This is the accepted version of the journal article: Jovells Vaqué, Sílvia; Bonilla Salomón, Isaac; Mažgút, O.; [et al.]. «Cricetid, eomyid and murid fauna from the Middle Miocene site (MN6) of Devínska Nová Ves - Bonanza (Slovakia)». Geobios, Vol. 79 (Aug. 2023), p. 1-15. DOI 10.1016/j.geobios.2023.05.008 This version is available at https://ddd.uab.cat/record/282012 under the terms of the license 1 Cricetid, Eomyid, and Murid fauna from the Middle Miocene site (MN6) of 2 Devínska Nová Ves – Bonanza (Slovakia) 3 4 S. Jovells-Vaqué a,b,*, I. Bonilla-Salomón a, O. Mažgút a, S. Čermák c, À.H. Luján b,d, P. 5 Joniak a, M. Sabol a 6 7 a Department of Geology and Palaeontology, Faculty of Natural Sciences, Comenius 8 University, Ilkovičova 6, SK - 84215 Bratislava, Slovak Republic 9 10 b Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, 11 Edifici ICTA-ICP, c / de les Columnes, s/n, Campus de la UAB, 08193 Cerdanyola del 12 Vallès, Barcelona 13 14 c Institute of Geology of the Czech Academy of Sciences, Rozvojová 269, 16500 Prague 6, 15 Czech Republic 16 17 d Department of Geological Sciences, Faculty of Sciences, Masaryk University, Kotlářská 18 267/2, 611 37 Brno, Czech Republic 19 20 * Corresponding author. E-mail address: silvia.jovells@icp.cat (S. Jovells-Vaqué). 21 22 1 23 Abstract 24 Rodents play an important role in all current and extinct ecosystems. In this study we present 25 the fossil record of cricetids and eomyids from Devínská Nová Ves–Bonanza (Vienna Basin, 26 Slovakia). A detailed morphometric analysis revealed the presence of Eumyarion cf. latior, 27 Democricetodon vindobonensis, Karydomys sp., Anomalomys sp., Neocometes cf. brunonis, 28 Keramidomys carpathicus, and Eomyops sp. This species association with other taxa available 29 from the locality indicates an early MN6, probably slightly younger in age than the 30 neighbouring locality Devínská Nová Ves–Štokeravská vápenka = Neudorf an dem March – 31 die Spaltenfüllung or Zapfe’s Fissures. Paleoenvironmental specificities of the locality dated 32 after the peak of the Mid-Miocene Climatic Optimum are discussed. The faunal assemblage, 33 together with geological setting, suggests a mixed insular or peninsular area with subtropical 34 forests close to freshwater bodies in the vicinity of a saltwater sea. 35 36 Keywords: 37 Rodents 38 Cricetids 39 Eomyids 40 Central Europe 41 MN6 42 Late Badenian 43 44 2 45 1. Introduction 46 Small mammal fauna, especially rodent fauna, plays a very significant role in current 47 and past ecosystems. They represent nowadays 40% of the mammal biodiversity worldwide 48 (Pardiñas et al., 2017; D’Elía et al., 2019). Rodents are commonly characterized as small and 49 diversified mammals, specialized in gnawing and with high reproductive rates. Their radiation 50 includes different types of species from subterranean species to arboreal and cursorial forms 51 (D’Elía et al., 2019). These characteristics make the rodent group one of the most adaptative 52 groups across the globe. Rodent fossil fauna are common components of the mammalian 53 fossil record in the Miocene terrestrial deposits in Europe. They also represent one of the most 54 biochronologically significant groups of fossil mammals due to their rapid evolutionary rates 55 and their wide geographical distributions. Therefore, the first and last appearances of 56 determinate rodent taxa, such as cricetids and eomyids, mark ‘the boundaries’ between 57 different biochronological units used in terrestrial record, including the Mammal Neogene 58 (MN) zone system in Europe (Mein, 1975, 1990, 1999; Fahlbusch, 1976; Bruijn et al., 1992; 59 Van Dam et al., 2001, 2006; Hilgen et al., 2012). Moreover, the cricetid and eomyid fauna 60 along with other groups of rodents, such as glirids and sciurids, are also important for 61 paleoecological and paleoenvironmental studies. 62 In Slovakia, the fossil record of rodents currently found in Miocene deposits is not as 63 rich as in other parts of Europe, as they have been found only in eight out of at least 27 64 Miocene mammal sites (Sabol et al., 2021). The record for the fossil rodents in the Slovak 65 territory begins during the Astaracian (Middle Miocene; MN6) since throughout the Early 66 Miocene most of the territory was submerged by the vast inland Central Paratethys sea 67 (Kováč et al., 2017, 2018). 68 This study focuses its attention on the cricetid and eomyid fauna from the Devínska 69 Nová Ves–Bonanza site (DNV–Bonanza; Vienna Basin) correlated to the early Astaracian 3 70 (Middle Miocene, MN6; see Sabol and Holec, 2002; Sabol, 2005a,b; Sabol et al., 2021). 71 DNV–Bonanza site is located in an area that had a peninsular to insular environment during 72 the Badenian, based on the characteristics of the faunal and geological record (Sabol et al., 73 2021). The sedimentary record of DNV–Bonanza is related to paleobiogeographic changes 74 that took place during the Middle Miocene in the Central Paratethys area (Kováč et al., 2017, 75 2018). 76 DNV–Bonanza site was discovered by amateur paleontologist Š. Meszároš in 1982. It 77 is located in the former Štokeravská vápenka (= Stockerau limestone quarry), situated in the 78 northern slope of Devínska Kobyla Hill near the Village of Devínska Nová Ves (Holec et al., 79 1987; Fejfar and Sabol, 2005; Sabol and Kováč, 2006; Mažgút, 2010). This site is a fissure 80 filling with high probability of Badenian sediment infill. It contains a rich marine and 81 terrestrial fossil fauna. DNV–Bonanza is known for its mammal fossil remains, especially for 82 the primitive Miocene seals (Holec et al., 1987; Koretsky and Holec, 2002; Koretsky and 83 Rahmat, 2015). These findings promoted new excavations in the area lead by a team from the 84 Comenius University (Bratislava) and the Smithsonian Institution (Washington) in 1997– 85 1998 (Holec et al., 1987), as well as by subsequent revision research teams in 2001–2022 86 (Sabol et al. 2021). 