Early Miocene small mammals from MWQ1/2001 Turtle Joint (Mokrá-Quarry, South Moravia, Czech Republic): biostratigraphical and palaeoecological considerations Isaac Bonilla-Salomón, Stanislav Čermák, Àngel H. Luján, Ivan Horáček, Martin Ivanov & Martin Sabol Mokrá-Quarry (South Moravia, Czech Republic) represents a unique fossil site with an extraordinary abundance and diversity of vertebrate fossil remains. Most research on Mokrá-Quarry localities was focused on herpetofauna. Despite its relevance, the mammal fauna has not been yet studied in detail. In this work, the small mammals from one of the karstic fissures, Mokrá-Western Quarry (1/2001 Turtle Joint), are thoroughly described for the first time, including eight different taxa: Prolagus schnaitheimensis, Prolagus cf. vasconinensis, Rhinolophus cf. cluzeli, Rhinolophus cf. grivensis, Galerix sp., Aliveria aff. luteyni, Megacricetodon sp., and Melissiodon dominans. The Megacricetodon finds represent one of the first appearances of this cricetid in Central Europe and sheds light on the early evolution of the genus. Moreover, the small mammal assemblage confirms an early Miocene age (Burdigalian, MN4) for MWQ1/2001. The paleoenvironment inferred shows a dry karst landscape, with patches of woodlands and open steppe, together with marshy areas. • Key words: Lagomorpha, Rodentia, Chiroptera, Eulipotyphla, Carpathian Foredeep Basin, Moravian Karst. Bonilla-Salomón, I., Čermák, S., Luján, À.H., Horáček, I., Ivanov, M. & Sabol, M. 2021. Early Miocene small mammals from MWQ1/2001 Turtle Joint (Mokrá-Quarry, South Moravia, Czech Republic): biostratigraphical and palaeoecological considerations. Bulletin of Geosciences 96(1), 99–122 (9 figures, 2 tables). Czech Geological Survey, Prague. ISSN 1214-1119. Manuscript received Mai 28, 2020; accepted in revised form November 25, 2020; published online December 20, 2020; issued January 10, 2021. Isaac Bonilla-Salomón & Martin Sabol, Comenius University, Faculty of Natural Science, Department of Geology and Paleontology, Mlynská dolina, Ilkovičova Str. 6, SK-84215 Bratislava, Slovak Republic; salomon1@uniba.sk • Stanislav Čermák, Institute of Geology of the Czech Academy of Sciences, Rozvojová 269, 16500 Prague 6, Czech Republic • Àngel H. Luján, Masaryk University, Faculty of Science, Department of Geological Sciences, Kotlářská 267/2, 611 37 Brno, Czech Republic & Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, Edifici ICTA, c/ Columnes s/n, Campus de la UAB, 08193 Cerdanyola del Vallès, Barcelona, Spain; angel.lujan@icp.cat • Ivan Horáček, Charles University, Faculty of Science, Department of Zoology, Viničná 7, CZ-12844 Praha 2, Czech Republic • Martin Ivanov, Masaryk University, Faculty of Science, Department of Geological Sciences, Kotlářská 267/2, 611 37 Brno, Czech Republic Early Miocene fossil mammal assemblages are quite Ottnangian, Burdigalian) based on the data obtained from frequent in Central Europe, however, only nine localities micromammals (Cicha et al. 1972, Fejfar 1990). However, are known from the territory of the Czech Republic (Cicha the most recent fossil bearing site discovered in the Czech et al. 1972; Fejfar, 1974, 1990; Fejfar & Roček 1986; Republic is the open-cast limestone mine of Mokrá-Quarry, Fejfar & Kvaček, 1993; Fejfar et al. 2003; Ekrt et al. which is placed 12 km ENE of Brno city. It is located on 2016). The older ones come from the early Burdigalian the Mokrá Plateau, situated in the SE of the Moravian deposits of the main brown coal seam of Ahníkov/Merkur Karst, close to the margin fault of the West Carpathian Mine (early MN3a) and the limestone quarry of Tuchořice Foredeep. The Mokrá Plateau is mostly formed by the (MN3b), both located in the Most Basin (Ohře/Eger massive and biodetritic Vilémovice Limestone (Givetian– Graben, NW Czech Republic) and of early Burdigalian Frasnian) of the Macocha Formation (Ivanov et al. 2006). (Eggenburgian) in age. Regarding MN4 sites, the three The western part of the Mokrá-Quarry is mainly made up Dolnice localities (Dolnice 1–3) from the Cheb Basin by the Vilémovice Limestone, whereas the central part of (Burdigalian, late Ottangian–Karpatian) have yielded the quarry is formed also by the Hády-Říčka Limestone abundant vertebrate remains (Fejfar 1974, 1990; Fejfar and the Křtiny Limestone of the Líšeň Formation, as well & Kvaček 1993). As for the South Moravian Region, the as breccia and sandy limestones with shales of the Březina site of Ořechov was correlated with the MN4 zone (late Formation (Rez 2010). Those carbonates, which pass into DOI 10.3140/bull.geosci.1801 99 Bulletin of Geosciences • Vol. 96, 1, 2021 the lower Carboniferous flysch facies, are often affected MWQ1/2001 and MWQ2/2003 was published (Ivanov by several phases of folding due to the Paleozoic orogeny. et al. 2006), but neither descriptions nor measurements of The chemical character of the carbonates, together with the fossil material were provided. As for small mammals, the process of pressure dissolution and the mechanical only a preliminary study of the material from MWQ1/2001 defor­mation (Hladil et al. 1987; Rez 2003, 2010), are the was conducted by Sabol et al. (2007). main factors responsible of karstic landscape of Mokrá- In this work, all micromammal remains from Quarry. MWQ1/2001 were revised for the first time. Moreover, In the southern part of the Moravian Karst, Neogene a thorough discussion on the age of this fossil assemblage freshwater and brackish sediments of Ottnangian age, and its paleoecological conditions are assessed. together with Badenian deltaic sands, gravels and marine clays, are abundant. Even though marine sediments dominated the Carpathian Foredeep in the South Moravia Material and methods Region during the Karpatian stage, those sediments are not preserved in the southern part of the Moravian Karst. Terminology. – The micromammal fauna from 1/2001 was The Badenian transgression affected the majority of the discovered during the field campaigns leaded by M. Ivanov Drahany Upland and the Moravian Karst (Procházka and R. Musil from the Masaryk University (Brno, Czech 1899, Brzobohatý 1997). Remnants of those sediments Republic) during the years 2002–2005. The remains were occur in karst fissures, as confirmed by several bore­ obtained by washing the fossil material, mixed in sand holes located between the central and western quarries, and clays, in sieves of 0.5 mm mesh (Ivanov et al. 2006). which yielded rich remains of foraminiferans, ostracods, Currently, the material is housed in the collections of the sponge spicules and fish otoliths (Brzobohatý et al. Department of Geological Sciences (Faculty of Science, 2000). Masaryk University) under the inventory numbers Indeed, Mokrá-Quarry consists of three separated SMM/009-09-11/ 372009, Pal. 3000–3846. Upper cheek quarries: the western (MWQ), the central (MCQ), and teeth are indicated by upper case letters (P4, M1, M2, the eastern one (MEQ: Fig. 1). Several karst fissures M3) and lower cheek teeth by lower case (p4, m1, m2, have been located in central and western quarries, one m3). When distinction between first and second molars is yielding a rich assemblage of the early Pleistocene age dubious, these are indicated as M1/2 or m1/2. (Fejfar & Horáček 1983). At the beginning of 21st century, Dental terminology and metrics employed in the commercial extraction works started (Ivanov & Musil present work follow the next authors: Sciuridae (Cuenca- 2004, Ivanov et al. 2006), leading to the discovery of two Bescós 1988); Cricetidae (Oliver & Peláez-Campomanes fossiliferous karst fissures in the western quarry: 1/2001 2013); Erinaceidae (Engesser 1980); Chiroptera (Hand Turtle Joint (hereinafter MWQ1/2001), and 2/2003 1985, Horáček & Špoutil 2012); Lagomorpha (Lopéz- Reptile Fissure (hereinafter MWQ2/2003). The last years Martínez 1989, Angelone & Sesé 2009, Hordijk & van of research in Mokrá-Quarry have provided three more der Meulen 2010a, Hordijk et al. 2010). Lagomorpha localities, which are the following: MCQ3/2005 from drawings and measures were taken with the aid of a Dino- the central quarry; MWQ-TC4/2001 and MWQ4/2018 Lite digital microscope. from the western quarry. It is noteworthy that turtle fossil Dental measurements of lagomorphs were taken only remains are especially abundant in the above-mentioned in adult specimens (recognized by their prismatic tooth localities (see Luján et al. 2017, in press). In any case, shape), as maximum antero-posterior or bucco-lingual all research efforts in Mokrá-Quarry have been focused dimensions of the respective two-dimensional dental on the herpetofauna to date. According to Ivanov (2008), structures orthogonal to prismatic shaft. For the rest of thirteen different amphibian taxa were reported from both groups, all measurements were taken with the Carl Zeiss fissures MWQ1/2001 and MWQ2/2003, including the Stemi 305 microscope and the Carl Zeiss W-PI 10x/23 first records of the West European species of Triturus cf. Microscope Focusable Eyepiece, providing cheek teeth marmoratus and Chioglossa meini in Central Europe as occlusal surface maximum length and width (L × W). early as MN4. Moreover, the recent description of a new All measured data are given in millimetres (mm). species of varanid, Varanus mokrensis from MWQ1/2001 Micrographs were taking using the JSM-6390 Scanning and MWQ2/2003, provided the first documented example Electron Microscope (SEM) at the Earth Science Institute of a European varanid for which osteological data allowed in Banská Bystrica (Slovakia). a well-supported assignment at the genus level (Ivanov et Neogene time scales boundaries follow the Inter­ al. 2018, Villa et al. 2018). Regarding mammals, the first national Chronostratographic Chart v2020/03 (www. preliminary ascriptions of some medium-sized specimens stra­tigraphy.org). MN zones, where their boundaries are were conducted by Ivanov and Musil (2004). Shortly after, defined based on selected appearance events, follows a faunal list of medium-sized and large mammals from Hilgen et al. (2012) in general. We follow Hilgen et al. 100 Isaac Bonilla-Salomón et al. • Early Miocene small mammals Figure 1. Location of Mokrá- A B Quarry. • A – geographical po­s-­ ition of Mokrá-Quarry. • B – schematic global map of the Czech Republic. • C – position of 1/2001 in Mokrá-Quarry. Modified from Luján et al. (in press, fig. 1). 