Palaeontographia Italica 90 2003 149-161 tavv. Pisa, 2005

Eomyidae and Gliridae from Rudabánya

GUDRUN DAXNER-HÖCK*

KEY WORDS — Rodents, Late Miocene, MN9, Hungary

ABSTRACT — Six glirid species are described from the Late Miocene of Rudabánya: Glirulus lissiensis Hugueney & Mein 1965, Paraglirulus werenfelsi Engesser 1972, Muscardinus hispanicus De Bruijn 1966, Muscardinus cf. vallesiensis (Hartenberger 1966), Glis minor Kowalski 1963 and Myoglis cf. ucrainicus Nesin & Kowalski 1997. Although descending from Early Miocene glirid lineages, three out of six species are restricted to the Late Miocene. The eomyid from Rudabánya, Eomyops catalaunicus, is known to range from MN9 to MN10. This glirid-eomyid association is characteristic for a Vallesian age (MN9). The Rudabánya glirids most probably lived in arboreal thickets and forests, feeding on fruits, seeds, berries and insects as their living relatives do.

RIASSUNTO — Vengono descritte sei specie di gliridi dalla località del Miocene superiore (MN9) di Rudabánya (Ungheria): Glirulus lis- siensis Hugueney & Mein 1965, Paraglirulus werenfelsi Engesser 1972, Muscardinus hispanicus De Bruijn 1966, Muscardinus cf. valle- siensis (Hartenberger 1966), Glis minor Kowalski 1963 and Myoglis cf. ucrainicus Nesin & Kowalski 1997. Sebbene rappresentanti di linee evolutive che risalgono al Miocene Inferiore, tre delle sei specie presenti a Rudabánya sono limitate al Miocene superiore. Un unico eomyide è presente a Rudabánya, Eomyops catalaunicus. Questa forma è nota per avere una distribuzione che abbraccia le zone MN9 e MN10. L’associazione gliridi-eomyidi di Rudabánya è indicativa di una età Vallesiana (MN9). I gliridi di Rudabánya molto probabilmente vivevano in boschi e foreste, cibandosi di frutta, semi ed insetti, come fanno i loro corrispondenti viventi.

* Gudrun Daxner-Höck, Dept. Geol.-Pal., Museum of Natural History, Burgring 7, A-1010 Vienna, Austria. e-mail: [email protected]

INTRODUCTION ments a Leica WILD M8 stereo microscope was used. The famous vertebrate locality Rudabánya is sit- uated in NE Hungary. Since the first finding of the TABLE 1 - The total number of teeth of glirids and primate Rudapithecus in 1965 by G. Hernyak, there eomyids from Rudabánya were field activities for more than 30 years. The first systematic excavations of fossils were carried Kretzoi Kordos Daxner-Höck out by M. Kretzoi from 1971 to 1978. Several exca- 1971-1978 1985-1998 1994 vations followed from 1985 to 1998 under the lead- ership of L. Kordos, R. Bernor, P. Andrews, M. Eomyops catalaunicus 4 13 Armour-Chelu and D.R. Begun (cfr. Bernor et al., Glis lissiensis 7 2003). During all these field seasons small mam- Paraglirulus werenfelsi 6 1 5 mals were collected by wet screening, but the result Muscardinus hispanicus 23 was relatively poor. Primarily small vertebrates Muscardinus cf. vallesiensis 11 were recovered from the black mud, black clay and Glis minor 28 315 gray mud layers of the localities I and II. Only for Myoglis cf. ucrainicus 48 13 28 small samples from different layers of locality II 89° 17* 72* were sieves with mesh sizes of 0.5 mm used in the summer 1994. From these samples even the finest fraction of washing remains was carefully sorted. It TAXONOMIC DESCRIPTION included small sized rodents (Glirulus lissiensis, Eomyops catalaunicus) which were poorly repre- Eomyidae Deperet & Douxami, 1902 sented or unknown from Rudabánya before. The Eomyops Engesser, 1979 total number of teeth of glirids and eomyids is list- Eomyops catalaunicus (Hartenberger, 1966) ed in Table 1. The specimens collected by M. (Text-fig. 1/1-8; Tab. 2) Kretzoi from 1971 to 1978 are indicated by (°) in tables and figures. All specimens collected by L. 1974 Leptodontomys catalaunicus (Hartenberger) - Kretzoi et Kordos from 1985 to 1998, and by Daxner-Höck in al.: p. 375 the 1994 will be stored in the collection of the Geological Institute of Hungary and are indicated Type locality by (*) in tables and figures. Can Llobateres (Spain), MN9 SEM-photos of the fossils were taken by a Philips XL 20 scanning electron microscope at the Measurements Biozentrum/University of Vienna. For measure- see table 2 150 GUDRUN DAXNER-HÖCK

Text-fig. 1 - Teeth of Eomyops catalaunicus (Hartenberger, 1966) from Rudabánya (Hungary). 1) °P4 left; 2) *M1/2 left; 3) *M1/2 left; 4) *M1/2 right; 5) *p right; 6) *m left; 7) *m left; 8) *M1/2 right. ° = Coll. Kretzoi / Budapest, * = Coll. Kordos / Budapest. All right side teeth are 4 1/2 1/2 figured as if they were from the left side.