87 88 2. Geological setting 89 The DNV–Bonanza site is a broad karst fissure located at the eastern margin of the 90 abandoned limestone quarry (the geographic coordinates of the site are 48°12’13” N and 91 17°00’11” E) but in the opposite quarry wall, as the most famous site of DNV – Štokeravská 92 vápenka = Neudorf an dem March – die Spaltenfüllung or Zapfe’s Fissures (Holec et al., 93 1987). The site itself is situated in the protective wall of the Lower Jurassic dark limestone, 94 oriented towards the railway line from Bratislava to Prague (Fig. 1(A–B)). Marine sands, 4 95 sandstones, and large limestone boulders fill the fissure (Fig. 1(C)). Single layers of this 96 sedimentary filling document change of coastal line position associated with several (possibly 97 orbital) cycles and contain fossils of terrestrial (incl. freshwater), semi-aquatic, and marine 98 vertebrates (Sabol and Kováč, 2006; Sabol et al., 2021). A detailed description of DNV– 99 Bonanza site, including the most important sites for the interregional correlations in Central 100 Europe, has been presented by Holec et al. (1987), Sabol and Kováč (2006), and Lehotský et 101 al. (2009). The DNV–Bonanza site, is a type locality of four species (Bufo priscus Špinar, 102 Klembara and Meszáros, 1993, Gerhardstorchia meszaroshi Sabol, 2005, Devinophoca 103 claytoni Koretsky and Holec, 2002, and D. emryi Koretsky and Rahmat, 2015) and it forms a 104 “transition” between DNV–Štokeravská vápenka and DNV–Sandberg sites from the 105 biostratigraphic point of view (Sabol and Kováč, 2006; Sabol et al., 2021). Moreover, these 106 sites show a direct relationship between the terrestrial and the marine record, based on the 107 collected data published in Sabol et al. (2021). The DNV–Bonanza site is dated 108 approximately around 13.8 Ma, while the oldest site of this area, i.e., DNV–Štokeravská 109 vápenka, has an approximate time span from 14.7 to 13.8 Ma (early Badenian, Middle 110 Miocene: Sabol et al., 2021). The remaining and youngest site of the area is DNV–Sandberg 111 and dates back from 13.8 Ma to 12.8 Ma (Sabol et al., 2021). The correlation of these sites is 112 important because it shows a complete record of terrestrial and marine fauna ranging from 113 14.7 to 12.8 Ma from the Central Paratethys area during a time of great changes, as was the 114 Mid Miocene Climatic Optimum (MMCO; Steinthorsdottir et al., 2021). 115 116 3. Material and methods 117 The material from DNV–Bonanza is housed at the Faculty of Natural Sciences of the 118 Comenius University in Bratislava (numbered as KGP-MSxx, KGP-xx.x and DB92xxxx), and 119 at the Slovak Natural Museum – Natural History Museum (labelled as Zxxxx) in the same 5 120 city. The micromammal assemblage described in this work comes from the thorough 121 excavations that took place during the years 1997–1998, 2001, and 2002. Part of these 122 materials was already figured in Sabol, Joniak and Holec (2004). The material numbered as 123 DB92xxxx was recovered in 1991-1993 field campaigns by PJ. Dental terminology for the 124 cricetid fauna follows Oliver and Peláez-Campomanes (2013), whereas the measurement 125 method is after Daams and Freudenthal (1988: 42, fig. 1). The dental terminology for eomyid 126 species follows Fahlbusch (1970) and Álvarez-Sierra (1987) for the measurements. The dental 127 terminology for Neocometes follows Fahlbusch (1966). All measurements are given in 128 millimetres and were taken with an optical micrometer to the nearest 0.01 mm. Estimated 129 measurements, because of minor damage or distortion, are between brackets. Summary 130 statistics and scatterplots were performed using the R software (R Core Team 2017). The 131 biochronologic/biostratigraphic terminology and boundaries follow Steininger (1999) and 132 Hilgen et al. (2012). Although the material under study stems from different layers of the 133 same fissure deposited in the time horizon at approximately 13.8 Ma (Sabol et al., 2021), it is 134 evaluated as a whole. 135 136 4. Systematic paleontology 137 Order Rodentia Bowdich, 1821 138 Family Cricetidae Fischer [von Waldheim], 1817 139 Genus Eumyarion Thaler, 1966 140 Type species: Cricetodon medius Lartet, 1851 (= Cricetodon helveticus Schaub, 1925). 141 142 Eumyarion cf. latior Schaub et Zapfe, 1953 143 Material and measurements: Partial skull with fragment of upper M1 (KGP-1: 144 unmeasurable); left M1 (KGP-6a.4: L= 2.05, W= 1.48); right m1 (KGP-9.2: L= 2.14, W= 6 145 1.34); m1 fragment (KGP-MS60: unmeasurable); right m2 (KGP-6a.5: L= 1.65, W= 1.45); 146 left m2 (KGP-6a.6: unmeasurable); fragment; left m3 (KGP-6a.9: L= [1.53], W= [1.29]), right 147 m3 (KGP-9.3: L= 1.5, W= 1.32). 148 Description: M1 (Fig. 2(A)). The roots are not preserved in specimen KGP-6a.4. The 149 anterocone is simple and well developed and has a labial position compared to the rest of the 150 tooth. The labial part of the anteroloph presents a bifurcated spur that occupies most of the 151 anterosinus but does not connect with the spur of the paracone. The lingual part of the 152 anterocone connects with the protocone by the anterolophule. The anterolophule presents two 153 little spurs strongly directed to the anterosinus, but they do not show any connection. The 154 paracone is well-developed and presents three different spurs: the first one is located in the 155 lower anterior part of the protolophule; the second one is a small spur directed to the labial 156 anterocone spur, but it does not show any connection; and the third one, the ectoloph, is a 157 ridge that follows the margin of the tooth and is directed to the mesoloph but without reaching 158 it. The mesoloph is long but does not reach the labial edge. The sinus is deep and anteriorly 159 directed. There is no presence of an entomesoloph in these specimens. The metalophule is 160 simple and connects anteriorly to the hypocone. The posteroloph is long and well-developed, 161 reaching the metacone in its posterior part: this results in a narrow and deep posterosinus. 162 m1 (Fig. 2(B)). There are two roots in these specimens: the anterior one is smaller and 163 located below the anteroconid, while the posterior one is more robust. The anteroconid is 164 simple and round-shaped. The anterosinusid is open and narrower than the protosinusid, 165 which is closed by a low cingulid that emerges from the anteroconid and would correspond to 166 the labial anterolophid. There is also a small protostylid at the end of the labial anterolophid. 167 The metalophid is double; the anterior arm connects just before the protoconid and the 168 posterior arm behind it. There is a short anteriorly directed mesolophid in the specimen KGP- 169 9.2 (Fig. 2(B)), the other specimen (KGP-MS60) is broken, being only the anterior part of the 7 170 tooth present, but a short mesolophid is preserved. The sinusid is wide and open and there is a 171 faint presence of an ectomesolophid. The hypolophulid is simple and connects anteriorly the 172 entoconid with the hypoconid. The posterolophid is well-developed and encloses a wide 173 posterosinusid. 174 m2 (Fig. 2(C)). There are two roots in these teeth, being the anterior smaller and less 175 robust. The lingual anterolophid is reduced and barely present. It is directly connected with 176 the metaconid, resulting in a very narrow and reduced anterosinusid as is shown in specimen 177 KGP-6a.5 (Fig. (2C)). The labial anterolophid is more developed than the lingual one and 178 encloses a narrow protosinusid. Anterior to the mesoconid, there are two ridges: the former is 179 posteriorly directed and corresponds to the protoconid hind arm, whereas the latter is smaller 180 and shorter, corresponds to the mesolophid, and connects to the lingual end of the protoconid 181 hind arm. The metalophulid is simple and anterior. The posterolophid is well-developed and 182 connects with the posterior part of the entoconid, resulting in a closed posterosinusid. The 183 sinusid is posteriorly directed and closed by a low cingulid. There is a faint presence of the 184 extomesolophid. 