500 km 50 km C 200 m (2012) for defining the basis for MN3 and MN5, but attribution to the sole genus Prolagus: (i) the complete Steininger (1999) for MN4. rootless hypsodonty in permanent teeth; (ii) isolated anter­o- conid in p3; (iii) absence of m3; (iv) hypoconulid in m2; Measurements and locality abbreviations. – Measurements (v) P2 with distinct meso- and paraflexus; (vi) P3 with abbreviations: AA – partial width; CV – coefficient of well-developed paraflexus and mesoflexus; (vii) enamel variation [%]; L – length; La – anteroconid length; Lme – fossettes in P4–M2. Two distinct species of Prolagus metaconid–entoconid length; Lpl – postlobus length; (P. schneitheimensis and P. cf. vasconiensis) were distin­ Ltrig – trigonid length; L × W – maximum length and g­uished in MWQ1/2001 assemblage, well differing from width of the cheek tooth occlusal surface; M/m – upper/ each other by the size, curvature, and relative length of lower molar; N – number of specimens; OR – observed tooth shafts; size and morphology of p3 anteroconid; range; P/p – upper/lower premolar; PH – hypoflexus width; com­­plexity of p3 protoconid; size and morphology of TH – distal hyperloph length; W – width; Wa – anteroconid P4–M2 fossettes; and length of M1–M2 hypoflexus (see width; Wdh – precone–distal hypercone width; Wtl – third “Descriptions” for details). lobe width of m2; Wtal – talonid width; Wtrig – trigonid The lagomorph fauna of MWQ1/2001 comprises width; x̄ – arithmetic mean. predominantly (ca. 85%) the species P. schnaitheimensis, Locality abbreviations: MWQ1/2001 – Mokrá-Western which is characterized particularly by: straight and short Quarry 1/2001 Turtle Joint (MN4); MWQ2/2003 – p3 shaft without antero-posterior bending; very shallow Mokrá-Western Quarry 2/2003 Reptile Fissure (MN4). p3 centroflexid; simple p3 protoconid; short P4–M2 shafts with a small radius of linguo-buccal curvature. This fossil lagomorph represents one of the prolagid linea­ges Systematic palaeontology that evolved in the “cricetid vacuum”, which was first described by Tobien (1975) from the early Miocene loca­l­- Class Mammalia Linnaeus, 1758 ity Schnaitheim (MN3, Germany). The species is also Order Lagomorpha Brandt, 1855 known from many other MN3–MN4 localities of Central Famil: Prolagidae Gureev, 1960 Europe (see e.g. Tobien 1975, Ziegler & Fahlbusch 1986, Sabol et al. 2007). The material presented here is fully Genus Prolagus Pomel, 1853 compatible with the one from the type locality, except for: a higher degree of evolution of M1–M2 parafossettes and Discussion. – Lagomorphs from MWQ1/2001 are ex­clu­ P3 lagiloph; the predominantly bifurcated P3 parafossette siv­ely represented by isolated teeth. The combination of with more plicated enamel walls; the longer lagiloph; and the following discriminant features clearly supports their the shorter mesial hyperloph. These features are most 101 Bulletin of Geosciences • Vol. 96, 1, 2021 similar to Prolagus recorded from the localities Limberg including the type series described by Viret in Roman & and Echzell (Tobien, 1975). Two M1 specimens in the Viret (1930, fig. 7b) and additional material described material under study possess highly crenulated enamel by Tobien (1975, fig. 58). The material from the older walls of parafossette (Fig. 2O), resembling Ptychoprolagus locality of Laugnac (MN2) presented by Tobien (1975) forsthartensis, a descendant of P. schnaitheimensis shows, on the other hand, more developed P3 lagiloph and (Tobien 1975, López Martínez 2001, Hordijk et al. 2010). longer and more curved P3 mesoflexus; this may indicate However, the mentioned parafossettes from MWQ1/2001 similar morphoclines (reduction of the lagiloph in the P3 are much less crenulated than those in the type material and enlargement and bending of the mesoflexus), such as originally described by Tobien (1975) from the MN4 in the Iberian vasconiensis-tobieni lineage. However, we locality Forsthart (Germany). Taking into account the are aware that the compositions of particular morphotypes larg­e ly accepted ancestor-descendant relationship of of particular teeth within European populations of the P. schnaitheimensis and Ptychoprolagus forsthartensis vasconiensis-oeningensis lineage are not yet known. and the simple not crenulated fossettes significantly Moreover, it should be noted that an evolution of pro­ prevailing in the whole material from MWQ1/2001, lagids, having considerable intra-specific and onto­ge­n­ these two specimens are regarded here as extreme etic variation, is not generally straightforward due to variants within the morphological range of the species mosaic, convergent, and parallel trends and/or changes P. schnaitheimensis. The morphology of p3 in the schnait­ (López Martínez 1989, 1997; Hordijk et al. 2010). Thus, hei­mensis-lineage seems to be quite conservative; in fact, we tentatively accept the hypothesis by Hordijk et al. the species of this clade never evolved a deep p3 centro­ (2010) that the Iberian P. lopezmartinezae lineage and flexid and p3 protoconid retained generally simple, mostly western Central European P. oeningensis lineage evolved rounded without protoconulid. from P. vasconiensis in allopatry, both reflecting ana­gene­- Specimens referred here to Prolagus cf. vasconiensis tic trends; i.e. that the Prolagus vasconiensis-P. lopez­ represent about the 11% of the material from MWQ martinezae-P. tobieni lineage is endemic to the Iberian 1/2001. The taxonomically important features in the Peninsula. In this context, we co-identify here, pro tem­ available material, such as the long and antero-posteriorly pore, the second species from MWQ1/2001 with the bent p3, very shallow p3 centroflexid with protoconid- nominal taxon described by Viret in Roman & Viret (1930) metaconid connection, small p3 anteroconid, simple from the MN3 locality of Estrepouy and refer the taxon p3 protoconid with short protoflexid and incipient under study as P. cf. vasconiensis. protoconulid, short P4–M2 shafts with a small radius of linguo-buccal curvature, and reduced simple P4–M2 Prolagus schnaitheimensis Tobien, 1975 fossettes are fully compatible with the species: Prolagus Figure 2A–I, O vasconiensis and P. lopezmartinezae. The former species, described by Viret in Roman & Viret (1930) from the Material. – One left P2 (Pal. 3071); 6 left P3 [Pal. 3072–77 MN3 locality of Estrepouy (France), was widespread (juv.)]; six right P3 (Pal. 3078–82, Pal. 3231); eight left throughout Central-Western Europe from the Czech P4 (Pal. 3083–90); three right P4 [Pal. 3091–92 (juv.), Republic to Spain (Boon-Kristkoiz & Kristkoiz 1999), and Pal. 3093]; three left M1 (Pal. 3094–96); seven right M1 further it is regarded to be the last common ancestor of all (Pal. 3097–3103); two left M2 (Pal. 3104–05); four right subsequent lineages of Prolagus which arose during the M2 (Pal. 3106–09); two left M1/M2 (Pal. 3110–11); three early Miocene (López-Martínez 1997). The latter species right M1/M2 (Pal. 3112–14); eleven left p3 [Pal. 3037–43, was described by Hordijk & van der Meulen (2010a) Pal. 3045–47, Pal. 3050 (juv.)]; three right p3 (Pal. 3044, from the early Miocene (MN4; local Zone B) locality Pal. 3048–49). of San Marcos (Spain), and at present, it is only known from Spanish localities (Hordijk et al. 2015). Hordijk & Measurements. – All the measurements are provided in van der Meulen (2010a) consider P. lopezmartinezae Tabs 1, 2. to be an intermediate chronospecies within the Iberian endemic evolution of P. vasconiensis to P. tobieni. In their Description. – All teeth are rootless. view, P. lopezmartinezae differs from P. vasconiensis in P 2 (F ig. 2A ): The sole available P2 in the lagomorph having a particular combination of features such as the P3 material from 1/2001 is damaged in its buccal part. The lagicone without a long lagiloph, less curved and generally tooth shaft is straight and relatively short. The hypercone, shorter P3 mesoflexus, and slightly more reduced enamel lagicone, and postlobus are short, all of similar length. The fossettes in the upper molars. Nevertheless, the first two paraflexus and mesoflexus are well-developed, both are features were recognized not only in the very limited relatively narrow, oriented posteriorly, the latter is longer. material from MWQ1/2001, but also in the material from The hypercone is rounded, without hypoflexus, and the type locality of Estrepouy (MN3) of P. vasconiensis, without mesial hyperloph. 