest early Miocene turtle occurrences come deposits, which are placed in the valley between the Western from the Most Basin (Ohře/Eger Graben, NW Czechia), and Central sectors, was carried out from analysis of several and more specifically from the limestone quarry of Tuchořice boreholes (Ivanov et al. 2006). More recently, Mokrá-Quarry and deposits of the main brown coal seam of the Ahníkov/ has provided three more turtle localities, which are named as Merkur-North Mine (both early Burdigalian: Eggenburgian, follows: TC/2001 and 4/2018 both from MWQ; and 3/2005 MN3). Otherwise, the youngest locality in Czechia would from MCQ. It is remarkable that the localities from Mokrá- be Dolnice site (Burdigalian: late Eggenburian-Ottnangian, Quarry have yielded one of the most abundant and complete MN4) placed in the Cheb Basin (Fejfar 1974, 1990). In herpetofauna of Central Europe, and more specifically from fact, Dolnice has been the only Czech site studied in detail the early Miocene (MN4), including: amphibians (Bufonidae, yielding fossil turtles, including descriptions and figures (see Pelobatidae, Proteidae, Ranidae, and Salamandridae: Ivanov Młynarski & Roček 1985). 2008); squamates (Lacertidae, Amphisbaenia, ?Scincoidea, 692 GEODIVERSITAS • 2021 • 43 (20) Fossil turtles from Mokrá-Quarry, Czech Republic (early Miocene) A B N Prague Brno N 500 km 50 km Legend C Proterozoic Cities MWQ 1/2001 and Paleozoic MWQ TC/2001 ‘Turtle joint’ (395 m a.s.l.) (380 m a.s.l.) Main fault Mesozoic MWQ 2/2003 MWQ 4/2018 ‘Reptile joint’ (365 m a.s.l.) (380 m a.s.l.) Main thrusts Paleogene MCQ 3/2005 Mokrá-Quarry Neogene Central; 200 m Western FIG. 1. — Approximate geographical position of Mokrá-Quarry (A). Schematic geological map of the Czech Republic (B). Position of the five Mokrá-Quarry locali- ties that have yielded fossil turtle remains (B, C). Anguidae, Varanidae, Boidae, Colubridae, Elapidae, Pythoni- folding as a result of the Variscan orogeny (Rez et al. 2011). dae, and Viperidae: Ivanov et al. 2018, 2020) and Testudines The karstic landscape of the Mokrá Plateau is then rather the (Geoemydidae and Testudinidae: Luján et al. 2017a; Bonilla- result of the chemical character of the carbonates, and more Salomón et al. 2021). specifically of the processes of pressure dissolution, as well as The first aim of this paper is to study the turtle composi- of the mechanical deformations of such carbonates (Hladil tion of all five Mokrá-Quarry localities. Secondly, we provide et al. 1987; Rez 2003). Neither Mesozoic nor Palaeogene descriptions and taxonomy of turtles based on all remains. sediments are present from the whole southern part of the Finally, we discuss the palaeoclimatic conditions given the Moravian Karst: instead, Neogene deposits are abundant. new data obtained from these fossil turtles. They consist of freshwater and brackish sediments (late Eggenburgian to Ottnangian), as well as deltaic sand, gravel and marine clay (Badenian). During the Karpatian, marine AGE AND GEOLOGICAL BACKGROUND sediments dominated the Moravian part of the Carpathian Foredeep, although deposits of that age are not preserved in Remains described in this paper come from Mokrá-Quarry the southern part of the Moravian Karst (Petrová et al. 2001). located about 12 km ENE of Brno. It is placed in the SE As the palaeobathymetric analysis of the Badenian sediments part of the Moravian Karst, on the Mokrá Plateau (Fig. 1). It pointed out, marine sedimentation in the foredeep moved lies in a close proximity of the margin fault of the West Car- far to the northwest. Consequently, Badenian transgression pathian Foredeep. The Mokrá Plateau is mainly formed by the reached its maximum affecting most of the Drahany Upland Vilémovice Limestone of the Macocha Formation (Givetian- and the Moravian Karst (Procházka 1899; Brzobohatý 1997). Frasnian), which is quite massive and biodetritic (Hladil et al. Remnants of such sediments are preserved in karst joints and 1987; Ivanov et al. 2006). However, Mokrá-Quarry is mainly open faults of the Mokrá Plateau, which are clearly recognized made up by Devonian carbonates (Hády-Říčka Limestones in the Vilémovice Limestone (e.g., limonitised layers or alter- of the Líšeň Formation) passing into the Lower Carbonif- natively for containing oyster fragments: Hladil et al. 1987). erous flysch facies, both often affected by several phases of More recently, this geological episode was also confirmed GEODIVERSITAS • 2021 • 43 (20) 693 Luján À. H. et al. by a number of sampling boreholes, which are rich mainly SYSTEMATIC PALAEONTOLOGY in foraminiferans, ostracods, sponge spicules and otoliths (Brzobohatý et al. 2000; Ivanov et al. 2006). Order TESTUDINES Linnaeus, 1758 Fossil mammals recovered from Mokrá-Quarry include Family GEOEMYDIDAE Theobald, 1868 remains of both micro- and macro-mammals. Thus far, only Subfamily PTYCHOGASTERINAE De Stefano, 1903 two localities from MWQ (1/2001 and 2/2003) have been preliminary studied. According to Ivanov et al. (2006), the first Ptychogaster Pomel, 1847 two fissures were reported including macro-mammals only, further indicating that these localities are of the early Miocene TYPE SPECIES. — Ptychogaster emydoides Pomel, 1847. in age (MN4). Shortly after, Sabol et al. (2007) confirmed the age for both fissures based on the micromammalian assemblage. As for rest of the localities (i.e., TC/2001, 4/2018 and 3/2005), Ptychogaster (Ptychogaster) sp. we tentatively attributed them to the early Miocene (MN4, (Figs 2; 3) Ottnangian) as well, due to the lack of more detailed studies. LOCALITIES. —MWQ1/2001, MWQ2/2003, MWQTC/2001, MWQ4/2018 and MCQ3/2005. MATERIAL AND METHODS STUDIED MATERIAL. — Czech Republic. South Moravia Region, Mokrá-Quarry, carapace remains (Fig. 2A-R’): Pal. 1300, nuchal Turtle remains of the five fossiliferous karst joints were recov- and left peripherals 1-3; Pal. 1301, nuchal; Pal. 1302, neural 4; Pal. 1303, neural 5; Pal. 1304, neural 7; Pal. 1305, suprapygal 1; ered during