185 m3 (Fig. 2(D)). There are two specimens available in the DNV–Bonanza site material. 186 These teeth present two roots, one located in the anterior position, just below the 187 anterolophids, and the other in the posterior part of teeth. The anterolophids are well- 188 developed, being the lingual anterolophid more reduced than the labial anterolophid, resulting 189 in a barely present anterosinusid and a small protosinusid. The mesolophid is medium-sized, 190 but it does not reach the lingual margin of the tooth in both preserved specimens (KGP-6a.9 191 and KGP-9.3; Fig. 2(D)). The hypolophulid is single and connects the entoconid with the 192 hypoconid. The posterolophid is well-developed enclosing a wide posterosinusid, whereas the 193 sinusid is narrow and posteriorly oriented. 8 194 Remarks: This genus has a combination of primitive dental characteristics that make them 195 easily identifiable from other Miocene cricetids (see Bruijn, 2009). The specimens recovered 196 in the DNV–Bonanza site, show several diagnostic features that allow to refer them to the 197 Eumyarion genus, such as the labial position of the anterocone and the bifurcated anteroloph 198 in the M1. 199 Apart from the isolated check teeth, a partial skull with the rostrum, part of the 200 palatine and the maxilla with an upper left M1 fragment was recovered as well (KGP-1; Fig. 201 3(A–B)). The presence of this tooth fragment allows the adscription of this skull to the genus 202 Eumyarion and most likely to Eumyarion cf. latior species due to the morphology of the 203 anterocone, anterolophule, and part of the metacone. 204 The genus Eumyarion is characterized by an archaic tooth morphology showing 205 similarities with some Oligocene cricetids such as Eucricetodon (Kälin, 1999). Several 206 species of the genus have been described during the Early and Middle Miocene in Europe, 207 which are the following: Eumyarion weinfurteri (Schaub and Zapfe, 1953), E. bifidus 208 (Fahlbusch, 1964), and E. latior (Schaub and Zapfe, 1953). The validity of both taxa E. latior 209 and E. weinfurteri has already been discussed by several authors (e.g., Engesser, 1972; Bruijn 210 and Saraç, 1991; Bruijn, 2009). In fact, the different size ranges for Eumyarion species is 211 depicted in Figure 4 as well, being E. weinfurteri the smaller species within the genus 212 Eumyarion. That figure further shows that there is great homogeneity among the Eumyarion 213 populations from DNV sites. Both Eumyarion weinfurteri and E. latior were described by 214 Schaub and Zapfe (1953) from the same fissures, being size the only remarkable difference 215 between the two species. From then on, smaller sized specimens of the genus Eumyarion from 216 the Early Miocene European fossil record were assigned to E. weinfurteri based solely on this 217 character (Fejfar, 1974; Bolliger, 1992; Daxner-Höck, 1998). Even though it is not within the 218 scope of this paper to elucidate the controversy regarding E. weinfurteri and E. latior, some 9 219 authors of this work are working on a revision of the populations of Eumyarion from DNV 220 sites, which may shed light on the subject. For the time being, the material is here referred to 221 as Eumyarion cf. latior. 222 223 Genus Democricetodon Fahlbusch, 1964 224 Type species: Democricetodon crassus Freudenthal in Freudenthal and Fahlbusch, 1969 225 (=Cricetodon minor Lartet, 1851 sensu Fahlbusch 1964). 226 227 Democricetodon vindobonensis (Schaub et Zapfe, 1953) 228 Holotype and type locality: Originally described as Cricetodon brevis vindobonensis nov. 229 ssp. (Schaub and Zapfe, 1953); holotype: mandible fragment with m1 and m2 sin.; type 230 locality: DNV–Štokeravská vápenka (originally, Horizont B der Neudorfer Spalte, Helvétien). 231 Material and measurements: partial skull; right maxillary fragment with M1-M3 KGP-17 232 (M1, L: 1.62, W: 0.87; M2, L: 1.32, W: 1.10; M3, L: 1.02, W: 0.85); M1: (left KGP-6a.7 (L: 233 1.92, W: 1.32), right KGP-6a.8 (L: 1.87, W: 1.30), right KGP-MS29 (L: 1.65, W: 1.10), right 234 KGP-MS30 (L: -, W: 1.10); M2: (left KGP-6a.12 (L: 1.37, W: 1.25), left KGP-MS31 (L: 1.4, 235 W: 1.22); M3: (right KGP-MS34 (L: 0.95, W: 1.23)); m1: (left KGP-6a.10 (L: 1.52, W: 236 1.12)), right KGP-MS35 (L: 1.52, W: 1.05), right KGP- MS36 (L: 1.47, W: 1.00), right KGP- 237 MS37 (unmeasurable), left Z14597 (L: 1.27, W: 0.90); m2: (left KGP-6a.11 (L: 1.37, W: 238 1.12), left KGP-6a.13 (L: 1.32, W: 1.15), left KGP-6a.14 (L: 1.23, W: 0.95), right KGP-MS38 239 (L: 1.37, W: 1.00), left KGP-MS39 (L: 1.37, W: 1.10), left KGP-MS40 (L: 1.37, W: 1.08), 240 left Z14597 (L: 1.28, W: 1.10), left Z14598 (L: 1.43, W: 1.12); m3: (left KGP-MS61 (L: 1.08, 241 W: 0.97), left KGP-MS62 (L: 1.20, W: 0.97), left Z14597 (L: 1.22, W: 0.86)). 242 Description: M1 (Fig. 2(E–G)). There are three roots available in the M1 specimens. The 243 larger ones are located under the labial and the lingual part of the tooth respectively, whereas 10 244 the third one is located under the anterocone. The anterocone is simple and well-developed 245 and has a labial position compared to the rest of the tooth. As is seen in unworn specimens 246 (e.g., Fig. 2(F–G)), the anterolophule shows a labial long spur that reaches the margin of the 247 teeth dividing the anterosinus into two small valleys. The protolophule is double in KGP-6a.7 248 (Fig. 2(E)), whereas in the other specimens studied it is simple, being the anterior arm 249 connected with the basis of the anterolophule labial spur (KGP-6a.14; Fig. 2(F)). The 250 protolophule hind arm connects with the entoloph posteriorly to the protocone. The mesoloph 251 is long but ends before connecting to a small mesostyle. The metalophule is simple and 252 connects directly to the posteroloph, except for the specimen KGP-6a.14 (Fig. 2(F)). The 253 posterosinus is narrow and deep, closed by the posteroloph. The sinus is narrow and deep and 254 is closed by a small cingulum. 255 M2 (Fig. 2(H–I)). There are two roots in the preserved two samples: one of them root 256 is placed labially and the other one on the lingual part of the tooth. The labial anterolophule is 257 well-developed enclosing a narrow anterosinus. Similarly, the lingual anteroloph is well 258 developed and encases a narrow and small protosinus. The protolophule is double following 259 the same pattern described for the M1 specimens (Fig. 2(E–G)). The mesoloph is long and 260 reaches the margin of the tooth forming a small mesostyle on the mesosinus. The sinus is 261 narrow and is closed by a low cingulum. The metalophule is simple and connects with the 262 entoloph posteriorly to the mesoloph as is seen in specimen KGP-6a.12 (Fig. 2(I)), the 263 metalophule is simple and is connected with the mesoloph through the entoloph. The 264 posteroloph is well-developed and encloses a deep and narrow posterosinus. 