102 Isaac Bonilla-Salomón et al. • Early Miocene small mammals A B C D E F G H O N M 1 mm I J K L Figure 2. Morphology of teeth in Prolagus taxa from MWQ1/2001. • A–I, O – Prolagus schnaitheimensis Tobien, 1975; A – left P2 (Pal. 3071), in occlusal and frontal views; B – left P3 (Pal. 3072), in occlusal and frontal views; C – left P4 (Pal. 3083), in occlusal and frontal views; D – left M1 (Pal. 3095), in occlusal and frontal views; E – left M2 (Pal.3104), in occlusal and frontal views; F – left p3 (Pal. 3038), in occlusal view; G – left p3 (Pal. 3039), in occlusal view; H – left p3 (Pal. 3040), in occlusal view; I – left p3 (Pal. 3037), in occlusal and buccal views; O – right M1 (Pal. 3102), in occlusal view, reversed. • J–N – Prolagus cf. vasconiensis Viret, 1930 in Roman & Viret (1930); J – right p3 (Pal. 3115), in occlusal and buccal views, reversed; K – left P3 (Pal. 3117), in occlusal and frontal views; L – left M1 (Pal. 3119), in occlusal and frontal views; M – left M2 (Pal. 3124), in occlusal and frontal views; N – right M1 (Pal. 3123), in occlusal view, reversed. Scale bars = 1 mm; grey for occlusal views, black for frontal and buccal views P 3 ( F ig. 2B): The tooth shaft (as with P4–M2) is rela­- to long. The postlobus is indented in its anterior side, t­ively short with a small radius of linguo-buccal curvature. well developed and extends distinctly on the labial side The lagicone has a simple shape and smooth outline, of the tooth. The distal hypercone is more developed often with a weak indentation at the base of the lagiloph. than the mesial one, which is slightly reduced in some The lagiloph is variably developed, from very reduced cases. The mesial hyperloph is distinctly developed and 103 Bulletin of Geosciences • Vol. 96, 1, 2021 Table 1. Measurements of upper teeth in Prolagus from 1/2001. Abbreviations: AA – partial width; CV – coefficient of variation [%]; L – length; Lpl – postlobus length; M – upper molar; N – number of specimens; OR – observed range; P – upper premolar; PH – hypoflexus width; TH – distal hyperloph length; W – width; Wdh – precone–distal hypercone width; x̄ – arithmetic mean. Prolagus schnaitheimensis Prolagus cf. vasconiensis N x̄ OR CV N x̄ OR CV P2 L 1 – 0.80 – – – – – Lpl 1 – 0.70 – – – – – W 1 – 1.47 – – – – – Wdh 1 – 0.63 – – – – – P3 L 11 1.56 1.45–1.70 4.88 3 1.46 1.40–1.50 – Lpl 10 0.92 0.80–1.00 7.34 3 0.88 0.85–0.90 – W 11 2.61 2.45–2.95 5.86 3 2.18 2.15–2.20 – Wdh 11 1.59 1.08–2.00 13.60 3 1.54 1.48–1.63 – P4 L 10 1.50 1.30–1.65 6.22 – – – – TH 10 0.82 0.75–0.90 5.78 – – – – W 10 2.49 2.40–2.60 2.65 – – – – AA 8 2.40 2.35–2.55 2.97 – – – – PH 10 0.87 0.75–1.00 8.65 – – – – M1 L 9 1.49 1.40–1.55 3.53 3 1.32 1.15–1.45 – W 10 2.32 2.20–2.58 4.86 5 2.05 1.85–2.25 7.52 AA 10 2.23 2.15–2.48 4.16 5 1.95 1.80–2.10 6.28 PH 9 1.08 0.93–1.25 8.71 5 1.26 0.95–1.55 20.33 M2 L 6 1.35 1.30–1.45 4.14 1 – 1.23 – W 6 2.18 2.10–2.25 2.77 1 – 1.80 – AA 6 2.06 2.00–2.15 2.63 1 – 1.70 – PH 6 1.01 0.95–1.05 4.35 1 – 1.40 – M1/M2 L 4 1.41 1.40–1.45 1.78 – – – – W 5 2.30 2.23–2.38 2.60 – – – – AA 2 – 2.15, 2.30 – – – – – PH 3 0.95 0.90–1.00 – – – – – long. The precone does not have enamel hiatus in 91% M1 (Fig. 2D, O): Both teeth, M1 and M2, possess a large of specimens (N = 11). The mesoflexus is of variable parafossette, whereas the mesofossette is lost. The para­ depth and curvature, in three cases it is closed into fossette in M1 is J-shaped, predominantly (80%, N = 10) mesofossette. The hypoflexus is shallow and mostly with widened and bifurcated buccal end of its anterior V-shaped. (i.e. longer) branch; in two cases this part is distinctly P 4 (F ig. 2C): Two fossettes are present in the tooth. crenulated. The hypoflexus is relatively short, it reaches The parafossette is large, wide, and exclusively J-shaped. an average of about 47% of tooth width; in vast majority The buccal end of its anterior (longer) branch is simple in (90%, N = 10) it is V-shaped in the lingual edge and 82% of specimens (N = 11), in the remaining individuals U-shaped in the labial one. In one case the hypoflexus is is bifurcated. In one case only, the anterior branch of fused with parafossette. parafossette completely covers the mesofossette. The M 2 ( F i g . 2 E ) : The parafossette is J-shaped. It is meso­fossette is large and very variable in shape, from oval less developed than in M1, i.e. shorter and predominantly (27%; N = 11) to L-shaped (27%) or C shaped (36%). The (67%, N = 6) with widened but simple (not bifurcated) hypoflexus is short, it reaches an average of about 35% of buccal end of its anterior (longer) branch. The hypoflexus tooth width, it is V-shaped in 45% of specimens (N = 11), is short and of comparable morphology as in M1. It in the remaining individuals V-shaped in the lingual edge reaches an average of about 46% of tooth width; in one and U-shaped in the labial one. case the hypoflexus is fused with the parafossette. 104 Isaac Bonilla-Salomón et al. • Early Miocene small mammals p 3 ( F i g . 2 F – I ) : Small-sized tooth. Its shaft is protoisthmid is rather wide. The entoconid is quite long, straight without antero-posterior bending and, in relation (i.e. longer than 1/ 5 of tooth length), without enamel to its occlusal size, relatively short (Fig. 2I). The available hiatus, and often indented in its anterior side. All flexids material consists of 14 p3s; one of them is a juvenile (shaft are smooth without undulation or even crenulation. with the typical conical appearance), the rest are adults (prismatic shaft). In adult p3s, four basic occlusal patterns Remarks. – Prolagus schnaitheimensis from MWQ1/2001 were distinguished, with the following combination of differs: from P. praevasconiensis Ringeade, 1979 in its features: (i) isolated anteroconid, metaconid connected via larger size, larger p3 anteroconid, and well-developed m2 metalophid with the protoconid, and occurring mesoflexid hypoconulid isolated from talonid along the full height (46% of cases; N = 13; Fig. 2F); (ii) isolated anteroconid, of the crown; from P. vasconiensis in its larger occlusal metaconid connected via metalophid with the protoconid, size of the teeth, more straight and relatively shorter p3 and occurring mesofossettid (31% of cases; N = 13; shaft, larger anteroconid of p3, simpler p3 protoconid Fig. 2I); (iii) isolated anteroconid, isolated metaconid, without protoflexid and protoconulid, P3–M2 shafts and occurring mesoflexid (15% of cases; N = 13; relatively shorter and with smaller radius of linguo-buccal Fig. 2G); (iv) anteroconid connected via anterolophid curvature, larger and more complex fossettes in P4–M2; with the protoconid, metaconid connected via metalophid from P. fortis López-Martínez & Sesé, 1991 in Álvarez- with the protoconid, and occurring mesoflexid (8% of Sierra et al. (1991) in its more straight and relatively cases; N = 13; Fig. 2H). In 12 prismatic teeth, the pattern shorter p3 shaft, larger and wider p3 anteroconid, simpler is stable along the entire tooth crown (i.e. the occlusal p3 protoconid without protoflexid and protoconulid; from pattern corresponds to the one in the root side); in one P. aguilari López-Martínez, 1997 in its less developed prismatic specimen the occlusal surface has pattern (i), p3 centroflexid, protoflexid, and protoconulid; from whereas the root side pattern (ii); the same occurs in the Ptychoprolagus forsthartensis Tobien, 1975 in its much sole juvenile specimen of the sample. The occlusal outline less developed enamel crenulation in upper teeth; from is not elongated (L/W = 0.96). The anteroconid is large, P. lopezmartinezae Hordijk & van der Meulen, 2010a in larger than metaconid, predominantly of oval shape, its larger size of teeth, larger, and wider p3 anteroconid, widened antero-lingually. The centroflexid is incipient, simpler p3 protoconid without protoflexid and proto­ very shallow. The protoconid is simple and rounded; conulid, simpler P2 hypercone, more developed P3 the protoflexid and protoconulid are not developed. The lagiloph, larger fossettes in P4–M2, shorter hypoflexus Table 2. Measurements of lower teeth in Prolagus from 1/2001. Abbreviations: CV – coefficient of variation [%]; L – length; La – anteroconid length; Lme – metaconid–entoconid length; Ltrig – trigonid length; m – lower molar; N – number of specimens; OR – observed range; p – lower premolar; W – width; Wa – anteroconid width; Wtl – third lobe width of m2; Wtal – talonid width; Wtrig – trigonid width; x̄ – arithmetic mean. Prolagus schnaitheimensis Prolagus cf. vasconiensis N x̄ OR CV N x̄ OR CV p3 L 12 1.71 1.63–1.85 4.16 1 – 1.35 – Lme 13 1.22 1.13–1.27 3.12 1 – 0.97 – La 12 0.59 0.53–0.67 7.11 1 – 0.40 – W 12 1.77 1.63–1.93 4.61 1 – 1.22 – Wa 12 0.92 0.75–1.00 9.04 1 – 0.50 – Prolagus schnaitheimensis seu P. cf. vasconiensis N x̄ OR CV p4/m1 L 14 1.11 0.93–1.27 10.28 Ltrig 14 0.62 0.57–0.68 7.24 Wtrig 14 1.12 1.00–1.25 6.53 Wtal 14 1.14 0.97–1.30 8.31 m2 L 6 1.57 1.33–1.70 8.63 Wtrig 7 1.06 0.93–1.17 7.91 Wtal 9 1.04 0.85–1.15 8.65 Wtl 8 0.73 0.60–0.87 12.61 105 Bulletin of Geosciences • Vol. 96, 1, 2021 in M1–M2; from P. oeningensis (König, 1825) in its with a size comparable to the anteroconid, and anteriorly larger anteroconid of p3, simpler p3 protoconid without concave. The centroflexid is incipient, very shallow. The protoflexid and protoconulid, absence of long centroflexid, protoconid is short and rounded, anteriorly with very short larger fossettes in P4–M2, shorter hypoflexus in M1–M2; protoflexid and incipient protoconulid. The hypoflexid is from P. vargasensis Hordijk & van der Meulen, 2010b long, strongly inclined towards the posterior side of tooth. in its larger and wider p3 anteroconid, and presence of The protoisthmid is rather narrow. The entoconid is quite protoconid-metaconid connection in p3. long, (about a third of the tooth length), without enamel hiatus and with antero-buccal protrusion. The mesoflexid Prolagus cf. vasconiensis Viret, 1930 in Roman & is very long and L-shaped. Viret (1930) Figure 2J–N Remarks. – Prolagus cf. vasconiensis from MWQ1/2001 differs: from P. praevasconiensis in its more complicated Material. – Two left P3 (Pal. 3116–17); one right P3 (Pal. p3 protoconid and well-developed m2 hypoconulid 3118); one left M1 (Pal. 3119); four right M1 (Pal. 3120–23); isolated from the talonid along the full height of the crown; one left M2 (Pal. 3124); one right p3 (Pal. 3115). from P. oeningensis in its smaller size, absence of long p3 centroflexid, reduced P3 lagiloph; from P. vargasensis Measurements. – All the teeth measurements are provided in its smaller size, presence of protoconid-metaconid in Tabs 1–2. connection in p3, shorter P3 lagiloph; from P. fortis in its smaller size, shorter P3 lagiloph; from P. aguilari Description. – All teeth are rootless. in its smaller size, smaller and relatively narrower p3 P 3 ( F ig. 2K ): The tooth shaft is relatively long with anteroconid, less developed p3 centroflexid, protoflexid, a large radius of linguo-buccal curvature. The lagicone and protoconulid; from