field campaigns (2002-2019). Thus far, over Pal. 1306, suprapygal 2; Pal. 1307, left costal 1; Pal. 1308, right costal 100 specimens of turtles have been recovered, of which a 6; Pal. 1309, right costal 6; Pal. 1310, left costal 8; Pal. 1311, right geoemydid and two testudinids (i.e., one small and another costal 8; Pal. 1312, right peripheral 1; Pal. 1313, left peripheral 7; large) were identified. In order to recover shell fragments and Pal. 1314, left peripheral 8. Plastral remains (Fig. 3A-R): Pal. 1315, bones mixed in calcareous sand and clays, all sediment was left hypoplastron; Pal. 1316, right hypoplastron; Pal. 1317, right hypoplastron; Pal. 1318, left xiphiplastron. washed in sieves of 0.5-2.0 mm mesh (Ivanov et al. 2006). H2O2 in aqueous solution was employed to dissolve the rock fragments and clay matrix, followed by the washing of the DESCRIPTION different samples with a regulated stream of running water. The ptychogasterid material (Pal. 1300-1318) from Mokrá- The material is currently held in the collections of the Depart- Quarry consists of numerous disarticulated plates, which present ment of Geological Sciences at Masaryk University (Brno, preserved parts of the almost the entire shell. However, some Czechia). It should be noted that some turtle remains were plates are missing from the carapace (i.e., pygal, costals 2-4 left unstudied, since they consisted mainly of poorly preserved and 7, neurals 1-3 and both medial and posterior peripher- carapace and plastron fragments (i.e., most of them juvenile als). The plastron is represented only by the hypoplastron individuals), and therefore, a precise anatomical identification and the xiphiplastron. The following description is based on for them was impossible. all available material, however, not all preserved plates are depicted in Figures 2 and 3. According to the dimensions of ABBREVIATIONS the nuchal plate (Pal. 1300-1301), costals (Pal. 1307-1311) Institutions and peripherals (Pal. 1312-1314), the shell would have been IGUW Institut für Geologische Wissenschaften, Universität relatively large (> 20 cm: Figs 2; 3). Ornamentation of both Wien (formerly Institut für Geologie), Vienna, Austria; carapace and plastron external surfaces is absent. As it is typi- Pal Department of Geological Sciences, Faculty of Sci- ence, Masaryk University, Brno, Czech Republic. cal for Ptychogaster plates, dermal grooves are well developed. Unlike Testudo, neither growth lines nor sutures (i.e., com- pletely fused to each other: see e.g., Fig. 2A-D) are preserved, Localities so it is impossible to evaluate them. 1/2001 Mokrá-Western Quarry 1/2001 Turtle Joint (MN4); The nuchal plate is hexagonal in outline, wider than long 2/2003 Mokrá-Western Quarry 2/2003 Reptile Joint (MN4); (Pal. 1301: Fig. 2E-H). It contacts the first pair of periph- 3/2005 Mokrá-Central Quarry 3/2005 (MN4); TC/2001 Mokrá-Western Quarry Turtle Cave (MN4); erals, the first pair of costals and the neural 1. The anterior 4/2018 Mokrá-Western Quarry 4/2018 (MN4). border possesses a shallow nuchal notch, affecting the nuchal border and the first peripherals (Pal. 1300: Fig. 2A, B). The TERMINOLOGY anteroposterolateral sides of the nuchal are rather equal in Anatomical shell nomenclature employed throughout the length, whereas the posterior border is narrow and slightly descriptions is based on Zangerl (1969), whereas the taxonomy convex anteriorly (Fig. 2E, F). In lateral view, the nuchal is used follows Luján et al. (2014, 2016, 2017b). vaguely curved (Fig. 2I, J). Two transversal thickenings are FIG. 2. — Carapace remains of Ptychogaster (Ptychogaster) sp. from Mokrá-Quarry localities: A-D, Pal. 1300 (nuchal and peripherals 1-3); A, B, dorsal; C, D, vis- ceral; E-L, Pal. 1301 (nuchal); E, F, dorsal; G, H, visceral; I, J, left lateral; K, L, anterior; M, N, Pal. 1302 (neural 4); M, dorsal; N, visceral; O-Q, Pal. 1303 (neural 5); O-P, dorsal; Q, visceral; R, S, Pal. 1304 (neural 7); R, dorsal; S, visceral. T, U, Pal. 1305 (suprapygals 1-2); T, dorsal; U, visceral; V, W, Pal. 1307 (costal 1); V, dor- 694 GEODIVERSITAS • 2021 • 43 (20) Fossil turtles from Mokrá-Quarry, Czech Republic (early Miocene) A B C D M E F I K G H J L N O P Q S’ T nu U per 1 Ce R S Ma 1 V Ve 1 X A-R’ co 1 ne 1 Pl 1 W Y A’ Z B’ C’ D’ F’ E’ sp 1 I’ J’ G’ H’ S’ py O’ P’ Q’ R’ K’ L’ M’ N’ sal; W, visceral; X-Z, Pal. 1308 (costal 5); X, Y, dorsal; Z, visceral; A’-C’, Pal. 1309 (costal 6); A’, B’, dorsal; C’, visceral; D’-F’, Pal. 1310 (costal 8); D’, E’, dorsal; F’, visceral; G’-J’, Pal. 1312 (peripheral 1); G’, H’, dorsal; I’, J’, visceral; K’-N’, Pal. 1313 (peripheral 7); K’, L’, dorsal; M’, N’, visceral; O’-R’, Pal. 1314 (peripheral 8); O’, P’, dorsal; Q’, R’, visceral views; S’, reconstruction of carapace. Thick lines indicate to scute sulci, dotted lines sutures and oblique lines denote missing plate portions. Abbreviations: Ce, cervical; co, costal; Ma, marginal; ne, neural; nu, nuchal; per, peripheral; Pl, pleural; py, pygal; sp, suprapygal; Ve, vertebral. Scale bars: 1 cm. GEODIVERSITAS • 2021 • 43 (20) 695 Luján À. H. et al. A B C G H I J D Gu epi W Hu hyo ent K Pec E L F S hyp Ab M Fe T xi An N W O P U V Q R A-V FIG. 3. — Plastral remains of Ptychogaster (Ptychogaster) sp. from Mokrá-Quarry localities: A-F, Pal. 1315 (hypoplastron); A, B, visceral; C, D, ventral; E, F, left lateral; G-L, Pal. 1316 (hypoplastron); G, H, visceral; I, J, ventral; K, L, right lateral; M-R, Pal. 1317 (hypoplastron); M, N, visceral; O, P, ventral; Q, R, right lat- eral; S-V, Pal. 1318 (xiphiplastron); S, T, visceral; U, V, ventral views; W, reconstruction of plastron. Thick lines indicate to scute sulci, dotted lines denote plate sutures, dashed lines denote completely ligamentous kinesis (between hypoplastra and peripherals and the hinge between hyoplastron and hypoplastron), and oblique lines indicate missing plate portions. Abbreviations: Ab, abdominal; An, anal; ent, entoplastron; epi, epiplastron; hyo, hyoplastron; hyp, hypoplastron; Fe, femoral; Gu, gular; Hu, humeral; Pe, pectoral; xi, xiphiplastron. Scale