265 M3 (Fig. 2(J)). Two M3 were recovered from the DNV–Bonanza site, being the 266 specimen (KGP-17) the only attached to a maxilla fragment that contains the complete tooth 267 row (M1 to M3). The other specimen (i.e., KGP-MS34; Fig. 2(J)) is a single right M3. The 268 root of KGP-MS34 is not preserved, whereas in KGP-17 it is not visible as it is attached to the 11 269 maxilla in its natural position. The general morphology of both M3 is rounded. The lingual 270 and labial anterolophs are well-developed, enclosing the anterosinus and the protosinus 271 respectively. The sinus is closed by a low cingulum. The paracone is connected to the 272 anterolophule by a single protoloph arm. The metaloph is simple. The posteroloph is well- 273 developed and reaches the posterior part of the metacone, enclosing the posterosinus. 274 Moreover, a partial skull embedded into sandstone matrix was also recovered from 275 DNV–Bonanza site. Only part of the skull is visible, including the right tooth row (KGP-17; 276 Fig. 3(C–D)). The latter can refer to Democricetodon vindobonensis due to the occlusal 277 morphology of the M1-M3 cheek teeth. Apart from the tooth row, the right incisor is also 278 present. Regarding the skull, the rostrum is preserved along with the right zygomatic arch, the 279 frontal bone, and the parietal bones. This skull is flattened dorsally (Fig. 3(C–D)). 280 m1 (Fig. 2(K–M)). The m1 specimens have two roots; the former is located on the 281 anterior part of the tooth and the latter is on the posterior part. Both roots have similar 282 dimensions. The anteroconid is simple and small compared to the anterocone of M1s. The 283 lingual anterolophid is short and defines a small round-shaped anterosinusid as shown in 284 specimen KGP-6a.10 (Fig. 2(K)). On the contrary, the labial anterolophid is longer and more 285 developed than the lingual one, which results in the enclosing of the protosinusid. The 286 metaconid presents two anterior spurs: the lingual one merges with the lingual anterolophid, 287 except in specimen KGP-MS36 (Fig. (2M)); and the labial one would correspond to the 288 protolophulid, which is simple and connects to the anterolophulid, leaving the protoconid 289 isolated from the metaconid and anteroconid. The mesolophid is long and reaches the lingual 290 margin of the tooth. The metalophulid is simple and anterior. The posterolophid is long and 291 well-developed, and merges with the entoconid resulting in a closed posterosinusid. The 292 sinusid is also closed by a low cingulid. 12 293 m2 (Fig. 2(N–S)). These specimens have two roots. The lingual anterolophid is very 294 short and almost absent in some cases (e.g., KGP-6a.13; Fig. (2O)); in fact, this results in very 295 reduced and in some cases absent anterosinusid. On the contrary, the labial anterolophid is 296 well-developed enclosing the protosinusid. The mesolophid is long and reaches the lingual 297 margin of the tooth. Both the metalophulid and hypolophulid are simple and anterior. As in 298 seen in the m1, the posterolophid is well-developed and enclosing the posterosinusid. The 299 sinusid is closed by a low cingulid as well. 300 m3 (Fig. 2(T)). There are three m3 available from DNV–Bonanza, all of them with 301 two roots. The lingual and labial anterolophids are well-developed, being the lingual one 302 smaller than the labial one: this results in a small anterosinusid and a longer, but narrow 303 protosinusid. There is no mesolophid in these specimens (Fig. 2(T)). The posterolophid is 304 well-developed and encloses a wide and deep posterosinusid. The sinusid is wide and 305 enclosed by a low cingulid. 306 Remarks: The specimens recovered from DNV–Bonanza site were originally named 307 Cricetodon brevis vindobonensis by Schaub and Zapfe (1953). However, they are here 308 referred to Democricetodon vindobonensis in having a long mesoloph/id that extends to the 309 margin of the teeth in both upper and lower molars forming a mesostyle/id, as well as the 310 presence of a long labial spur of the anterolophule which divides the anterosinus into two 311 smaller valleys (also called as “anteromesoloph” in Sabol, 2005a). Democricetodon 312 vindobonensis mainly differs from D. franconicus by the presence of a long mesoloph 313 reaching the margin of the teeth, and longer anteromesoloph. 314 As is depicted in Figure 5, the material recovered from DNV–Bonanza, despite scarce, 315 fits well within the size range of Democricetodon vindobonensis, especially regarding the 316 upper and lower M1/m1, which can be compared with the already published data from DNV– 317 Štokeravská vápenka (see Fejfar, 1974). Moreover, we included more data from other 13 318 Democricetodon species such D. franconicus from Dolnice 1-3 and Ořechov (Fejfar, 1974), 319 D. gracilis and D. mutilus from Sandelzhausen (Wessels and Reumer, 2009) and other Central 320 European localities to compare our specimens with these other assemblages. We concluded 321 that D. vindobonensis material from DNV–Bonanza site coincides in size within the range for 322 the D. vindobonensis from DNV–Štokeravská vápenka, as well it is quite similar in size with 323 D. franconicus from the Czech sites of Dolnice 1-3 and Ořechov (Fig. (5)), despite showing 324 clear morphological differences (see above). 325 326 Genus Karydomys Theocharopoulos, 2000 327 Type species: Karydomys symeonidisi Theocharopoulos, 2000. Karydia 1, Early Miocene, 328 Greece. 329 330 Karydomys sp. 331 Material and measurements: right M2 (KGP-MS63: L=2.50, W=2.05) 332 Description: M2 (Fig. 2(U)). There is only one single specimen represented by an isolated 333 upper M2 (KGP-MS63). The tooth is quite worn and makes it difficult to describe and 334 evaluate with confidence. It is square-shaped with a well-developed labial anteroloph that 335 follows the margin of the tooth and encloses the labial anterosinus. Similarly, the lingual 336 anteroloph is also well-developed and encloses the protosinus with a low ridge. The 337 protolophule is double. The anterior arm connects with the anterolophule right before the 338 protocone. The protolophule posterior arm connects directly to the mesocone and the lower 339 part of the mesoloph. The mesoloph is long and almost reaches the labial margin of the tooth, 340 but without the presence of mesostyle. Instead, the mesoloph connects directly with an 341 anterior metacone spur. The metaloph is simple and is joined directly with the posteroloph 342 just behind the hypocone. The posteroloph is well-developed and enclose the posterosinus. 14 343 The sinus is narrow and straight, enclosed by a low cingulum on the lingual margin of the 344 tooth, and with no endostyle present in the sinus. 