P. schnaitheimensis in its smaller has a simple shape and smooth outline. The lagiloph is occlusal size of teeth, relatively longer, and p3 shaft more very reduced. The postlobus is short and does not extend curved antero-posteriorly, smaller anteroconid of p3, widely on the labial side of the tooth. The pointed distal more complex p3 protoconid with incipient protoflexid hypercone is more developed than the rounded mesial and protoconulid, relatively longer P3–M2 shafts with one. The mesial hyperloph is distinctly developed and larger radius of linguo-buccal curvature, P4–M2 with very long. The precone does not have enamel hiatus. The smaller, narrower, and less developed fossettes; from mesoflexus is short and slightly curved. The hypoflexus is Ptychoprolagus forsthartensis in its much less developed shallow and mostly V-shaped. enamel crenulation in upper teeth. The very scanty M 1–M 2 (F ig. 2L–N ): The tooth shafts are relatively material of the taxon under comparison does not show any long with a large radius of linguo-buccal curvature. The significant differences in size and morphology of teeth mesofossettes are missing. The parafossettes are small from those found in the central European populations of (compared to P. schnaitheimensis) and towards the P. vasconiensis and the Iberian species P. lopezmartinezae posterior end of the tooth row are progressively more (see “Discussion to the lagomorph material” for justifi­ reduced; from J- to small oval-shaped. The anterior branch cation of the species attribution). of the parafossettes are narrow and simple, without any bifurcation. The length of the hypoflexus is very variable, Prolagus schnaitheimensis seu P. cf. vasconiensis though it is possible to individuate two distinct variants; (i) four of six teeth have a long hypoflexus reaching about Material. – Seven left p4/m1 (Pal. 3051–55, Pal. 3232–33); two thirds of W (Fig. 2L–M), (ii) two of six teeth have seven right p4/m1 (Pal. 3056–62); three left m2 a relatively short hypoflexus reaching about half of W [Pal.3063–64 (without trigonid), Pal. 3234]; six right m2 (Fig. 2N). The latter variant is associated with a larger and [Pal. 3065–69 (without hypoconulid), Pal. 3070 (without more centrally positioned parafossette (Fig. 2N). trigonid)]. p 3 ( F i g . 2 J ) : A sole specimen is available in the lagomorph material from MWQ1/2001. The tooth shaft Measurements. – See Tab. 2. is rootless, prismatic, antero-posteriorly bent, and, in relation to its small occlusal size, relatively long. The Description. – p 4 – m 1 : Rootless tooth consisting of occlusal outline is elongated (L/W = 1.10). The tooth trigonid and talonid fused together by cement. Both morphology can be assimilated to pattern I (see Remarks lobes are of similar width and posteriorly have thickened of P. schnaitheimensis), and it is stable along the entire enamel. The trigonid is of rhomboid shape and longer than tooth crown. The anteroconid is diamond-shaped and the talonid. The talonid possesses a variably developed small, lying in a rather central position with respect to anterior protrusion having an antero-labial flexid and the tooth’s antero-posterior axis. The metaconid is small often a shallow anterior notch. 106 Isaac Bonilla-Salomón et al. • Early Miocene small mammals m 2 : The three-lobed, rootless tooth has the trigonid Description. – m1 (F ig. 3A ): The anteroconid complex and talonid similar to those in p4–m1 with the addition of protrudes mesially. A deep well-marked notch isolates a hypoconulid that is fully developed and isolated from both anteroconids. The labial anteroconid shows one the talonid along the total height of the crown. The hypo­ posterior spur that connects to a massive anterolophulid conulid is variable in shape, but generally simple and oval. cuspid. The lingual anteroconid shows two posterior spurs: one connecting to a mesial spur from the metaconid, Remarks. – Lower molariforms do not bear discriminant likely the metalophulid, and the other one attaching to features allowing a clear distinction between the above the anterolophulid cuspid. The labial anterolophulid described prolagid taxa, therefore, the available material does not connect to the labial anteroconid, and it shows of lower molariforms from MWQ1/2001 is referred here a small thickening that constitutes the labial anterostylid. to Prolagus schnaitheimensis seu P. cf. vasconiensis. The protosinusid is open. The metaconid shows two well-defined spurs running distolingually from it. The Order Rodentia Bowdich, 1821 protoconid hind arm reaches the most labial spur from the Family Cricetidae Fischer [de Waldheim], 1817 metaconid, which joins with a similar spur running antero- Subfamily Cricetopinae Matthew & Granger, 1923 lingually from the entoconid, leaving a small sinusid Tribe Melissiodontini Schaub, 1925 open lingually. The mesostylid is weak but present. The mesoconid is very large. It has a short mesolophid that Genus Melissiodon Schaub, 1920 ends without reaching the mesial spur of the entoconid. The mesoconid is also connected by a short anterior spur Melissiodon dominans Dehm, 1950 to the protoconid hind arm. A well-developed ectolophid Figure 3A–F connects with the mesoconid. The sinusid is M-shaped and open labially. The posterolophid shows a hypoconulid- Material and measurements (L × W). – One left m1 (Pal. like bulge on its most distal side. The entoconid shows 3005: 2.39 × 1.9); four left m2 (Pal. 3000: 2.28 × 1.79; two short posterior spurs. Pal. 3002: 2.25 × 1.75; Pal. 3003: 2.28 × 1.79; Pal. 3004: m2 (F ig. 3B–F ): The outline of the teeth is rectan­- 2.23 × 1.68); one right m2 (Pal. 3001: 2.2 × 1.71). gu­lar. From the protoconid departs the anterolophulid, A B 1 mm C D E F Figure 3. Scanning electron micrographs of Melissiodon dominans Dehm, 1950 from MWQ1/2001 in occlusal view. A – left m1 (Pal. 3005); B – left m2 (Pal.3003); C – left m2 (Pal. 3000); D – left m2 (Pal. 3002); E – left m2 (Pal. 3004); F – right m2, reversed (Pal. 3001). 107 Bulletin of Geosciences • Vol. 96, 1, 2021 longer in Pal. 3002–03, and connects to a well-developed Subfamily Cricetodontinae Schaub, 1925 anteroconid. The protosinusid is open. The labial ante­ro­ lophid connects to the metalophulid. The anterosinusid Genus Megacricetodon Fahlbusch, 1964 is open labially. The protoconid hind arm is long and connects with two long spurs departing from the metaconid Megacricetodon sp. in Pal. 3003; in the other specimens the more labial spur Figure 4A, B does not reach the protoconid hind arm. The mesoconid links to the protoconid hind arm. The mesolophid connects Material and measurements (L × W). – Left maxillary to the protoconid hind arm enclosing a very small sinusid, fragment with M1 (Pal. 3013: 1.49 × 0.93); one fragment different in each tooth: very short and arched in Pal. of left maxilla with M1 (Pal. 3014: 1.5 × 0.98) and M2 3000 (Fig. 3C); slightly longer and arched in Pal. 3001 (Pal. 3015: 1.06 × 0.94). (Fig. 3F); and longer enclosing a wider sinusid in Pal. 3002–03. In specimen Pal. 3004 (Fig. 3E), the mesolophid Description. – M 1 ( F i g . 4 A , B ) : The anterocone is is reduced to a very short spur running lingually from very large, with a small notch that does not completely the mesoconid. The sinusid is open labially despite divide the cusp into two. There is a small platform in the low cristids running distally from the protoconid front of the anterocone. A short but thick anterolophule and mesially from the hypoconid. There is a hint of connects the lingual anterocone with the protocone. a very short hypolophulid in Pal. 3000 and Pal. 3003, Both cusps are also linked by a low lingual anteroloph. and a longer one that does not connect to the entoconid in Pal. 3014 shows a well-marked protostyle, whereas Pal. Pal. 3002. The entoconid shows a well-developed 3013 (Fig. 4A) only shows a small thickening. The labial anteriorly directed spur that attaches to the protoconid anteroloph connects the anterocone with the paracone, hind arm, delimiting a small sinusid open labially. The showing a very small labial anterostyle, and encloses the posterolophid is long; in Pal. 3001 shows a thickening anterosinus. The protocone connects with the paracone that constitutes a distinct hypoconulid. The entoconid by means of a very short protolophule. Pal. 3014 shows presents two short posterior spurs, the lingual one joins a very short forward paracone spur that does not connect the posterolophid. to a short labial spur of the anterolophule. The mesoloph reaches the labial edge of the tooth in Pal. 3014 (Fig. 4B) Remarks. – Melissiodon dominans Dehm, 1950 is a widely and ends close to the base of the metacone in Pal. 3013. studied species known exclusively from Europe ranging There is no distinct mesostyle. A small entostyl is present from the early Oligocene to the latest early Miocene. at the base of the hypocone, much more marked in Pal. During the Miocene, only two Melissiodon species are 3013. The hypocone is connected to the protocone by known: M. schlosseri and M. dominans. Melissiodon a lingual mesocingulum and to the metacone through the arambourgi Crusafont et al., 1955, another Miocene posteroloph. The mesosinus is shallow, narrower in Pal. species of the genus described from the Spanish Miocene 3014. The posterosinus is narrow and open distally. site of el Fallol, has been recently reviewed by Jovells- M 2 (F ig. 4B): The outline of the tooth is rectangular. Vaqué & Casanovas-Vilar (2018) and synonymized with The lingual cusps are more abraded than the labial ones. M. dominans. The m1 from MWQ1/2001 lies within The anterosinus is very narrow and there is no protosinus. the size variability range of M. dominans from all three The protocone is the largest cusp and is connected to the Dolnice localities and is slightly smaller than the sole paracone by a short anterior protolophule. Protocone and tooth measured from Ořechov (Fejfar 1990). Melissiodon hypocone are connected by