bars: 1 cm. 696 GEODIVERSITAS • 2021 • 43 (20) Fossil turtles from Mokrá-Quarry, Czech Republic (early Miocene) developed on the internal surface of this bone (Fig. 2C, D, peripheral plates, at least in both anterior and posterior part 2G-L). The cervical scute is present anteriorly, both dorsally of the carapace (Fig. 2A, B, E, F, V, A’, B’, D’, E’). and viscerally. It is a relatively large and trapezoidal element The peripheral plates 1-3 are longer than wide and slightly that is longer than wide (Fig. 2E, F). The lateral edges of the trapezoidal (Fig. 2A-D, G’-J’). They are completely fused cervical are slightly curved medially both in dorsal and vis- together (i.e., sutures are not visible) and crossed not only ceral sides (Fig. 2A, B, E-H). The overlap of this scute is less by the intermarginal sulcus, but also by the pleuro-marginal developed on the ventral surface (Fig. 2G, H). sulcus, unlike in Testudo. Peripherals 1-2 are prominent According to the preserved neural plates, an alternating in anterior direction in Pal. 1300. Peripheral 7, which is between octagonal and hexagonal plates forms the neural partially preserved, is rectangular (Pal. 1313: Fig. 2K’-N’). series: neural 4 octagonal (Pal. 1302: Fig. 2M, N) and neural 5 In internal view, it displays a rough elongated area for the and 7 hexagonal with short sides behind (Pal. 1303-1304: cartilaginous union of the inguinal process (Fig. 2M’, N’). Fig. 2O-S). A weak medial keel develops on the neural 7 Pal. 1313 is the last peripheral involved in the shell bridge (Pal. 1304: Fig. 2R, S) and the suprapygals 1-2 (Pal. 1305-1306: and also displays a weak lateral ridge on its external side. Fig. 2T, U). In visceral view, remains of the thoracic vertebrae Peripheral 8 is rectangular and hosts both the pleuromar- attachments are present in all neural plates (Fig. 2N, Q, S). ginal sulcus and intermarginal sulci (i.e., between margin- The suprapygal plate 1 is trapezoidal with a wider posterior als 8-9), which are situated far from the costoperipheral suture part (Pal. 1305: Fig. 2T). The anterior border is concave, (Pal. 1314: Fig. 2O’-R’). The marginal scute 1 is rectangular, whereas the posterior one is slightly convex. It contacts the slightly wider than long, whereas marginal 2 is trapezoidal. neural 8 anteriorly, costals 8 laterally and suprapygal 2 pos- Marginal 3 is approximately as wide as long. Marginal 8, teriorly. Suprapygal 2 is hexagonal, wider than long, and the only complete scute from the bridge area, is rectangular much wider than suprapygal 1 (Fig. 2T, U). It contacts the and higher than wide. The ventral overlap of all preserved suprapygal 1 and the posteromedial sides of costals 8 anteriorly, marginals is well developed. peripherals 11 laterally and pygal posteriorly. The anterolateral The preserved portion of the hyo-hypoplastral hinge is sides of the suprapygal 2 are slightly longer anteroposteriorly, straight and approximately transversal (Pal. 1377: Fig. 3M-P), compared to the posterolateral ones. The posterior side is which is located at the mid of the peripheral 6. The hypo- vaguely convex anteriorly (Fig. 2T, U). plastron contact anteriorly with the hypoplastron through a The vertebral scute series is partially preserved, which is hinge and laterally with the peripherals 6-7 through a com- quadrangular and slightly narrower than the costal series. pletely ligamentous union in the inguinal process (Pal. 1315- Vertebral 1 contacts the cervical and marginals 1-2 anteriorly 17: Fig. 3A-R). The visceral overlap of both abdominal and (Fig. 2B, F). It seems to be lyre-shaped and covers the lateral femoral scutes on hypoplastra is well developed (Fig. 3M, N). corners of the nuchal and costals 1 (Pal. 1300-1301: Fig. 2A, The xiphiplastron is trapezoidal and its posterior tip is rather B, E, F). According to preserved portion of the vertebral 3, rounded. The abdominofemoral sulcus, developed on the it expanded at least on costals 5 and neural 5 (Fig. 2X, Y). ventral surface of each hypoplastron, is concave (Fig. 2N, O). The sulcus between the vertebrals 3-4 is wavy in its medial Moreover, the latter does not reach the inguinal process later- part, and more specifically in the part that is crossing the neu- ally, but is located slightly below it. The femoroanal sulcus ral 5 (Fig. 2O, P). Vertebral 4 likely contacts the vertebral 3 is oblique, whereas the anal scute is triangular with rounded anteriorly, pleurals 3-4 laterally and vertebral 5 posteriorly lateral borders (Fig. 2S-V). The dorsal overlap of the anal (Fig. 2P, A’, B’, D’, E’). Vertebral 5 is the widest vertebral scute is moderately developed (Fig. 2S, T). scute, contacting with the vertebral 4 anteriorly, pleural 4 anterolaterally and marginals 11-12 posteriorly. It expands REMARKS on costals 8, neural 8, peripheral 12 and pygal, and therefore Ptychogasteridae is a geoemydid family which includes turtles covers the entire surface of the suprapygals 1-2. of medium body size. This clade originated in Europe dur- Although not fully preserved costals, the costal plate 1 is ing the Eocene and its members are characterized mainly by much longer than the rest of costal (Pal. 1307: Fig. 2V, W). a plastral kinesis (Lapparent de Broin 2001; Claude 2006). It is trapezoidal and always contacts the peripheral plates 1-3 Thus, the anterior part of the plastron is firmly connected to anterolaterally, nuchal anteromedially and neurals 1-2 medi- the carapace, whereas the posterior one is movable thanks ally. The anterior border is sinuous to articulate with the cor- to a hinge situated between hyo- hypoplastra and peripher- responding peripherals. Costal 6 is similar in regards of its als 6. Some studies claimed that ptychogasterids constitute a shape, being much wider than long (Pal. 1308-9: Fig. 2X-B’). monophyletic clade diagnosable by several synapomorphies The medial side of Pal. 1308 shows a short anteromedial and (Hervet 