345 Remarks: The solely recovered specimen shows high crown wear, which renders difficult its 346 taxonomical attribution. Despite this, it still retains several morphological characteristics that 347 allow for its tentative attribution to a genus level. Based on the dimensions of the M2, 348 together with some specific morphological characteristics (i.e., square-shaped morphology of 349 the teeth with the rounded edges of its crown, and long mesoloph without mesostyle at the 350 labial margin of the teeth and the absence of endostyle), we concluded that KGP-MS63 351 belongs to Karydomys sp. Moreover, the dimension of this tooth fits within the Karydomys 352 previously reported from DNV–Štokeravská vápenka published by Fejfar (1974) as 353 Lartetomys cf. zapfei. Later, Mörs and Kalthoff (2004) re-assigned the specimens found in 354 DNV–Štokeravská vápenka to Karydomys wigharti (Prieto, 2012). Anyhow, KGP-MS63 355 constitutes one of the largest cricetids recovered from DNV–Bonanza site. 356 357 Family Eomyidae Winge, 1887 358 Genus Keramidomys Hartenberger, 1966 359 Type species: originally described as Pseudotheridomys carpathicus nov. sp.; holotype: 360 maxilla dext. with P4 and M1; type locality: DNV–Štokeravská vápenka (originally, Horizont 361 D der Neudorfer Spalte, Helvétien: Schaub and Zapfe, 1953). 362 363 Keramidomys carpathicus Schaub and Zapfe, 1953 364 Material and measurements: two right M1 (KGP-MS69: L=0.85, W=0.90 and KGP-6a.18: 365 L=0.85, W=0.80). 366 Description: M1 (Fig. 2(V–W)). There are only two specimens available from DNV– 367 Bonanza site, and both correspond to the right M1. KGP-MS69 (Fig. 2(V)) is almost complete 15 368 while KGP-6a.18 (Fig. 2(W)) is quite damaged, with the labial part of the tooth completely 369 eroded. Both molars show a well-developed anteroloph that encloses the anterior syncline in 370 KGP-6a.18, yet remains open in KGP-MS69. The protoloph is thick in both molars reaching 371 the paracone. The mesoloph is long and well-developed reaching the lingual margin of the 372 teeth. There is no longitudinal crest connecting the mesoloph and the protoloph. Only a small 373 spur emerging from the mesoloph is discerned, thus resulting in an open labial syncline that 374 divides the molar into two parts. The base of the mesoloph is in contact with the hypocone. 375 The metaloph is also well-developed and connects the hypocone with the metacone. The 376 posteroloph is thick and follows the posterior margin of the tooth. In KGP-MS69, the 377 posterior syncline is left open, while in KGP-6a.18 is closed by the connection between the 378 posteroloph and the metacone probably due to the wear. 379 Remarks: The genus Keramidomys is one of the smallest eomyids known so far. This group 380 shows a very lophodont tooth pattern with the cups incorporated into the ridges and very 381 marked synclines between them. Keramidomys is also a biochronologically wide-ranged 382 genus being present from MN2/3 to MN14, thus resulting in several evolutionary lines 383 (Engesser, 1999; Kimura et al. 2019, Daxner-Höck et al. 2022). Originally erected by Schaub 384 and Zapfe (1953) as Pseudotheridomys carpathicus based on the material from DNV– 385 Štokeravská vápenka, Keramidomys carpathicus shares many features with K. thaleri, such as 386 the long mesoloph/id. However, in K. thaleri, the longitudinal crest is always present and in 387 many specimens this connection is complete, while in K. carpathicus this ridge is always 388 interrupted (Mein, 2009). The teeth recovered from DNV–Bonanza (Fig. (6)) coincide in size 389 with the K. carpathicus type material from DNV–Štokeravská vápenka published in Schaub 390 and Zapfe (1953) and Fejfar (1974). However, the scarcity of the assemblage recovered in 391 DNV–Bonanza is here referred to K. cf. carpathicus. 392 16 393 Genus Eomyops Engesser, 1979 394 Type species: Eomyops catalaunicus Hartenberger, 1966, Can Llobateres (Late Miocene), 395 Spain. 396 397 Eomyops sp. 398 Material and measurements: right P4 (KGP-6a.17: L=0.80, W=0.75) 399 Description: P4 (Fig. 2(X)). There is only one specimen available, but the poor state of 400 preservation renders it difficult to give an exhaustive description of the tooth. KGP-6a.17 401 (Fig. 2(X)) has a well-developed anteroloph that encloses the anterior syncline, probably due 402 to wear. The protoloph is well- developed and reaches the paracone. The mesoloph is short 403 and is posteriorly directed almost reaching the metacone. There is an indication of the 404 longitudinal ridge that connects the basis of the mesoloph with the protoloph. The metaloph is 405 thick and connects the hypocone with the metacone. The posterior part of the tooth is highly 406 worn so it is difficult to evaluate this area confidently. The labial syncline is wide and is 407 anteriorly directed. 408 Remarks: The genus Eomyops is one of the stratigraphically long-ranged eomyids known so 409 far in Europe. Its time span ranges from the Middle Miocene (MN5) to the Early Pleistocene 410 (MN17; Engesser, 1999), registered little changes regarding the cheek teeth morphology. One 411 of the main characteristics of Eomyops is its primitive bunodont cheek teeth structure 412 (Engesser, 1999). Particularly, KGP-6a.17 shows some of the important characteristics of the 413 genus Eomyops, such as a well-developed anteroloph and the presence of a second synclinal 414 extending to the central part of the teeth. There is a lot of discussion regarding the validity of 415 the genus Eomyops. Several authors, such as Engesser (1979) and Kimura et al. (2019), 416 consider Eomyops and Leptodontomys to be two separate genera. However, other authors, 417 (e.g., Kalthoff et al., 2022), consider these genera synonyms based mainly on the external 17 418 morphology and microscopic structure of the incisors. The genus Eomyops sp. was already 419 recovered in the nearby site DNV-Bonanza, DNV-Štokeravská vápenka (MN6; Schaub and 420 Zapfe, 1953; Kälin, 1997; Prieto, 2011). Since the single tooth recovered from this site is very 421 poorly preserved, a more precise attribution cannot be determined, therefore being referred to 422 as Eomyops sp. 423 424 Family Muridae Illiger, 1811 425 Subfamily Platacanthomyinae Alston, 1876 426 Genus Neocometes Schaub et Zapfe, 1953 427 Type species: Neocometes brunonis Schaub et Zapfe, 1953; Neudorf Spalte 1, Middle 428 Miocene, Slovakia. 429 430 Neocometes brunonis (Schaub and Zapfe, 1953) 431 Holotype and type locality: left mandible with m1-3 (O.E. 307; housed in the Natural 432 History Museum Basel); Devínska Nová Ves “Spalte 1”, MN6. = DNV–Štokeravská vápenka 433 (originally, Horizont D der Neudorfer Spalte, Helvétien). 