a short entoloph and a low dominans from MWQ1/2001 is similar in size to lingual mesocingulum. The sinus is deep and wider than populations of the MN4 of Germany, although the m2 the protosinus. The mesoloph is short and running mesio- is somewhat narrower than the material from Forsthart l­abially. The labial mesocingulum is short, with absence (MN4). Moreover, the m1 fits in the shorter range of the of mesostyle. The metalophule runs slightly anteriorly type material from Wintershof-West (MN3; Ziegler & and connects to the hypocone. The distal cusps are also Fahlbusch 1986), and it is notably smaller than the sole connected by a low posteroloph, which is more developed m1 from Maigen, a MN3 Austrian locality (Mein 1989). than in M1s. The posterosinus is wider than in M1s. The m1 of M. dominans from MWQ1/2001 slightly differs from the m1 material from Maigen, e.g. the mesoconid is Remarks. – The genus Megacricetodon, first defined by weaker and the anterolophid cuspid is absent. Altogether, Fahlbusch (1964) as a subgenus within Democricetodon, the ascription of the material to M. dominans is supported is known in Europe from the early to the late Miocene. by both metrics and morphology, like the two robust Peláez-Campomanes & Daams (2002) define three anterior cuspids in the m1 and the reduced mesolophid in different groups of Megacricetodon species, according the lower molars. to their size. The small-sized group, where the available 108 Isaac Bonilla-Salomón et al. • Early Miocene small mammals A 1 mm B Figure 4. Scanning electron micrographs of Megacricetodon sp. from MWQ1/2001 in occlusal view. A – left M1 (Pal. 3013); B – fragment of mandible with left M1 (Pal. 3014) and M2 (Pal. 3015). material of Megacricetodon sp. from MWQ1/2001 this genus have been proposed by Oliver & Peláez- allocates, includes: M. minor Lartet, 1851, M. collongensis Campomanes (2014, p. 953), with a strong geographic Mein, 1958, M. debruijni Freudenthal, 1968, M. primitivus component. According to Oliver & Peláez-Campomanes Mein & Freudenthal, 1971, M. sinensis Qiu et al. 1981, (2016), three different lineages are proposed based on M. pusillus Qiu, 1996, and the recently discovered three different migration events. The first one corresponds M. hellenicus Oliver & Peláez-Campomanes, 2016. to the appearance of the small-sized M. hellenicus Megacricetodon pusillus and M. sinensis are restricted from Aliveri (earliest MN4), evolving as an endemic to Asia, and M. debruijni belongs to late Miocene Greek form. The second dispersal event involves the faunas. Furthermore, there is no record of M. minor in M. bavaricus Fahlbusch, 1964 group, found in different MN4 faunas, as it appears in MN5 Central European Central European basins. Although the localities Dolnice assemblages, such as in Apistobel 520 m and Mühlbach 3 and Ořechov are similar to Aliveri in age (Fejfar (Swiss and Austrian sites, respectively). Kälin & Kempf 1990), the Czech Megacricetodon cannot be assigned (2009) also mentioned a very small new species of to the Greek species due to its larger size and different Megacricetodon from Apistobel 520 m and older MN5 dental morphology. The third dispersal event involves localities, which may belong to the same lineage of M. primitivus, so far restricted to southwestern Europe. M. minor. Therefore, the Megacricetodon from MWQ According to Oliver & Peláez-Campomanes (2016), the 1/2001 compares with M. primitivus, M. collongensis and M. bavaricus group has the earliest representatives from M. hellenicus. the Czech assemblages such as Ořechov and Dolnice 3, The two M1 from MWQ1/2001 fit well within the which are related to the small-sized M. aff. collongensis dimensional range of M. hellenicus from Aliveri North and (Swiss and German forms), and M. bezianensis from Aliveri South (Fig. 5), and the M2 is somewhat smaller France. The material from MWQ1/2001 is situated than the sole M2 recovered from Aliveri North. Despite slightly below the size range of M. aff. collongensis from the similarity in size, there are some differences that Langenau 1 (Fig. 5), but it fits on the lowest range of the prevent the assignation of the material from MWQ1/2001 samples from Forsthart and Rauscherod, all of them MN4 to M. hellenicus, such as the not so well divided localities from Germany. Compared with the other Czech anterocone in M1, the presence of a small cingulum assemblages, Megacricetodon sp. from MWQ1/2001 is in front of the anterocone surrounding it, and the short somewhat larger than the sole M2 from Ořechov, but its mesoloph and straight entoloph in M2. The material from morphology shows some clear differences: the presence of MWQ1/2001 also fits well within the range of the small mesostyle and posterior spur running from the protocone, Megacricetodon from Antonios (Vasileiadou & Koufos and the longer protolophule in the specimen from 2005), but it differs from the latter in the not completely Ořechov (see Fejfar 1974, fig. 29, nr. 21). Both M1 and divided anterocone, the less developed anterior cingulum, M2 from Dolnice 3 are noticeably larger than the material the shorter anterolophule, the less developed paracone from Mokrá-Quarry locality (Fig. 5), moreover there are spur in the M1, and the short mesoloph in the M2. some marked differences between both assemblages: the Megacricetodon sp. from MWQ1/2001 differs from completely split anterocone and a paracone spur running M. primitivus from western European localities for the distally that connects to the mesoloph, which are present absence of a completely split anterocone in M1, and in the material from Dolnice 3. the absence of a strong mesocingulum running along The scarce material from MWQ1/2001 is noticeably the hypocone to the anterocone. Several lineages of smaller than the rest of assemblages from the Czech 109 Bulletin of Geosciences • Vol. 96, 1, 2021 A B Figure 5. Scatterplot (length and width) of first and second upper molars of several species of Megacricetodon from various localities; A – upper M1; B – upper M2. Type locality and number of specimens is indicated next to locality and species name. In the upper M1, the localities of Antonios and Bezian (M. primitivus) show the full range of length and width, but the means are not represented. Measurements of Megacricetodon species were obtained from the following publications (Fahlbusch 1964, Fejfar 1974, Bulot 1980, Ziegler & Fahlbusch 1986, Vasileiadou & Koufos 2005, Oliver & Peláez-Campomanes 2014, Oliver & Peláez-Campomanes 2016; Volker J.S. unpublished data). 110 Isaac Bonilla-Salomón et al. • Early Miocene small mammals Republic and other Central European localities (Fig. 5), Description. – P 4 ( F i g . 6 A ) : The outline of the tooth all of them referred to the M. bavaricus group (Peláez- is triangular. The parastyle is well-developed; a very low Campomanes & Daams 2002). The Megacricetodon anteroloph runs from it, connecting to the base of the material from the rest of fissures from Mokrá-Quarry protocone. Protoloph and metaloph converge towards is still under study by one of the authors (I. Bonilla- the protocone. The protoconule is much smaller than the Salomón) and will help elucidate the phylogeny of the metaconule, which is massive. The metaloph is lingually group as well as its different dispersal events during the constricted. The mesostyle is small and isolated from the early Miocene of Central Europe. Basing on all the above labial cusps. The hypocone is very reduced, integrated stated information, we decided to tentatively attribute the into the posteroloph. scarce material from MWQ1/2001 to Megacricetodon sp. M 1 ( F i g . 6 A – C ) : The protocone is very large, and from it runs a thin and long anteroloph (except in Pal. Family Sciuridae Fischer [de Waldheim], 1817 3010, in which it is thicker), which reaches the base of Subfamily Pteromyinae Brandt, 1855 the paracone without connecting to it. Both cusps are con­- nected through the protoloph, which thickens into a proto­ Genus Aliveria de Bruijn, van der Meulen, & conule in Pal. 3008. The mesostyle is well-developed and Katsikatsos, 1980 isolated from the labial cusps. The metaloph runs towards the protocone, with an especially thick and elongated Aliveria aff. luteyni de Bruijn, van der Meulen, & metaconule in Pal. 3006 (Fig. 6B). A short endoloph Katsikatsos, 1980 connects the protocone with a not so well-developed Figure 6A–F hypocone. The posteroloph is low, leaving the sinus open. M 2 (F ig. 6D ): The outline of the tooth is square. There Material and measurements (L × W). – One left P4 is no parastyle. Protocone and paracone are connected (Pal. 3009: 1.75 × 1.87); two left M1 (Pal. 3010: 1.78 × through a low anteroloph and a well-developed protoloph. 2.02; Pal. 3008: 1.84 × 2.18); one right M1 (Pal. 3006: There is a thickening on the latter that could constitute 1.74 × 2.02); one right M2 (Pal. 3007: 1.9 × 2.15); one a distinct protoconule. There is a worn but distinct left m1 (Pal. 3011: 1.76 × 1.95); one left m3 (Pal. 3012: mesostyle attached to the paracone. The metaloph runs 2.19 × 1.98). mesiolingually to the protocone and shows a well- A B C 1 mm D E F Figure 6. Scanning electron micrographs of Aliveria aff. luteyni de Bruijn, van der Meulen, & Katsikatsos, 1980 from MWQ1/2001 in occlusal view; A – fragment of left maxilla with P4 (Pal. 3014) and M1 (Pal. 3015); B – right M1, reversed (Pal. 3006); C – left M1 (Pal. 3008); D – right M2, reversed (Pal. 3007); E – left m1 (Pal.3011); F – left m3 (Pal. 3012). 111 Bulletin of Geosciences • Vol. 96, 1, 2021 developed and single metaconule. The hypocone is reduced hypocone in M1–2, despite several features reduced, only constitutes a small bulge fused with the are markedly different. The material from MWQ1/2001 endoloph that connects to the protocone. The posteroloph differs from A. luteyni from Aliveri in the presence of runs very low, leaving the sinus open. a very large parastyle on the P4; an elongated metaconule m 1 ( F i g . 