2004a, b, 2006). However, the inclusion within long posteromedial sides (Fig. 2X-Z). However, the medial side ptychogasterids of non-kinetic extinct geoemydids genera, of Pal. 1308 is most likely rounded (Fig. 2A’-C’). Costal 8 is such as Merovemys, Clemmydopsis and Hummelemys, has been narrow, slightly wider than long, contacting the costal 7 ante- questioned (see Claude & Tong 2004). A more comprehen- riorly, neural 8 and suprapygal 1 medially, and suprapygal 2 sive phylogenetic analysis would be required in the future to and peripherals 11-12 posteriorly (Pal. 1310-11: Fig. 2D’-F’). know what genera belong to subfamily Ptychogasterinae, as The pleural scutes are not preserved with the exception of the well as to further clarify which are the closest extinct relatives first one. The marginopleural sulcus is entirely situated on of the latter. GEODIVERSITAS • 2021 • 43 (20) 697 Luján À. H. et al. A B C D sf sf A-D E F G I J K L sc fi E-G sc fi H M H, I-M sc fi FIG. 4. — Shell and postcranial remains of Testudinidae indet. from MWQ4/2018: A-D, Pal. 1363 (shell fragment); A, B, dorsal; C, D, visceral; E-H, Pal. 1364 (scapula); E, lateral; F, medial; G, anterior; H, dorsal; I-M, Pal. 1365 (fibula); I, dorsal; J, ventral; K, posterior; L, anterior; M, lateral views. Dotted lines indicate plate sutures and oblique lines denote missing plate portions. Abbreviations: fi, fibula; sc, scapula; sf, shell fragment. Scale bars: 1 cm. As for the genus Ptychogaster, it was originally erected by DESCRIPTION Pomel (1847) from the early Miocene (MN2) of Saint-Gerand- Only a shell fragment formed by two portions of plates is known le-Puy, France. Two subgenera are currently distinguished (Fig. 4A-D). Pal. 1363 corresponds most likely to a carapace por- within Ptychogaster: P. (Ptychogaster); and P. (Temnoclemmys). tion, but it is poorly preserved and is not possible to assess this Ptychogaster (Ptychogaster) emydoides Pomel, 1847 (i.e., type confidently. The length and maximum width of the preserved species of subgenus Ptychogaster) has a complex taxonomical plate fragment is 6 cm. The external part is completely smooth and nomenclatural history, with many junior subjective syno- and is not crossed by any sulcus (Fig. 4C, D). A suture is recog- nyms (Schäfer 2013; Luján et al. 2014). Although one revision nized on top of plate, which is concave and approximately 2 cm of the genus Ptychogaster has recently been performed, the wide. A partial bone, belonging to the shoulder girdle, has been results of this PhD thesis are still unpublished (Schäfer 2013). identified (Pal. 1364: Fig. 4E-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 Subfamily TESTUDININAE Batsch, 1788 area to join with the visceral part of the carapace (Fig. 4H). The hind limb skeleton is restricted to one partial fibula (Fig. 4I-M) Testudinidae indet. (Fig. 4) that is elliptical in cross-section. Its distal articular surface is slightly small, oval and convex (Fig. 4M). Both postcranial bones LOCALITIES. — MWQ4/2018. are poorly preserved and no significant details can be discerned. STUDIED MATERIAL. — Czech Republic. South Moravia Region, Mokrá-Quarry, shell and postcranial remains (Fig. 4A-M): Pal. 1363, REMARKS shell fragment; Pal. 1364, scapula (i.e., anterodorsal process frag- Fossil remains of giant tortoises are not very common ment); Pal. 1365, right fibula. in Miocene assemblages of Central Europe; their record 698 GEODIVERSITAS • 2021 • 43 (20) Fossil turtles from Mokrá-Quarry, Czech Republic (early Miocene) being limited to few localities from Austria, Germany, eral 2; Pal. 1337, right peripheral 2; Pal. 1338, right peripheral Hungary and Switzerland (Alba et al. 2010, 2011; Car- 5; Pal. 1339, left peripheral 7; Pal. 1340, right peripheral 7; Pal. 1341, right peripheral 8; Pal. 1342, left peripheral 9; Pal. 1343, mona et al. 2011; Luján 2015). Loveridge & Williams right peripheral 9; Pal. 1344, left peripheral 10; Pal. 1345, left (1957) proposed that all European giant tortoises should peripheral 10; Pal. 1346, left peripheral 10; Pal. 1347, left be transferred into the extant genus Geochelone. This pro- peripheral 11; Pal. 1348, left peripheral 11. Pal. 1349, left posal was adopted for some time, and consequently, large peripheral 11. Plastral remains (Fig. 6A-H’): Pal. 1351A, left tortoise remains in Europe are still frequently referred to epiplastron; Pal. 1352, entoplastron; Pal. 1353, entoplastron; Pal. 1354, entoplastron; Pal. 1351B, right hyoplastron; Pal. 1355, in the literature as Geochelone sp. (e.g., Auffenberg 1974; right hyoplastron; Pal. 1356, left hypoplastron; Pal. 1357, right Młynarski 1976). However, current phylogenies do not hypoplastron; Pal. 1358, right hypoplastron; Pal. 1359, right support a close relationship between Mio- Pleistocene hypoplastron; Pal. 1360, left xiphiplastron; Pal. 1361, left xiphi- large tortoises and Geochelone. More recently, Bourgat & plastron; Pal. 1362, left xiphiplastron. Bour (1983) referred all giant fossil tortoises to the genus Cheirogaster. Most subsequent works accepted this genus DESCRIPTION attribution (e.g., Luján et al. 2010, 2014), until recently Testudo material recovered from Mokrá-Quarry consists when Pérez-García & Vlachos (2014) proposed that mainly of disarticulated plates (Pal. 1319-1362). How- European Neogene giant tortoises constitute a clade that ever, some costals, peripherals and neurals are missing. is more derived than the type species of Cheirogaster. To They belong to a medium-sized Testudo, which reached allocate these taxa, Pérez-García & Vlachos (2014) erected approximately 20-25 cm. The following description is the genus Titanochelon, with Ti. bolivari (Hernández- based on the entire available material, but not all the Pacheco, 1917) as its type species. This genus is charac- plates are depicted (Figs 5; 6). Most of the dermal grooves terized by a shell reaching over 100 cm and the fusion and sutures are visible. There is no ornamentation on the of marginal scutes 12 (i.e., constituting a supracaudal carapace or plastron. Growth lines are discernible in some scute). However, the evolution of gigantism amongst carapace plates, and more specifically in both peripheral fossil tortoises is clearly a homoplastic phenomenon, and costal plates (e.g., Fig. 5K, A’). mainly related to insular conditions, or adaptation to The nuchal plate is hexagonal, slightly wider than long, either global or local environmental changes (Kear 2010; and its anterior edge is pointed (Fig. 5A-D). The poste- Luján et al. 2010, 2017b; Itescu et al. 2014). Similarly, the rolateral edges are straight, whereas the anterior ones are fusion of marginal scutes 12 occurs in many extant and curved in medial direction. A transverse thickening is rec- extinct genera and cannot be considered autapomorphic ognizable on the visceral surface of the nuchal (Fig. 5C, D). for the genus Titanochelon. In summary, the taxonomy The cervical scute is longer than wide and its total length of the Miocene giant tortoises of Europe is still a subject constitutes less than a half of the nuchal plate. It is well of debate and will require improvement of existing data developed both dorsally and viscerally according to the matrix (e.g., including more skull characters) in order to three preserved plates (Pal. 1319-21). The anterior edge decipher the phylogenetic relationships of Titanochelon is narrower than the posterior ones. The lateral sulci are (Luján et al. 2017b). almost straight and parallel to each other (Fig. 5A, B). Only three neurals plates are preserved, which vary in shape from subsquare (i.e., neural 4) to hexagonal (i.e., Genus Testudo Linnaeus, 1758 neurals 6 to 7: Fig. 5E-G). It is noteworthy that none of them is more than twice as wide as long, and that they Subgenus Chersine Merrem, 1820 are encroached transversally by the intervertebral sulci (Fig. 5E-G). TYPE SPECIES. — Testudo (Chersine) hermanni Gmelin, 1789. The pygal plate is trapezoidal with slightly concave ante- rolateral margins. Its external surface is moderately convex, whereas its internal one is rather concave (Fig. 5H-J). The Testudo (Chersine) cf. kalksburgensis Toula, 1896 marginal scutes 12 are missing, and therefore the supracau- (Figs 5; 6) dal scute is not divided by a sagittal groove (Pal. 1324-25: Fig. 5H-J). The shape of the vertebral scutes cannot be LOCALITIES. — MWQ1/2001, MWQ2/2003, MWQTC/2001, ascertained with a confidence because they are incomplete. MWQ4/2018 and MCQ3/2005. Only five of the eight costal plates are present (i.e., costals STUDIED MATERIAL. — Czech Republic. South Moravia Re- 1-3 and 5-6), which are trapezoidal. Costals 1 and 3 host gion, Mokrá-Quarry, carapace remains (Fig. 5A-V’): Pal. 1319, the intervertebral sulci (Fig. 5K, L, O, P, W’), whereas nuchal; Pal. 1320, nuchal; Pal. 1321, nuchal; Pal. 1322, neural the costals 2 and 6 host the interpleural sulci (Fig. 5M, 4; Pal. 1323, neural 6; Pal. 1350, neural 7; Pal. 1322, neural N, Q, R, W’). The pleural scutes are poorly preserved, so 4; Pal. 1324, pygal; Pal. 1325, pygal; Pal. 1328, right costal 1; no significant details can be discerned. Despite this, the 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, pleuromarginal sulcus coincides with the costoperipheral left peripheral 1; Pal. 1333, right peripheral 1; Pal. 1334, left suture all along the preserved peripheral and costal plates peripheral 2; Pal. 1335, left peripheral 2; Pal. 1336, right periph- (e.g., Fig. 5K, L, W’). GEODIVERSITAS • 2021 • 43 (20) 699 Luján À. H. et al. Peripheral plates 1-3, together with the nuchal, make up that this was trapezoidal. The abdominal seems to be the the anterior opening of the shell. Peripheral 1 is heptagonal, largest scute of the plastron. Moreover, its medial sector while the peripheral 2 is subtriangular. Peripheral 1 displays is slightly oblique (Fig. 6Y-Z). The hypo-xiphiplastral a moderately developed spike at about the middle of the suture is roughly straight and well developed, which anterior edge. The presence of protrusions on the remaining means that a plastral hinge is absent. The xiphiplastron peripherals (i.e., 2 and 3) cannot be evaluated. Peripheral 5 is trapezoidal and its ventral side is very flat and crossed (Pal. 1338: Fig. 5A’, B’), which is the only preserved plate by the femoroanal sulcus (Fig. 6A’-C’). It is noteworthy involved in the shell bridge, is rectangular and rather flat, that the distinct notch in their lateral margins, between unlike posterior peripherals. Pal. 1338 displays a very weak the anal and femoral scutes, is missing (Fig. 6A’, C’). The longitudinal lateral ridge that is placed slightly above of the femoral is trapezoidal and much longer medially than the marginoabdominal sulcus. Peripherals 7-11 (Fig. 5C’-W’), anal scutes, which are subrectangular and wider than long. together with the pygal plate (Fig. 5H-J), form the posterior The femoroanal sulcus is slightly sinuous and obliquely opening of the shell. An elongated and subvertical scar of oriented relative to the sagittal plane. The anal notch is the dorsal projection of the hypoplastron is discernible in wider than long, and its visceral area covered by the anal peripheral 7 internally (Fig. 5E’, F’). As a rule, the periph- scutes is variable, from moderately (Pal. 1361: Fig. 6A’, B’) erals 8-10 in Testudo are rectangular with dorsal surfaces to well developed (Pal. 1362: Fig. 6E’, F’). slightly concave, whereas in Ptychogaster, they are rather subrectangular and the dorsal concavity is well-developed. REMARKS Peripheral 11 hosts the lateral edge of the supracaudal scute The genus Testudo s.l. is a clade with five extant and multiple (Fig. 5S’-W’). The marginal scute 1 is trapezoidal, whereas extinct species of terrestrial tortoises of western Palearctic the remaining posterior marginals are either subsquare or distribution (Lapparent de Broin et al. 2006a, b; Fritz & subrectangular (Fig. 5A-D, S-V, W’). Based on two nuchal Bininda-Emonds 2007; Corsini et al. 2014; Delfino