434 Material and measurements: right M1 (KGP-6a.1: L=2.17, W=1.15); right M2 (KGP-6a.2: 435 L=1.92, W=1.55); right m1-m3 (Z14596/1: m1 – L= -, W=1.23; m2 – L=1.71, W=1.24; m3 – 436 L=1.36, W=1.16); left mandible fragment m2-m3 (DB920002, m2 – L=1.69, W=1.51; m3 – 437 L=1.65, W=1.41) right m1 (KGP-MS64: L=2.18, W=1.28); right m2 (Z14596/2: L=1.81, 438 W=1.56); left m3 (KGP-6a.3: L=1.65 ,W=1.32). 439 Description: M1 (Fig. 7(A)). There is only one single M1 available in DNV–Bonanza site. 440 This molar has an elongated overall morphology, with a rounded anterior part. The anteroloph 441 is long and well-developed. Between the anterior transversal ridge and the anteroloph, the 442 syncline Ia is left open. The paracone and protoloph are thick leaving a deep and narrow 18 443 syncline I. The syncline II is deep and completely splits the tooth into two well-differentiated 444 parts. The mesoloph is thick, structure anteriorly directed that reaches the margin of the tooth. 445 The posteroloph is thin and connects directly with the metacone, thus resulting in a narrow 446 and short IV syncline. All synclines are open labially. 447 M2 (Fig. 7(B)). There is only one M2 recovered from this site, and it shows a medium 448 stage of wear. The syncline Ia is vaguely present, being completely closed and almost 449 integrated into the anterior transversal ridge. The syncline I is deep and narrow. The protoloph 450 connects the paracone and the protocone directly. The tooth is divided into two well- 451 differentiated parts due to the deep and narrow syncline II, as seen in the M1. The posteroloph 452 is partly broken in KGP-6a.2. It reaches the metacone, resulting in a closed syncline IV, 453 probably due to its state of wear. The syncline III is short and open in its labial parts, as in the 454 syncline II and I. 455 m1 (Fig. 7(C)). There are two m1 available from DNV–Bonanza: the specimen KGP- 456 MS64 is well-preserved, whereas Z14596/1 presents a high level of wear, making it difficult 457 an accurate description of its occlusal morphology. The m1 has a more elongated overall 458 morphology compared to the upper molars. The syncline Ia and I are joined forming a loop in 459 the anterior part of the tooth, which are closed labially. The anterior transversal ridge is 460 narrow and connects with the labial margin of the tooth. The synclinal II and III are both open 461 lingually and labially. The posterolophid is well-developed and encloses the synclinal IV by 462 reaching the entoconid. The hypolophulid is indistinguishable from the entoconid and the 463 posterolophid. 464 m2. One of the two m2 recovered is associated with a mandible (Z14596/1) with the 465 m1 (previously mentioned) and m3. This tooth row shows a high level of wear. The other 466 specimen available is Z14596/2: it also shows a high level of wear and therefore is difficult to 467 evaluate properly its occlusal morphology. These teeth show an overall square-shaped 19 468 morphology with rounded edges. Synclines I o IV are closed lingually due to the high level of 469 wear. In both specimens, syncline Ia is not visible. Syncline III is open labially in Z14596/2. 470 m3 (Fig. 7(D)). There are two m3 available: the specimen associated with the 471 mandible and show a high level of wear, whereas KGP-6a.3 is less worn. The anterolophid is 472 well-developed following the anterior margin of the teeth. The syncline I is narrow, deep, and 473 turns backwards on the lingual side. The synclines II and III also narrows and are only open 474 labially. The posterolophid is well-developed and encloses the synclinal IV, reaching the 475 entoconid. 476 Remarks: Although Neocometes has been reported in several Central European Early to 477 Middle Miocene localities, it is still a rare genus. In DNV–Bonanza, just a few molars were 478 recovered and ascribed to Neocometes brunonis. In fact, that genus does not register great 479 changes in the morphologic pattern of the molars, and only a few differences have been 480 proposed between N. brunonis and N. similis during the Middle-Late Miocene (Fejfar, 1999; 481 Mörs, 2006; Pineker and Mörs, 2011). As for the upper molars of N. brunonis, the syncline Ia 482 is not present and the anteroloph is being fused with the anteroloph transversal ridge. As a 483 rule, the synclines I, II, III, and IV are open labially in the upper molars in the Neocometes 484 genus (Fejfar, 1999; Mörs, 2006; Pineker and Mörs, 2011). The lower molars of N. brunonis, 485 in turn, have a syncline Ia reduced and anteriorly opened. Conversely, in N. similis the 486 syncline I and Ia are parallel, and they show no anterior opening (Schaub and Zapfe 1953; 487 Fejfar, 1974, 1999). Concerning molar size, there is a significant gradual increase of size 488 registered in this genus through the Miocene. As is seen in the Figure 8, the material 489 recovered from DVN–Bonanza site coincide within the size range for the N. brunonis type 490 material from DNV–Štokeravská vápenka and are larger compared with N. similis from 491 Erkerstshofen used for comparison (Fejfar, 1974). 492 20 493 Subfamily Anomalomyinae Schaub, 1925 494 Genus Anomalomys Gaillard, 1900 495 Type species: Anomalomys gaudryi Gaillard, 1900. La Grive-Saint-Alban (Ìsere). Middle 496 Miocene, France. 497 498 Anomalomys sp. 499 Material and measurements: right M2 (KGP-6a.21: L=1.62, W=1.20) 500 Description: M2 (Fig. 7(E)). A single specimen was recovered from DVN–Bonanza site: it 501 shows a medium stage of wear, but it still retains many occlusal morphological structures to 502 allow an accurate description. KGP-6a.21 is square-shaped with a very high level of 503 hypsodonty. The anteroloph is wide and occupies the anterior margin of the tooth from the 504 protocone to the paracone. Due to the high degree of wear in the anterior part of the tooth, the 505 anteromesoloph is almost and almost completely merged with the anteroloph. The anterior 506 anterosinus is highly reduced and the posterior anterosinus is narrow and deep. The protoloph 507 is wide and merges the paracone with the entoloph. The sinus is narrow and is anteriorly 508 directed. The mesoloph is of medium length and does not merge with the paracone posterior 509 spur, leaving the mesosinus open labially. The posteroloph is wide and merges in a 510 continuous crest the hypocone and the metacone following the posterior margin of the tooth. 511 The metaloph is connected with the metacone but does not reach the entoloph, leaving an 512 open connection between the mesosinus and the posterosinus. 513 Remarks: The taxonomical position of the genus Anomalomys has been under discussion 514 several times in the literature. Some authors considered Anomalomys to be part of the 515 subfamily Anomalomyinae within the family Cricetidae (Mörs, 2008). However, Fejfar 516 (1972) attributed this rodent clade within the family Spalacidae. Nowadays, Anomalomys is 517 mostly considered as a member of the separate family Anomalomyidae (Bruijn and Saraç 21 518 1991; Bolliger, 1996, 1999; Kalthoff, 2000; Hír, 2001; Nesin and Kovalchuk, 2021). 