6 E ) : The protoconid is round, large and in M1–2, instead of double like in the specimens from attached to a well-developed anteroconulid, which is the type locality; a well-marked mesostyle, isolated also connected to the base of the metaconid by a short from the paracone in the upper molars; a more rounded anterolophid. Metaconid and protoconid are also con­ protocone; absence of a complete metalophid on m1–2. nected by a very low and weak metalophid. There is Despite the similar size of the material from MWQ1/2001 a small but distinct mesostylid, attached to the metaconid with Aliveria luteyni, the morphological differences stated and isolated from the entoconid by a well-marked notch. above prevents the assignation of these sciurid remains The ectolophid is low and continuous, connecting the two to this species. However, the scarcity of the remains labial cuspids and enclosing the central valley. There is recovered from MWQ1/2001 prevents the erection of a massive mesoconid. The hypoconid is well-developed a new species. For that reason, it is assigned to Aliveria although abraded. From it a well-developed posterolophid aff. luteyni. connects with the entoconid, which is the smallest cuspid. The entolophid is absent. The posterolophid is higher on Order Erinaceomorpha Gregory, 1910 its lingual side. Family Erinaceidae Fischer [de Waldheim], 1817 m3 (F ig. 6F ): This tooth is slightly longer than m1, Subfamily Galericinae Pomel, 1848 with a more rounded distal side. The enamel is slightly crenulated. The protoconid shows a short spur running Genus Galerix Pomel, 1848 lingually from it. The anterolophid is high and connects to the metaconid. There is no anteroconid. There is a small Galerix sp. but distinct mesostylid, connected to the metaconid but Figure 7A–D isolated from the entoconid by a pronounced notch. The ectolophid connects both labial cuspids and shows a well- Material and measurements (L × W). – One right M2 (Pal. developed mesoconid. The hypoconid is massive and 3016: 2.13 × 2.69); one fragment of left M2 (Pal. 3017); connects to the entoconid by a well-developed and thick two left m2 (Pal. 3018: 2.56 × 1.56; Pal. 3019: 2.72 × 1.65). posterolophid. Description. – M 2 ( F i g . 7 A , B ) : The crown is only Remarks. – Aliveria de Bruijn, van der Meulen, & Katsi­ slightly worn, almost undamaged in Pal. 3016. In Pal. katsos, 1980 is a genus of medium-sized flying squirrels, 3017 (Fig. 7B) only the distal half is preserved. There described from the locality of Aliveri. Two different is a very small preparacrista running mesiolabially from species have been described so far from the type locality: the paracone, but here is no distinct parastyle. The postpa­ A. luteyni and A. brinkerinki. Both species differ mainly rac­rista joins the premetacrista; a mesostyle is absent. It in size, the latter being larger (de Bruijn et al. 1980). has a U-shaped postmetacrista running to the distobuccal Aliveria has been mostly recorded in several sites from the corner of the tooth. At the end of it there is a small worn Anatolian region and Balkan area (Doukas 2003, Bosma metastyle, which is almost unnoticeable in Pal. 3017. At et al. 2019) but remains outside this area are scarce. the base of the metacone there is a thin crest that connects Besides the type locality it has only been found out of the it to a large and round metaconule. The protocone is large; Anatolian region in Sibnica (Marković et al. 2016).The the preprotocrista running from it does not connect to the sciurid material recovered from MWQ1/2001, formerly base of the paracone. The trigon basin enclosed is deep. attributed to Palaeosciurus sp. (Sabol et al. 2007) is The hypocone connects through the prehypocrista to the ascribed to the genus Aliveria based on the presence of postprotocrista. There is a well-marked concavity on the a well-delimited hypocone in M1–2, the convergence of lingual outline of the tooth, between the lingual cusps. protoloph and metaloph towards the protocone, and the Both mesial and distal cingula are well-developed. presence of a well-developed anteroconid in the lower m 2 ( F i g . 7 C , D ) : The crown is slightly worn. The molars. The sciurid remains from MWQ1/2001 fit within lingual cuspids are somewhat higher than the labial ones. the size range of A. luteyni from Aliveri and it is clearly The paraconid is completely merged into the paralophid smaller than A. brinkerinki from the same locality. It is in Pal. 3019 (Fig. 7D) and only constitutes a small bulge also somewhat larger than the material from Sibnica 4 in Pal. 3018. The paralophid is round and runs from the assigned to Aliveri aff. luteyni by Marković et al. (2016). It protoconid towards the base of the metaconid but without shows closer affinity to A. luteyni, like the m3 morphology reaching it. In Pal. 3019, the protoconid and metaconid from Klapangi 1 (Bosma et al. 2019, fig. 7d) and the are also connected by a strong protolophid that links to 112 Isaac Bonilla-Salomón et al. • Early Miocene small mammals 1 mm A B C D Figure 7. Scanning electron micrographs of Galerix sp. from MWQ1/2001 in occlusal view; A – right M2, reversed (Pal. 3016); B – fragment of left M2 (Pal. 3017); C – left m2 (Pal. 3018); D – left m2 (Pal. 3019). the metalophid. There is no metacristid. The postcristid recognized morphological and dimensional similarities connecting the hypoconid with the entoconid is much with G. symeonidisi, for a precise attribution of the lower in Pal. 3019. Galerix remains at the species level, the morphology of the upper and lower premolars has to be taken into account. Remarks. – The clearly wider than longer M2 morphology, Given that those teeth are missing in the assemblage from together with the crescent shape of the metaconule, MWQ1/2001, the material is here referred to Galerix sp. is indicative of the tribe Galericini, which is very common in the Miocene of Europe. The genus Galerix Order Chiroptera Blumenbach, 1779 is characterized by the posterior arm of the metaconule Family Rhinolophidae Gray, 1825 not reaching the posterolabial side of the tooth, feature that is clearly present in the material from MWQ1/2001. Genus Rhinolophus Lacépède, 1799 During the early Miocene, two species of Galerix are present in Central Europe: G. symeonidisi Doukas, 1986 Rhinolophus cf. cluzeli Hugueney, 1965 and G. aurelianensis Ziegler, 1990. The sole complete Figure 8A–C M2 from MWQ1/2001 is clearly smaller than that of G. aurelianensis from Erkertshofen 2 and Petersbuch 2, Material and measurements. – Left humerus [Pal. 3036: whereas the two m2 fit on the smaller range of the sample 3.70 (width of epiphysis), 1.43 (width of epicondyle), 1.53 from the above-mentioned localities (Ziegler 1990). (capitulum width), 1.3 (width of diaphysis)]; left mandible Compared with the sole m2 recovered from the Czech site with (p3-root), p4 (Pal. 3035: L 1.36 × W 1.03), m1 (Pal. of Ahníkov/Merkur north Mine (MN3), the sample from 3844: L 2.05 × Wtr 1.4, Wtl 1.45), and m2 (Pal. 3845: L 2.06 × MWQ1/2001 is somewhat narrower (see van den Hoek Wtr 1.35, Wtl 1.43). Ostende & Fejfar 2006). Besides the size differences, G. aurelianensis usually has a connection between the Description. – H u m e r u s ( F i g . 8 A ) : The preserved protocone and the metaconule (Ziegler 1990), which is portion of the humerus (distal epiphysis) shows all the absent in the M2 and fragment of M2 from MWQ1/2001. characters of large-sized horseshoe bats: broad medial Compared with G. symeonidisi from other Central epicondyle with processus spinosus (that is broken in European localities, the M2 from MWQ1/2001 is larger Pal. 3036); sharp medial margin of trochlea; and broad than the material from Erkertshofen 2, in fact, it fits in the capitulum extending to a cylindric lateral epicondyle. upper size range of the material from Erkertshofen 1 and Compared to R. delphinensis from Devínska Nová Ves, the in the lower size range of the specimens from Petersbuch lateral epicondyle is relatively narrow, but its diameter is 2 (Ziegler & Fahlbusch 1986). The M2 is similar in close to the one of the capitulum, while its lateral fossa is size to the specimens of G. symeonidisi from the late deeper with a slit-like proximal opening (as in R. kowalskii). early Miocene localities from Germany, e.g. Fortshart, M a n d i b u l a r f r a g m e n t ( F i g . 8 B , C ) : Minute p3 Rembach and Rauscheröd (Ziegler & Fahlbusch 1986). (root) is displaced at labial margin of p4, molariform Both m2s are larger than those from the above-mentioned dentition is robust, molars are nyctalodont with sharp German late early Miocene sites. In spite of the above entoconid crests. 113 Bulletin of Geosciences • Vol. 96, 1, 2021 p4 (F ig. 8B, C): the base of the tooth is rectangular. from several Miocene sites, including the specimens The cuspid is pyramidal in shape, sharply pointed and from its type locality (La Grive) and with R. cluzeli from separated by vertical distal wall from spacious distal Vieux-Collonges (both kindly provided to IH by Pierre cingular basin with conspicuous incision in its labial base. Mein). Significant differences have been found in the Both posterior and lingual faces are roughly concave. shape of p4 between the mandibles from MWQ1/2001 A cingulid surrounds the base of the tooth, which is and R. delphinensis, but a there is instead a good match thicker on its distal side. between the mandibles from MWQ1/2001 and R. cluzeli. m1 (F ig. 8B, C): The tooth is only slightly worn. The The morphology of the humerus of R. aff. lemanensis protoconid is massive, being the highest cuspid; it connects from Wintershof-West figured by Ziegler (Ziegler 1993, to the paraconid and a distinct low staying paracristid pl. 3.3), is almost identical with Pal. 3036 from Mokrá through a narrower preprotocristid. Correspondingly, MWQ1/2001, particularly as concerns its heavily build a thicker postprotocristid connects protoconid to the meta- lateral epicondyle, yet dimensions of the latter are much conid and a short but thick metacristid. The precingulid is smaller (W = 3.70 cm in Pal. 3036 vs. 4.86–5.68 cm in wide and connects to the postcingulid by means of a well- Wintershof-West). developed labial cingulid. The entoconid is thinner than the other cuspids and links to the base of the metaconid Rhinolophus cf. grivensis (Depéret, 1892) through a preentocristid that runs along the lingual margin Figure 8D–F of the tooth. The hypoconid is very large but lower than the protoconid, and it is connected to it by a straight crista Material and measurements. – Right mandible with m2 obliqua. From the hypoconid runs a sharp postcristid that (Pal. 3033: L 1.34 × Wtr 0.74, Wtl 0.89) and m3 (Pal. 3846: attaches to a small hypoconulid. L 1.31 × Wtr 0.87, Wtl 0.89); toothless fragment mandible m 2 ( F i g . 