et al. plates (Pal. 1319-21), two peripherals 1 (Pal. 1332-33) and 2012; Luján et al. 2016). Despite that, in the past, three three costals 1 (Pal. 1326-28), the triple junction amongst extant genus-group taxa were distinguished as genera (see the pleural 1, vertebral 1, and marginal 1 is located outside e.g., Turtle Taxonomy Working Group 2014), currently, the nuchal plate. Noticeably, the posterior border of the the use of a single genus (Testudo s.l.) and three subgenera marginal 5 (Pal. 1338; Fig. 5A’, B’) is parallel (i.e., instead (Testudo, Agrionemys and Chersine) is being better accepted of oblique) relative to those anteroposterior edges of the (see Luján et al. 2016; Graciá et al. 2017; Turtle Taxonomy peripheral 5. Working Group 2017). Among extinct taxa, the taxonomy A partial epiplastron, together with the hyoplastron of Testudo s.l. also included the distinction of the extinct (Fig. 6A-F, O-R), forms the anterior plastral lobe, which genus Paleotestudo Lapparent de Broin, 2000. Although is rather trapezoidal (Fig. 6I’). Despite not being entirely the latter has not been the object of an exhaustive review, preserved, the epiplastral dorsal pad seems rectangular and results presented by Luján et al. (2016) clearly pointed out longer than wide. It is moderately developed posteroventrally that Paleotestudo is a junior subjective synonym of subgenus and overhangs it slightly. Consequently, a small gular pocket Chersine. is present (Fig. 6A-C). The gular scutes are triangular, with Regarding the species Testudo kalksburgensis Toula, 1896, it slightly sinuous lateral margins, and form an angle of less was originally described by Toula (1896) based on one speci- than 45° relative to the sagittal axis (Fig. 6B-D). men from Kalksburg, Vienna: a partial shell currently housed The three available entoplastra (Pal. 1352-54: Fig. 6G-N) at the IGUW. At the beginning of the twentieth century, new are hexagonal both ventrally and viscerally. They are par- fragmentary material, coming from the late Miocene local- tially covered by the gular scutes, which generally extend ity of Au am Leithaberge, was referred to T. kalksburgensis up to the middle of the entoplastron. The ventral surface by Siebenrock (1914). Shortly after, Staesche (1931) also of the epiplastra, covered by the gular scutes, is not in erected Testudo kalksburgensis var. steinheimensis based on relief. In all specimens, the gular scutes are not crossed by various specimens from the middle Miocene (MN7) local- the humeropectoral sulcus transversally (Fig. 6H, I, K, L). ity of Steinheim (Germany). Although T. kalksburgensis was Pal. 1351B is the best preserved hyoplastron (Fig. 6O-R), considered a junior subjective synonym of T. antiqua Bronn, which hosts entirely both the humeropectoral and pec- 1831 by Glaessner (1933), subsequently Młynarski (1976) toroabdominal scutes: the former is nearly straight and resurrected the species once again. Indeed, the validity of obliquely oriented relative to the sagittal plane (i.e., only T. kalksburgensis was also confirmed by latter studies (see slightly sinuous: Fig. 6R), while the latter is curved and Młynarski 1980; Bachmayer & Młynarski 1981; Schleich transversally oriented relative to the sagittal plane (Fig. 6R, 1981; Gemel & Rauscher 2000; Gemel 2002; Danilov et al. V). The preserved portion of the humeral scute indicates 2012; Luján et al. 2016; Březina et al. 2021). FIG. 5. — Carapace remains of Testudo (Testudo) cf. kalksburgensis from Mokrá-Quarry localities: A-D, Pal. 1320 (nuchal); A, B, dorsal; C, D, visceral; E, Pal. 1322 (neural 4) in dorsal; F, Pal. 1323 (neural 6) in dorsal; G, Pal. 1350 (neural 7) in dorsal; H-J, Pal. 1325 (pygal); H, external; I, J, internal; K, L, Pal. 1326 (costal 1) in dorsal; M, N, Pal. 1329 (costal 2) in dorsal; O, P, Pal. 1330 (costal 3) in dorsal; Q, R, Pal. 1331 (costal 6) in dorsal; S-V, Pal. 1333 (peripheral 1); S, T, external; U, V, internal; W-Z, Pal. 1334 (peripheral 2); W, X, external; Y, Z, internal; A’, B’, Pal. 1338 (peripheral 5) in external; C’-F’, Pal. 1340 (peripheral 7); C’, D’, exter- 700 GEODIVERSITAS • 2021 • 43 (20) Fossil turtles from Mokrá-Quarry, Czech Republic (early Miocene) A B C C E F G Ma 1 Ce Ma 1 Ce n4 n6 n7 H I Ve 1 nu J K L Ve 1 Pl 1 M N py Ve 2 O P Ve 2 Ve 2 co 2 Pl 2 Spr co 1 Pl 1 Ma 1 Ce co 3 Q Pl 2 Ma 2 per 1 R Ve 3 W’ nu per 2 Ve 4 Ma 3 Pl 3 co 6 Ve 1 S T Ma 1 per 3 Pl 4 Ma 4 ne 1 Ma 1 co 1 Ma 2 U V Pl 1 Ve 2 co 2 Ma 2 Ma 5 ne 2 per 4 per 1 W X co 3 per 5 Ma 3 Ma 6 ne 3 Y Z Pl 2 Ve 3 co 4 ne 4 Ma 7 Ma 3 per 6 Ma 2 Ma 2 co 5 per 2 ne 5 Ma 8 per 7 A’ B’ Ve 4 Pl 3 ne 6 co 6 C’ D’ per 8 ne 7 Ma 9 ne 8 co 7 per 9 Pl 4 sp 1 co 8 Ve 5 Ma 10 per 10 sp 2 per 5 py W’ Ma 11 E’ Spr per 11 per 7 F’ K’ L’ M’ G’ H’ I’ J’ per 9 N’ per 8 per 11 per 10 S’ T’ A-V’ U’ V’ O’ P’ Q’ R’ nal; E’, F’, internal, G’-J’, Pal. 1341 (peripheral 8); G’, H’, external; I’, J’, internal; K’-N’, Pal. 1343 (peripheral 9); K’, L’, external; M’, N’, internal; O’-R’, Pal. 1345 (peripheral 10); O’, P’, external; Q’, R’, internal; S’-V’, Pal. 1348 (peripheral 11); S’, T’, external; U’, V’, internal views; W’, reconstruction of carapace Thick lines correspond to scute sulci, dotted lines denote plate sutures, and oblique lines indicate missing plate portions. Abbreviations: Ce, cervical; co, costal; Ma, mar- ginal; ne, neural; nu, nuchal; per, peripheral; Pl, pleural; py, pygal; sp, suprapygal; Spr, supracaudal; Ve, vertebral. Scale bars: A-V’, 1 cm; W’, 2.5 cm. GEODIVERSITAS • 2021 • 43 (20) 701 Luján À. H. et al. A B Gu E G I K M O epi epi ent Hu D Gu Gu H L ent C F J N Gu Hu Hu Hu ent hyo Hu Q A-H’ P Pe Gu I’ epi Hu ent hyo R hyo S Pe Hu Pe hyo Pe Ab T Ab hyp U Hu V hyo W Fe Pe xi X An I’ hyp Z Fe Y Ab A’ B’ C’ xi An Fe D’ E’ G’ H’ Fe Fe F’ xi xi An An An FIG. 6. — Plastral remains of Testudo (Testudo) cf. kalksburgensis: A-D’, from Mokrá-Quarry localities; A-F, Pal. 1351A (left epiplastron); A, C, visceral; B, D, ventral; E, F, left lateral; G-I, Pal. 1354 (entoplatron); G, visceral; H, I, ventral; J-L, Pal. 1353 (entoplatron); J, visceral; K, L, ventral; M, N, Pal. 1352 (entoplatron); M, visceral; N, ventral; O-R, Pal. 1351B (hyoplastron); O, P, visceral; Q, R, ventral; S-V, Pal. 1355 (hyoplastron); S, T, visceral; U, V, ventral; W-Z, Pal. 1356 (hypoplastron); W, X, visceral; Y, Z, ven- tral; A’-D’, Pal. 1360 (xiphiplastron); A’, B’, visceral; C’, D’, ventral; E’-H’, Pal. 1361 (xiphiplastron); E’, F’, visceral; G’, H’, ventral views; I’, reconstruction of plastron. Thick lines correspond to scute sulci, dotted lines denote plate sutures, and oblique lines indicate missing plate portions. Abbreviations: Ab, abdominal; An, anal; ent, ento- plastron; epi, epiplastron; hyo, hyoplastron; hyp, hypoplastron; Fe, femoral; Gu, gular; Hu, humeral; Pe, pectoral; xi, xiphiplastron. Scale bars: A-H’, 1 cm; I’, 2.5 cm. 702 GEODIVERSITAS • 2021 • 43 (20) Fossil turtles from Mokrá-Quarry, Czech Republic (early Miocene) DISCUSSION that, apart from Titanochelon, a second giant tortoise inhab- ited Central Europe during the early Miocene (Luján pers. TAXONOMIC ATTRIBUTION OF THE DESCRIBED REMAINS observ.). In any case, the material presented here is the Ptychogaster (Ptychogaster) sp.: Almost half of the fossil turtle second report of giant testudinids in Czechia (see Březina material from Mokrá-Quarry can be referred to the genus et al. 2019), and certainly, more material will be required Ptychogaster. This is the case, not only for the material com- to clarify the taxonomy of this finding. ing from the localities of 1/2001 and 2/2003, but also from the newly recovered ones, from both the western (TC/2001 Testudo (Chersine) cf. kalksburgensis and MWQ4/2018), and central part (3/2005). Ptychogas- The second most abundant turtle material from Mokrá- terid remains of both localities (1/2001 and 2/2003) were Quarry is a small testudinid, ascribed to genus Testudo. It originally referred to extant genus Chrysemys by Ivanov & belongs to subgenus Chersine (see Luján et al. 2016), which Musil (2004), or alternatively to family Emydidae subse- is characterized by: a trapezoidal (instead of rounded) ante- quently (see Ivanov et al. 2006). In neither of these two rior plastral lobe; in the posterior border of marginal scute works, did the authors provide a discussion, nor figured 5 being parallel (instead of oblique) to those of peripheral these shell remains, and therefore this attribution cannot be plate 5; and the absence of a plastral hinge, along with a justified. However, the geoemydid material described here not elevated shell bridge. Among the species of the subge- can be referred to the genus Ptychogaster (Ptychogasteridae: nus Chersine from Central Europe, the material described Ptychogasterinae) on the basis of the well developed pair here most closely resembles T. (Ch.) kalksburgensis in the of internal thickenings on the ventral surface of nuchal, a morphology of the epiplastral dorsal pad (i.e., moderately completely ligamentous union between hypoplastron and developed posteroventrally and overhanging it slightly), peripherals, and the presence of a hinge between hyo- and and by the presence of a small gular pocket (Luján et al. hypoplastron at the level of the peripheral 6. Moreover, the 2016). Due to incomplete preservation of the anterior most complete ptychogasterid remains show longer ante- edge of epiplastron (i.e., protruding in T. steinheimensis) rior peripherals, irregular neural series formed mainly by and the absence of the neural 1 (i.e., with a marked pos- octagonal and hexagonal plates with short sides in front, terior constriction in T. catalaunica), we cannot conclude and a nuchal that is longer than wide. The latter features that specimens from Mokrá-Quarry belong to T. (Ch.) characterize the subgenus Ptychogaster, and further discounts kalksburgensis. an alternative attribution to the subgenus Temmnoclemmys (Luján et al. 2014). Given that the most diagnostic shell part PALAEOECOLOGICAL RECONSTRUCTION in ptychogasterids is missing (i.e., the dorsal epiplastral lip), Mokrá-Quarry localities represent one of the most diverse the remains from Mokrá-Quarry are too scarce to provide a and well-documented early Miocene (late Eggenburgian- taxonomic assignment at the species level. In any case, the Ottnangian) vertebrate assemblages in Central Europe ptychogasterid material presented here enables to confirm that (Ivanov & Musil 2004; Ivanov 2008; Ivanov et al. 2006, Ptychogaster was also present in the South Moravian Region 2018, 2020). To date, 40 ectothermic taxa belonging to during the early Miocene (MN4). To date, the ptychogas- amphibians (13 taxa) and reptiles (three turtle taxa and 24 terids had only been reported in Ohře/Eger Graben (NW squamate taxa) have been recorded. Although the number Czechia), mainly on the basis of described material from of turtle remains recovered from Mokrá-Quarry is relatively Dolnice site (Cheb Basin: Młynarski & Roček 1985). This high (over 100 shell plates), it shows a rather low diversity, agrees with the range of subgenus Ptychogaster in Czechia, including two tortoises (of which one is gigantic) and one where it is already recorded, but has yet to be studied in geoemydid. Turtle fauna is consistent with three different detail, including the following early Miocene (MN3) locali- palaeoenvironments, which are further supported by the ties: Ahníkov/Merkur-North Mine, Břešťany, Most/Brüx, faunal assemblages of mammals, amphibians, and the rest Tuchorice and Želeč (Laube 1900, 1901; Schlosser 1910; of reptiles (i.e., squamates). The members of the genus Liebus 1930; Fejfar & Kvaček 1993; Kvaček et al. 2004: Ptychogaster were classified as aquatic turtles by Reinach Böhme & Ilg 2003; Schäfer 2013; Ivanov 2015; Bonilla- (1900), which means they spend most of their lives in the Salomón et al. 2021). water bodies, only venturing ashore to lay their eggs or to thermoregulate their bodies. Conversely, De Stefano (1903) Testudinidae indet. proposed