519 Although KGP-6a.21 found in DNV–Bonanza shows sufficient morphological characteristics 520 that allow its attribution to the genus Anomalomys, an attribution at the species level is not 521 possible. 522 523 5. Discussion 524 Devínska Nová Ves set of sites have a key role in understanding the biostratigraphy of 525 the Badenian Paratethys regional stage (15.97-12.70 Ma, early Astaracian, MN6; Kováč et al., 526 2018; Sabol et al., 2021). Devínska Nová Ves is composed by three different sites DNV– 527 Štokeravská vápenka, DNV–Sandberg and DNV–Bonanza. DNV–Štokeravská vápenka 528 recorded terrestrial fossil vertebrates, DNV–Sandberg yielded predominantly a marine fossil 529 record, while DNV – Bonanza shows a poor transition between the two different 530 environments. DNV– Bonanza has a short mammal record, and in this work, we focused our 531 attention on the cricetid and eomyid fauna recovered from this site, as well as on its 532 biostratigraphic/biochronologic and paleoenvironmental implications. 533 The main cricetid representative in this site is Democricetodon vindobonensis, a small 534 sized fossil rodent only known so far from the DNV–Štokeravská vápenka. The DNV– 535 Bonanza remains of D. vindobonensis coincide in morphology and size with the specimens 536 found in DVN–Štokeravská vápenka. This fact allows us to give an approximate correlation 537 to the lower MN6. There is also the presence of ?Megacricetodon sp. in DNV–Bonanza, but 538 only mandible fragments have been recovered so far (Sabol, 2005a). 539 The representatives of both genera (Megacricetodon and Democricetodon) represent, 540 together with other rodent taxa such as Eumyarion weinfurteri, Anomalomys sp., and 541 Neocometes brunonis the result of immigration waves from Asia into Europe during the Early 542 Miocene (Kälin, 1999). The presence of these taxa in DNV–Bonanza is indicative of a great 22 543 biodiversity of rodents, indicating a rich environment. In DNV–Bonanza there are several 544 representatives of these genera (Sabol, 2005a; Mažgút, 2010), probably descendants of Early 545 Miocene species. Some remains of Neocometes brunonis were also recovered from this site. 546 The latter genus is recorded in Europe from the MN4, although there are some findings dated 547 from the MN3 such as in Petersbuch 28 (Mörs, 2008) to the MN7/8: N. brunonis is well 548 known from deposits dated from MN6-MN7/8; and N. aff. brunonis from MN5 site of 549 Schellenfeld 2 (Ziegler, 1995). As to Eumyarion, several isolated teeth were found in DNV– 550 Bonanza site and were referred to Eumyarion cf. latior. A sole specimen recovered from 551 Bonanza was attributed to Anomalomys sp. Karydomys, is known in several localities from 552 Central Europe during a short period of time, i.e., during the MN5 until MN6, corresponding 553 to the Orleanian/Astaracian transition (Mörs and Kalthoff, 2004). 554 Concerning the eomyid fauna, two different taxa have been recovered in DNV– 555 Bonanza: Eomyops sp. (one molar) and Keramidomys carpathicus (two molars). The latter 556 taxon is well known in Central Europe during the MN6 (Sabol and Holec, 2002). 557 Based on the rodent assemblage from the DNV–Bonanza site, not limited to the 558 cricetid and eomyid fauna, but also considering the presence of glirids such as Myoglis meini 559 and Muscardinus sansanensis, among others (Sabol, 2005a; Sabol et al., 2021), as well as of 560 the sciurid Csakvaromys bredai (Sabol, 2005a), the site can be correlated to the MN6 zone 561 (see also Sabol et al., 2021). This is further supported by the insectivore records from the 562 same site (Sabol, 2005b). 563 However, the rodent assemblage from DNV–Bonanza shows several differences with 564 the fauna recorded at the nearby site of DNV–Štokeravská vápenka, therefore appearing to be 565 a subset of the DNV–Štokeravská vápenka site (Sabol, 2005a,b; Mažgút, 2010; Sabol et al., 566 2021). As mentioned before, the Devínska Nová Ves set of sites are dated after the peak of 567 the MMCO after which a major and permanent cooling step occurs. This was one of the major 23 568 paleoclimatic events that took place in Europe at that time. Particularly, in Devínska Nová 569 Ves this is reflected on a decrease in representation of open land elements recorded in the 570 different sites (Sabol and Kováč, 2006). 571 When considering the paleoenvironmental aspects, the rodent assemblage suggests a 572 forested environment, especially with the presence of the eomyids Keramidomys carpathicus 573 and Eomyops sp., as well as a good assemblage of glirids, and the sciurid Csakvaromys bredai 574 (Fejfar, 1974; Sabol 2005 a,b; Sabol et al., 2021). The presence of several taxa of cricetid 575 rodents, which are characterized as generalist species (Kälin, 1999), cannot provide further 576 information on the palaeoecology. Therefore, for a complete paleoenvironmental 577 reconstruction, the whole fossil assemblage must be considered. The study of the 578 Eulipotyphla depicted the palaeoenvironment as a forested area close to water bodies (Sabol, 579 2005b), which supports the results presented here. Moreover, the presence of several marine 580 taxa, such as fish, sharks, seals, and marine bivalves, indicates a mixed insular or peninsular 581 area, with subtropical forests close to freshwater bodies and in the vicinity of a saltwater sea 582 (Sabol 2005a,b; Sabol et al., 2021). 583 584 6. Conclusions 585 The rodent assemblage of DNV– Bonanza site shows a significant number of cricetid 586 and eomyid specimens. In this study, we were able to identify and describe several cricetid 587 taxa such as Eumyarion cf. latior, Democricetodon vindobonensis, and Karydomys sp., 588 together with Anomalomys sp. and Neocometes brunonis. A total of two eomyid species have 589 been identified, being Keramidomys carpathicus and Eomyops sp. Apart from the described 590 rodent taxa, DNV–Bonanza has a larger rodent assemblage with several taxa of glirid and 591 sciurids. This association allows us to correlate this site, as well as its neighbouring locality, 592 to the lower part of the MN6. In terms of the paleoenvironment, the rodent assemblage 24 593 confirms a forested dominated environment, especially given the presence of eomyids such as 594 Keramidomys carpathicus, and Eomyops sp., but also a good assemblage of glirids and the 595 sciurid Csakvaromys bredai. Meanwhile, the cricetid fauna is considered to be generalist, so it 596 cannot give further information on this behalf. Previous studies on the insectivore fauna of 597 this locality further support the forested environment suggested. 