8 B – C ) : The tooth is nearly unworn. The (Pal. 3034). protoconid is thicker than the m1 while the trigonidal fossid is somewhat narrower than in m1. The cingulid Description. – Both mandibles show nearly identical runs continuously from the anterior part of the tooth to the alveolar configuration: a very large rounded alveolus of hypoconulid. The entoconid is small and attaches to the p2; relatively large alveolus of p3 not compressed and base of the metaconid through the sharp preentocristid. occupying a labial half of toothrow between p2 and p4; The crista obliqua connects the hypoconid with the relatively long p4; and equal size of all molars. The teeth distolingual side of the base of the protoconid. The in mandible Pal. 3033 (Fig. 8D–F) are strongly worn, postcristid is even sharper than in m1 while hypoconulid nyctalodont, with distinct entoconid crests and indistinct is somewhat less distinct. hypoconulids connected by a straight postcristids to mas­ sive hypoconids. Trigonids on both teeth are relatively Remarks. – In Miocene karstic sites, rhinolophids are narrow, with straight mesial walls. Compared to the high often the only component of the bat fossil record (Ziegler and pointed metaconids, the paraconids are much less 1993, 2003). Typically, this clade is represented there with distinct. The base of trigonidal fossid is low and marked a large-sized form resembling the extant Rhinolophus by a distinct basal cingulid. The protoconid is the highest ferrumequinum. As a rule, it is coidentified with R. delph­i­- cuspid and connects to the paraconid and the metaconid nensis Gaillard, 1899 from the MN7+8 site of La Grive. through the protocristid and a thick metacristid and thinner This species was reported from at least 15 Miocene sites paracristid. The cristid obliqua connecting the hypoconids (MN3–MN13). Besides, two other species of the same with the distal base of protoconids are straight, without size category were distinguished: R. lemanensis Revilliod, any undulation. Hypoconids are heavily worn. In both 1920 (type locality MN2, Saint-Gérand-le-Puy) reported teeth, the talonids are noticeably broader and larger in also from Winterhof-West and three other sites of square than the trigonids. In contrast to the extant species MN3–MN4 age (Ziegler 1993); and R. cluzeli Hugueney, of R. euryale, R. hipposideros or R. lepidus groups, the 1965 from Coderet-Branssat (MP30) and Vieux- m3 of the specimens Pal. 3033 shows almost no trace of Collonges (MN4: as R. aff. delphinensis in Mein 1958). reduction, except for a slight medial shift in position of its Both these forms differ from R. delphinensis by a lesser entoconid. degree of premolar reduction and a slenderer form of p4 with a sharply pointed main cuspid and an enlarged Remarks. – The small-sized representative of the genus distal cingulid basin with ventral flexion of its labial in the present sample. is tentatively attributed to R. gri­ base. Rhinolophus lemanensis is somewhat larger than vensis (including similis Zapfe, 1950), the species R. delphinensis and R. cluzeli appears at the lower range reported from 13 Miocene (MN3–MN10) sites. Both the of R. delphinensis metric variation. We directly compared two specimens from Mokrá show distinctly smaller di­ the specimen from MWQ1/2001 with R. delphinensis mens­ions than R. dehmi Ziegler, 1993 from Wintershof- 114 Isaac Bonilla-Salomón et al. • Early Miocene small mammals A 1 mm B C D E F Figure 8. A, C – scanning electron micrographs and drawings of Rhinolophus cf. cluzeli Hugueney, 1965 from MWQ1/2001 in occlusal view; A – left humerus (Pal. 3036); B, C – left mandible with (p3), p4 (Pal. 3035), m1 (Pal. 3844) and m2 (Pal. 3845). • D, F – scanning electron micrographs and drawings of Rhinolophus cf. grivensis from MWQ1/2001 in occlusal view; D, E – right mandible with m2 (Pal.3033) and m3 (Pal. 3846); F – toothless fragment mandible (Pal. 3034). West but fall in the dimensional range of R. grivensis Merkur Mine. Regarding the MN4, there are three levels of as reported for populations of several early to middle Dolnice, while in South Moravia Region two localities are Miocene sites by Ziegler (2003). The low degree of m3 reported: Ořechov and Mokrá-Quarry. On the preliminary reduction conspicuous in our specimen and supporting study devoted to small mammals from MWQ1/2001 by the suggested identification is also worth of mentioning in Sabol et al. (2007), the MN4 age for this karst fissure was these connections. proposed. It was based mainly on the co-occurrence of Anyhow, the actual taxonomic identity of the Miocene the genera Melissiodon (its last occurrence is at the end of representatives of the genus (including the large-sized MN4) and Megacricetodon, which first appears in Central forms mentioned above) is in no way a trivial problem. Europe during the MN4. In addition, the small-mammal Regarding an excessive phylogenetic diversity of extant assemblage was correlated with other early Miocene rhinolophids in tropic and subtropic habitats, we may Czech localities such as Dolnice 3 and Ořechov, both expect a large number of species also in the Neogene MN4b in age, based on the small mammal assemblage European fossil record, similarly as multiple cases of (Fejfar 1990). Dolnice 3 and Ořechov have yielded parallel morphoclines, and quite minute phenotypic dif- scarce remains of Megacricetodon; this genus is absent in ferences among individual species. The detailed analyses Dolnice 1 and 2. The mammalian fossiliferous sequence of large population samples are required to resolve these of Ořechov is located below the Oncophora (Rzehakia) questions. beds, which limits the boundary between Ottnangian and Karpatian stages (Cicha et al. 1972). It has close affinities to the assemblage of Dolnice 3, especially for the presence Discussion of Megacricetodon. The similar age for both Czech sites was also supported by Ziegler & Fahlbusch (1986), when Biostratigraphic considerations comparing the MN4b localities from Germany (Rembach, Fortshart and Rauscheröd) to both Ořechov and Dolnice The early Miocene continental fossil record of the Czech localities. Moreover, these authors also confirmed that the Republic is quite well-documented. To date, there are upper limit of the Oncophora beds marked the boundary two well-known MN3 localities: Tuchořice and Ahníkov/ between the Ottnangian and Karpatian stages. The other 115 Bulletin of Geosciences • Vol. 96, 1, 2021 two Dolnice assemblages (1 and 2), which yielded remains Nevertheless, the German locality Achen 17 did not of Democricetodon, Ligerimys antiquus and L. florancei, support the age of Ořechov proposed by Abdul Aziz et al. lack remains of the rodent genus Megacricetodon (Fejfar (2010), which is on the strontium content of otoliths. In 1990). Therefore, both assemblages were referred to fact, Achen 17 shows a similar assemblage to the faunas a somewhat older age (MN4a: see van der Meulen et al. registered from Dolnice 1 and 2, where Democricetodon 2011). and Pseudotheridomys are common, and Megacricetodon Kälin & Kempf (2009) define the Democricetodon is missing. Considering that the site was dated around franconicus-Megacricetodon collongensis interval zone, 17.8 ± 0.3 Ma, it would make the 18 Ma date for Ořechov with Glovelier as the reference locality, in the Northern unsustainable, considering the faunal differences between Alpine Foreland Basin of Switzerland. This local mammal both assemblages. zone is characterized by the first occurrence of the modern Recently, Reichenbacher et al. (2013) presented a new cricetid Democricetodon, and the genus Ligerimys, being magnetostratigraphic and biostratigraphic correlation the latter represented by L. antiquus and L. florancei. It is of the Swiss and South German Molasse Basins. This noteworthy that the top of this mammal zone is marked work proposed a new Ottnangian–Karpatian boundary, by the first appearance datum (FAD) of Megacricetodon. pushing the limit of these two stages half a million Based on the absence of Megacricetodon, Dolnice 1 years younger. Thus, the Kirchberg Formation and the and 2 were dated older than Ořechov and Dolnice 3, in age of the Oncophora beds were reconsidered, being agreement with the work of van der Meulen et al. (2011). the new correlation aged between 17.2 and 17.3 Ma. Furthermore, the above-mentioned biozone was placed Con­s equently, the age of Ořechov, located below the between 17.55 Ma and 17.15 Ma. This age agrees with the Oncophora beds, was dated around 17.3 Ma. Since Dol­ one proposed for Dolnice 3 and Ořechov, which remain in nice 3 shows an assemblage very similar to Ořechov, an the late MN4. alike age for it is expected. Moreover, the latest studies However, works focusing on the stratigraphy of Swiss focusing on Megacricetodon from Central European and German Molasse Basins have yielded different results localities (Fejfar 1990, Abdul Aziz et al. 2010, Oliver & regarding the correlation with the age and MN zonation Peláez-Campomanes 2016) agree with the latest proposal of the Czech localities. Research conducted on the by