that ptychogasterids were actually semi-terrestrial, Large-sized testudinid material recovered from MWQ4/2018 which would imply that they will spend a significant amount is similar in size to that of the genus Titanochelon (Pérez- of time on land near bodies of water, or even close to the García & Vlachos 2014). Particularly, the morphology of shoreline. He justified this hypothesis mainly on the propor- preserved postcranial bones (i.e., scapula and tibia), along tions and robustness of its forelimbs. Subsequent authors with the shell plate, resembles that of the middle Miocene (e.g., Młynarski 1976; Gemel 2002) also supported this (MN6) species Ti. vitodurana from Switzerland. However, lifestyle, since they had a high vaulted carapace: nevertheless, specimens from MWQ4/2018 cannot be referred to genus they did not discard that some species could be semiaquatic. Titanochelon on the basis of size, as material from Ahníkov/ As a rule, all semiaquatic and semi-terrestrial turtles (i.e., Merkur Mine (Ohře/Eger Graben, NW Czechia) suggests Emydoidea, Kinosternon, Terrapene and some species of Rhi- GEODIVERSITAS • 2021 • 43 (20) 703 Luján À. H. et al. noclemmys) display a relatively vaulted shell (Marmi & Luján CONCLUSIONS 2012). However, all extant turtles with a domed shell are not semi-terrestrial, and therefore this feature is not useful Turtle remains from the early Miocene (MN4) localities of to infer the lifestyle (e.g., Joyce & Gauthier; Benson et al. Mokrá-Quarry (Czech Republic) are herein described and 2011). Moreover, different species within the same genus attributed to three taxa: Ptychogaster (Ptychogaster) sp.; Testu- may have different natural histories and, as a result, some dinidae indet.; and Testudo (Chersine) cf. kalksburgensis. Large of them have a greater tendency to travel overland (Steen testudinids were recovered in MWQ4/2018 only, which is likely et al. 2012). Despite the fact that we cannot confirm all attributable to a fossil record bias. Both small testudinid and ptychogasterid species to have been semi-terrestrial turtles ptychogasterid remains represent the first fossil record from (not within the scope of this paper), the Ptychogaster mate- the South Moravian Region. The turtle assemblage recovered rial from Mokrá-Quarry is consistent with the nearby pres- from Mokrá-Quarry is similar to those from the Vienna Basin ence of freshwater (Alba et al. 2018; Casanovas-Vilar et al. and the North Alpine Foreland Basin. Finally, the presence of 2018). According to the palaeoreconstruction conducted by two terrestrial tortoises and one semi-terrestrial geoemydid is Ivanov et al. (2018, 2020) based on amphibian and reptiles, entirely consistent with the palaeoenvironmental reconstruction ptychogasterids from Mokrá-Quarry would have lived in the of Mokrá-Quarry localities, as a karst landscape surrounded surroundings of damp, or even marsh-like habitats. by open forest and steppe. Two tortoise taxa (Testudinidae indet. and Testudo cf. kalksburgensis), adapted to dry and open environments have been found in Mokrá-Quarry. The giant tortoise, Acknowledgements recovered only in MWQ4/2018, had a large body and The research reported in this work was funded by the Span- would had needed some kind of vegetation cover or caves ish Agencia Estatal de Investigación – European Regional (i.e., shaded places) in order to cool down their body tem- Development Fund of the European Union (ÀHL: CGL2016- perature (Schleich 1988; Böhme 2008). In any case, the 76431-P, CGL2017-82654-P and AEI/FEDER EU), the giant tortoises would have inhabited areas near to open Generalitat de Catalunya (ÀHL: CERCA Programme), the forest environments. In turn, the small tortoise Testudo VEGA Scientific Agency (AČ: 1/0209/18; IBS: 1/0164/19), cf. kalksburgensis would have preferred an even more open the APVV grant agency of the Slovak Republic (IBS: APVV- and dryer environment, such as a steppe (e.g., Baruš et al. 16-0121), the Ministry of Culture of the Czech Republic (JB: 1992; Čerňanský et al. 2012). DKRVO, MK000094862), the SPECIFIC research project Therefore, Mokrá-Quarry localities should be interpreted at the Faculty of Science at Masaryk University, Brno (MI, as a karst landscape surrounded by open forest environments JB: MUNI/A/0944/2019), and from the Operational Pro- and steppe, as shown by the abundant remains of amphibians gramme Research, Development and Education – Project and reptiles (Ivanov 2008; Ivanov et al. 2018; 2020). Some “Postdoc@MUNI” (No. CZ.02.2.69/0.0/0.0/16_027/000 small mammal taxa recovered from both localities (1/2001 8360). ÀHL acknowledges a research visit to the University and 2/2003) have been associated with open environment of Comenius in Bratislava, Slovakia (National Scholarship conditions and warm weather, including the cricetodontids Programme of the Slovak Republic for the support of mobil- genera Democricetodon and Megacricetodon, and the sciurid ity of researchers, Autumn Call 2017). We thank the editors Palaeosciurus (Sabol et al. 2007). The record of Melissiodon M. Augé, G. Metais and J.-S. Steyer for preparing a special also suggests wooden areas and somewhat more humid condi- volume tribute to the memory of Jean-Claude Rage. Finally, tions (Jovells-Vaqué & Casanovas-Vilar 2018). With regards we are grateful to Walter Joyce and to an anonymous reviewer to the large mammal association, they consist of cervids for many helpful comments and suggestions that helped us and one suid, which confirm forested and swampy environ- to improve a previous version of this paper. ments (Rössner 2004). Despite the similar palaeoecology of both fissures (1/2001 and 2/2003), however herpetofauna of 2/2003 points to a slightly more humid palaeoenviron- ment, supported by the abundance of amphibians such as REFERENCES Triturus, Chioglossa and Mertensiella as well (Ivanov 2008; ALBA D. M., CARMONA R., BOLET A., ROBLES J. M., CASANOVAS- Ivanov et al. 2018). 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