598 599 Acknowledgements 600 We thank Dr. Ursula Gölich from the Vienna Natural History Museum and Dr. Jan 601 Wagner, and Dr. Boris Ekrt from the Prague National Museum for the opportunity to review 602 the classic rodent fauna collection from Devínska Nová Ves –Štokeravská vápenka and use it 603 as comparison material for this study. We also thank the reviewers Dr. János Hír and Dr. 604 Thomas Mörs and the editor, Dr. Gilles Escarguel, for the comments that greatly improved 605 the manuscript. The present work has been supported by the Spanish Agencia Estatal de 606 Investigación (I+D+i PID2020-117289GBI00 funded by MCIN/ 607 AEI/10.13039/501100011033/) and the Generalitat de Catalunya (CERCA Program). It is also 608 funded by the NSP (National Scholarship program of Slovak Republic for the Support of 609 Mobility of University Researchers, ID 33758) to S.J.V. The research project has also been 610 supported by the UK grant of the Comenius University in Bratislava (Project No. 611 UK/75/2021; I.B.S), the APVV grant agency of the Slovak Republic (Project No. APVV-20- 612 0079; I.B.S; M.S), by the institutional support RVO67985831 of the Institute of Geology of 613 the Czech Academy of Sciences, the Vedecká Grantová Agentúra MŠVVaŠ SR a SAV 614 (1/0533/21) to PJ. ÀHL is financially supported through the postdoctoral grant ‘Programa 615 Postdoctoral Beatriu de Pinós de la Secretaria d’Universitats i Recerca del Departament 616 d’Empresa i Coneixement de la Generalitat de Catalunya’ (2019 BP 00154). 617 25 618 References 619 Alston, E. R., 1876. On the classification of the order Glires. Proceedings of the Zoological 620 Society of London, 1876, 61–98. 621 Álvarez Sierra, M. Á., 1987. Estudio sistemático y bioestratigráfico de los Eomyidae 622 (Rodentia) del Oligoceno superior y el Mioceno inferior español. 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Journal of Vertebrate 820 Paleontology 21, 367–385. 821 Van Dam, J.A., Abdul Aziz, H., Álvarez Sierra, M.A., Hilgen, F.J., Van den Hoek Ostende, 822 L.W., Lourens, L., Mein, P., Van der Meulen, A.J., Peláez-Campomanes, P., 2006. 823 Long-period astronomical forcing of mammal turnover. Nature 443, 687–691. 824 Wessels, W., Reumer, B.M., 2009. Democricetodon and Megacricetodon (Mammalia, 1531 825 Cricetidae) from the Miocene of Sandelzhausen, Southern Germany. Palaeontologische 826 Zeitschrift 83, 187–205. 827 Winge, H., 1887. Jordfundne og nulevende gnavere (Rodentia) fra Lagoa Santa, Minas 828 Gerais, Brasilien. E. Museo Lund 1, 1–200. 829 Ziegler, R., 1995. Die untermiozänen Kleinsäugerfaunen aus den Süßwasserkalken von 830 Engelswies und Schellenfeld bei Sigmaringen (Baden-Württemberg). Stuttgarter 831 Beiträge zur Naturkunde B 228, 1–53. 832 833 32 834 Figure captions 835 836 Fig. 1. A–B: Location of the DNV–Bonanza site on northern slopes of Devínska Kobyla Hill 837 near Devínska Nova Ves – Štokeravská vápenka. C: Generalized section through the 838 sediments of Bonanza (according to Ivanov, 1998). I – fine limestone debris; 2 – white lime 839 sand; 3 – disaggregating sandstone with a higher content of muscovite; 4 – solid, light yellow 840 marlstone with a great quantity of fossils; 5 – big boulders with white lime matter: 6 – 841 greenish sand with interbeds of white lime matter; 7–11 – layers with coarse-grained, 842 disaggregating sandstone without fossils to the fossiliferous marl, rich in fossils; 12 – white 843 calciferous sandstone; 13 – yellowish-white sand with a large quantity of fauna; 14–17 – 844 greenish to light sandstone, the biggest quantity of fossils are contained in the layer No. 17; a 845 – Holocene humus-carbonate soil; b – Lias limestone; c – tectonic faults. Modified from 846 Sabol (2005a). 33 847 848 Fig. 2. Scanning electron microscope (SEM) micrographs. Eumyarion cf. latior. A: left M1 849 KGP-6a.4, B: right m1 KGP-9.2 (reversed), C: right m2 KGP-6a.5 (reversed), D: right m3 850 KGP-9.3 (reversed). Democricetodon vindobonensis. E: left M1 KGP-6a.7, F: right M1 KGP- 851 6a.14 (reversed), G: left M1 KGP-MS29, H: left M2 KGP-MS31, I: left M2 KGP-6a.12, J: 34 852 right M3 KGP-MS34 (reversed), K: left m1 KGP-6a.10, L: right m1 KGP-MS35 (reversed), 853 M: right m1 KGP-MS36 (reversed), N: left m2 KGP-6a.11, O: left m2 KGP-6a.13, P: left m2 854 6a.14, Q: right m2 KGP-MS38 (reversed), R: left m2 KGP-MS39, S: left m2 KGP-MS40, T: 855 left m3 KGP-MS61. Karydomys sp. U: right M2 KGP-MS63 (reversed). Keramidomys 856 carpathicus. V: right M1 KGP-MS69 (reversed), W: right M1 KGP-6a.18 (reversed). 857 Eomyops sp. X: right P4 KGP-6a.17 (reversed). 35 858 859 Fig. 3. Partial skull of Eumyarion cf. latior (KGP-1). A: left lateral view with M1 fragment. 860 B: Ventral view of the skull with fragment of left M1. Partial skull remains of 861 Democricetodon vindobonensis (KGP-17, C–D). C: dorsal view of the skull highly damaged. 862 D: Ventral view of the skull with the right tooth row preserved. 36 863 864 Fig. 4. Length/Width scatter plot of Eumyarion cf. latior from DNV- Bonanza site (black 865 dots), DNV- Štokeravská vápenka (squares: material from Vienna Natural History Museum; 866 triangles: material from Prague National Museum). The ellipses show the 95% confidence 867 interval for Eumyarion latior (Anwil. Fejfar, 1974), Eumyarion weinfurteri (Sandelzhausen. 868 Bruijn, 2009), and Eumyarion bifidus (Sandelzhausen, and other sites. Bruijn, 2009; Heissig, 869 1990). 37 870 871 Fig. 5. Length/Width scatter plot of Democricetodon vindobonensis from DNV–Bonanza site 872 (black dots), DNV–Štokeravská vápenka (squares: material from Vienna Natural History 873 Museum; triangles: material from Prague National Museum). The ellipses show the 95% 874 confidence interval for Democricetodon gracilis (Sandelzhausen. Wessels and Reumer, 38 875 2009), Democricetodon franconicus (Dolnice 1-3 and Ořechov. Fejfar, 1974), 876 Democricetodon mutilus (Sandelzhausen. Wessels and Reumer, 2009), Democricetodon 877 vindobonensis (DNV–Štokeravská vápenka, Neudorf. Fejfar, 1974). 39 878 879 Fig. 6: Length/Width scatter plots of Keramidomys carpathicus from DNV–Bonanza site 880 (black dots), DNV–Štokeravská vápenka (squares: material from Vienna Natural History 881 Museum; triangles: material from Prague National Museum), and Keramidomys thaleri is 40 882 represented by blue dots. The ellipses show the 95% confidence interval for Keramidomys 883 carpathicus from DNV–Štokeravská vápenka (Fejfar, 1974). 884 885 Fig. 7. Scanning electron microscope (SEM) micrographs. Neocometes brunonis. A: right M1 886 KGP-6a.1 (reversed), B: right M2 KGP-6a.2 (reversed), C: right m1 KGP-MS64 (reversed), 887 D: left m3 KGP-6a.3. Anomalomys sp. E: right M2 KGP-6a.21 (reversed). 888 889 Fig. 8. Length/Width scatter plot of Neocometes brunonis from DNV- Bonanza site (black 890 dots), DNV- Štokeravská vápenka (squares: material from Vienna Natural History Museum). 891 The ellipses show the 95% confidence interval for Neocometes brunonis (DNV–Štokeravská 41 892 vápenka. Schaub and Zapfe, 1953; Fejfar, 1974), and Neocometes similis (Erkertshofen. 893 Fahlbusch, 1966; Fejfar, 1974). 894 42