Reichenbacher et al. (2013), placing Dolnice 3 and stratigraphy of the Upper Freshwater Molasse in western Ořechov on the early MN4, and consider these remains Bavaria propose an older age for the Czech locality of to be the first representatives of the M. bavaricus group Ořechov (Abdul Aziz et al. 2010). The authors pointed (Oliver & Peláez-Campomanes 2016). out that the Oncophora beds, which overlain the Ořechov section, are correlated to the Kirchberg Formation (Brackish Water Molasse), and therefore, they assume The age of MWQ1/2001 Turtle Joint a middle–late Ottnangian age (around 18 Ma: MN4a) for Ořechov. Considering this information, Ořechov marks In general terms, the composition of faunal assemblage the FAD of Megacricetodon in Central Europe; the size of from MWQ1/2001 is similar to the other early Miocene Megacricetodon from Ořechov is smaller than that from Czech localities. Our results presented here confirm Langenau, a locality recording the earliest appearance of an early Miocene age (MN4) for this locality based on this genus in the North Alpine Foreland Basin (NAFB), the composition of the small mammal assemblage (see correlative to the base of the Kirchberg Formation, and Systematic palaeontology above), as tentatively stated therefore slightly younger than Ořechov. An age of about by previous works. The final allocation of MWQ1/2001 18 Ma for both Ořechov and Dolnice 3 would make the within the MN4 zone (early or late) remains to be specified. localities of Dolnice 1 and 2 even older. On the other It is remarkable that the scarce Megacricetodon material hand, other studies did not support the age for Ořechov recovered from MWQ1/2001 is significantly different from given on the latter study (see Kälin & Kempf 2009, van those from Ořechov and Dolnice localities (see remarks of der Meulen et al. 2011). According to these studies, the Megacricetodon sp. above), in both size and morphology. presence of Ligerimys antiquus, Melissiodon and the first Despite this, the alpha taxonomy of the remains from record of Democricetodon define Spanish Zone A, which MWQ1/2001 together with the evolutionary history and ends around 17 Ma. Furthermore, this faunal association different dispersals of Megacricetodon in Central Europe can be correlated with other localities from Central is still uncertain. Although there is a temporal trend in size Europe such as Glovelier (Switzerland), Petersbuch 2 increase along the different lineages of Megacricetodon and Erkersthofen 1 and 2 (Germany), Dolnice 1 and 2 (Abdul Aziz et al. 2010, fig. 3), further research on the (Czech Republic), and Oberdorf 3 and 4 (Austria). Given early evolution of the genus is required. The smaller size that Dolnice 3 and Ořechov reported Megacricetodon of the remains from MWQ1/2001 compared to those from remains, they should be considered the youngest sites. Dolnice 3 and Ořechov (and even smaller than M. aff. 116 Isaac Bonilla-Salomón et al. • Early Miocene small mammals collongensis from Langenau 1) could agree with an older conditions (Minwer-Barakat et al. 2009, Furió et al. 2011). age of the fossil assemblage. Besides, the Megacricetodon At any rate, the presence of the erinaceids in MWQ1/2001 from MWQ1/2001 shows clear morphological differences agrees with semi-open karstic vegetation surrounded with the species from Aliveri but resembles it in size. This by water bodies. The large mammal association is still could represent the earliest dispersal of this genus into under study by one of the authors of the present work Central Europe from Asia. The study of additional remains (I. Bonilla-Salomón) and mainly consists of cervids and of Megacricetodon, as well as the genus Ligerimys (of one suid. Although they require forest biotopes, it cannot great biostratigraphical value) in the rest of fissures will be be excluded that they could have lived close to a swampy very useful to elucidate the correct allocation of Mokrá- environment (Rössner 2004). Essentially, the mammal Quarry localities in the early Miocene. assemblages from MWQ1/2001 indicate warm weather conditions with patches of open forest and open steppe with the presence of marshy areas, which is typical of Palaeoecology karst landscapes. Palaeoecological reconstruction based on the study of As stated in the Systematic Palaeontology section, the small mammals reinforces the results obtained by the small mammal assemblage recovered from MWQ1/2001 study of other vertebrate groups i.e. amphibians and rep­ is relatively diverse. To date, eight different taxa have tiles. The diverse community of peri- or semi-aquatic been recovered: two lagomorphs; two chiropterans; one amphibians (Chelotriton, Triturus, Bufo and Pelophylax; hedgehog; and three rodents. Over 75% of that assem- Ivanov 2008) and reptiles (Ptychogaster, Natrix, Colu­ b­l age belongs to remains of Lagomorpha. The early brinae; Alba et al. 2018; Ivanov et al. 2018, 2020; Miocene prolagids from Central Europe can be regarded Luján et al. 2014, in press) in MWQ1/2001 indicates as forest-dwellers or as a peri-aquatic/riparian-dwellers, the presence of damp and marshy habitats with stagnant although their presence in more open-forest areas cannot or slow flowing waters in the locality. The absence of be excluded (cf. López-Martínez 2001). The genus crocodilians and most of freshwater turtle genera (i.e. Aliveria belongs to the flying squirrel group. Flying Trionyx, Rafetus, Clemmydopsis, Mauremys, Chely­ squirrels, which lived in forest environments, are in dropsis) at MWQ1/2001 suggest the presence of shallow general characterized by its nocturnal activity (de Bruijn and non-permanent water masses (Díaz Aráez et al. 1999). The record of Melissiodon is indicative of the 2017, Luján et al. 2019). Some water reservoirs were presence of wooden areas and somewhat humid conditions most probably surrounded by sandy shores with well- according to Jovells-Vaqué & Casanovas-Vilar (2018), drained substrates as documented by the occurrence of the although its feeding behaviour is still under discussion. extinct pelobatid frog Pelobates sanchizi (Venczel 2004, Mein & Freudenthal (1981) suggest a frugivorous diet Böhme & Vasilyan 2014). Species well-adapted to more for this genus, whereas Agustí & Antón (2002) advocate humid terrestrial environments have been also reported, for an arboreal behaviour and a diet based on leaves. such as Chelotriton sp., whose adults probably inhabited However, recent works (Wessels et al. 2018) suggested damp habitats with dense vegetation (Estes 1981, Ivanov that the specialization of the morphology of the chewing 2008). Similarly, Pseudopus laurillardi inhabited humid apparatus of the Melissiodontinae is rather an adaptation woodland environments. However, diverse assemblage to feeding on small invertebrates. As for Megacricetodon, of terrestrial heliophilic squamates [e.g. Ophisaurus, given its opportunistic condition, habitat preferences are Varanus mokrensis, Bavarioboa cf. hermi, Vipera still uncertain. In any case, as a ground dweller genus it (‘Oriental vipers’ group)] and a testudinid (Testudo cf. could indicate open-forest conditions (Kälin 1999). kalksburgensis; Luján et al. 2016, 2017) show on the open Regarding the rhinolophid bats, they are strict cave- woodland, bushes and grassland in close vicinity of dwellers foraging invertebrates in foliage (Storch 1999). water reservoirs. The presence of thermophilic taxa in This is consistent with a forest or semi-opened karstic MWQ1/2001 including Python, Varanus, Bavarioboa, vegetation. The presence of these bats indicates a spacious as well as elapids and ‘Oriental vipers’ indicate rather cave in time of deposition, close to these forest and semi- warm climatic conditions during the Miocene Climatic open karstic vegetation. Ziegler (1999) pointed out that the Optimum with mean annual temperatures reaching at least close relatives of Galerix live in forested environments, 18‒19 °C (Ivanov et al. 2018, 2020). close to water bodies, where they feed on invertebrates, frogs and some plant matter. However, other works point out that Miocene Galericini, such as Galerix and Para­ Conclusions sorex, were adapted to a high diversity of habitats (García- Alix et al. 2008; Furió et al. 2011, 2018); these taxa were The small mammal remains from the early Miocene event present in Miocene assemblages typical of arid loca­lity of MWQ1/2001 (Mokrá-Western Quarry, South 117 Bulletin of Geosciences • Vol. 96, 1, 2021 Moravia, Czech Republic) are described and attributed to Angelone, C., Jovells-Vaqué, S., Luján, À.H., Almécija, S. eight taxa: Prolagus schnaitheimensis, P. cf. vasconinensis, & Moyà-S olà , S. 2018. Can Pallars i Llobateres: A new Melissiodon dominans, Megacricetodon sp., Aliveria hominoid-bearing locality from the late Miocene of the aff. luteyni, Galerix sp., Rhinolophus cf. cluzeli, R. cf. Vallès-Penedès Basin (NE Iberian Peninsula). Journal of grivensis. The faunal assemblage corresponds to the Human Evolution 121, 193–203. MN4, based on one of the last occurrences of Melissiodon DOI 10.1016/j.jhevol.2018.04.008 Á lvarez -S ierra , M.A., D aams , R., L acomba , J.I., L opez - together with the presence of the cricetid Megacricetodon. Martinez, N., van der Meulen, A.J., Sese, C. & de Visser, J. MWQ1/2001 shows a micromammal assemblage similar 1991. Palaeontology and biostratigraphy (micromammals) of to those of Dolnice 3 and Ořechov. Although scarce, the continental Oligocene-Miocene deposits of the North-Cen­ the remains of Megacricetodon sp. from MWQ1/2001 tral Ebro Basin (Huesca, Spain). Scripta Geologica 94, 1–77. are essential to the understanding the appearance and A ngelone , C. & S esé , C. 2009. New characters for species different migration waves of this genus in Europe. More discri­m ination within the genus Prolagus (Ochotonidae, remains from the unpublished Mokrá-Quarry localities, Lagomorpha, Mammalia). 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