TABLE 2 - Measurements (in mm) of Eomyops catalaunicus from Rudabánya

E. catalaunicus Fig. P4 M1/2 p4 m1/2 m3 LWLWLWLWLW

°P4l 1/1 0.90 1.00 °M1/2l 0.90 1.00 *M1/2l 1/2 0.90 0.95 *M1/2l 1/3 0.90 1.00 *M1/2r 1/8 0.90 1.00 *M1/2r 1/4 0.90 1.00 *M1/2l 0.85 0.90 *M1/2l 0.85 1.00 *M1/2r 0.80 0.95 *M1/2r 0.85 1.00 *p4r 1/5 0.80 0.75 °m1/2r 0.92 0.97 *m1/2l 1/6 1.05 1.00 *m1/2l 1/7 1.05 0.90 *m1/2r 1.00 0.90 *m1/2r 1.00 0.95 °m3l 0.85 0.83

Description arm of the protocone. The protoloph and the met- Maxillary P4 (Text-fig. 1/1) is trapezoidal in aloph contact the anterior arms of the protocone occlusal view, and the buccal wall is slightly longer and hypocone, respectively. The mesoloph is short than the lingual one. Only one, strongly worn tooth and directed forwards. The V-shaped lingual sinus exists which does not show dental morphology in is directed slightly forwards in three of four M1/2. all details. The mesoloph is long and ends in a There are three roots: a lingual and two buccal small mesostyle. There is no anteroloph. The lin- ones. gual sinus is transversely directed. P4 has three Mandibular p (Text-fig. 1/5) is longer than 4 roots: a lingual one and two buccal ones. wide. There is no anterolophid. The large proto- Maxillary M1/2 (Text-fig. 1/2-4, 8) is almost conid and the small metaconid are situated very square in occlusal outline. The corners are round- close to one another and the short metalophid is ed. The buccal part of the anteroloph is longer than interrupted. The mesolophid is long, it ends in a the lingual one. It is connected with the anterior small mesostyle. There is a small mesoconulid EOMYIDAE AND GLIRIDAE FROM RUDABÁNYA 151 anterior to the mesolophid. The strong hypolophid TABLE 3 - Measurements (in mm) of Glirulus lissiensis extends to the posterior arm of the hypoconid. The from Rudabánya posterolophid is short. P has two roots, an anteri- 4 or and a posterior one. G. lissiensis Fig. M1 m1 m2 m3 Mandibular m (Text-fig. 1/6-7) are rectangular LWL WLWLW 1/2 in occlusal outline. The corners are rounded. The *M1r 2/1 0.90 0.90 lingual cusps are situated opposite to the buccal *m1r 2/2 0.95 0.85 cusps. The anterolophid has a lingual and a buccal *m1r 1.00 0.85 arm. The metalophid extends to the protoconid, *m2l 2/4 0.95 0.90 the hypolophid extends to the posterior arm of the *m2l 0.95 0.95 hypoconid. The mesolophid is long in one and *m2r 2/3 0.95 0.95 short in three other teeth. The m have three 1/2 *m3l 2/5 0.90 0.85 roots: a posterior and two anterior ones. Mandibular m is strongly worn. The lingual 3 arm of the anterolophid is longer than the buccal one. The mesolophid is strong and ends in a mesostylid on the lingual border. Description M2 (Text-fig. 2/1) is rhomboidal in occlusal view, Discussion small and concave. It is wider anteriorly than pos- In the Early Miocene eomyids were represented teriorly. The five transverse ridges are directed by eight genera which were widely distributed all slightly forwards and lingually connected with the over Europe. Successively these Early Miocene endoloph. The endoloph is complete. The anterior genera became extinct. However, at the MN4/5 centroloph is long and connected with the transition two new genera appeared, the smallest endoloph. Buccally it ends free forming a small that ever existed in Europe: Eomyops and Kerami- conulus. The posterior centroloph is bent and con- domys. nected with the anterior centroloph by a small lon- Eomyops is represented in many vertebrate fau- gitudinal ridge. There are three long extra ridges. nas by some isolated teeth only, but in a few locali- There are buccal connections of the anteroloph ties it is abundant. It persisted from MN5 (late and the protoloph, and of posteroloph, metaloph Early Miocene) to MN17 (Pleistocene). There are and posterior centroloph, respectively. Roots are four species (Engesser, 1999): Eomyops hebeiseni not preserved. Mandibular m (Text-fig. 2/2-5) is rectangular Kälin, 1997, E. oppligeri Engesser, 1990, E. catalau- 1/2 in occlusal view. The endolophid of m is complete, nicus (Hartenberger, 1966) and E. bodvanus 1 but it shows a notch behind the centrolophid in m . (Janossy, 1972). 2 Eomyops hebeiseni (type locality Katzloch, The centrolophid is long in all specimens, but Switzerland; MN6) is the oldest one. It is restricted never reaches the buccal border of the occlusal sur- face. The m and two out of three m ’s have two to MN6. Eomyops oppligeri (type locality Anwil, 1 2 Switzerland; MN8) ranges from MN7 to MN8. It is extra ridges between their anterolophid and met- alophid. All m ’s and m ’s have three further extra uncertain whether or not Eomyops from Podlesice 1 2 (MN14) belongs to this species. Eomyops catalauni- ridges: one on each side of the centrolophid and cus (type locality Can Llobateres, Spain; MN9) one in the posterior valley. Mandibular m (Text-fig. 2/6) is similar with m , ranges from MN9 to MN10. Eomyops aff. or cf. 3 2 catalaunicus determinations have longer chrono- but narrow in its posterior part. The lower molars logic ranges. have two roots. In molar morphology there are only small dif- ferences between the four species. However, E. Discussion catalaunicus is somewhat larger than E. oppligeri Morphology and size indicate that the Glirulus and smaller than E. hebeiseni and E. bodvanus. E. from Rudabánya is referable to G. lissiensis. The catalaunicus is a characteristic element of species ranges from MN4 (Oberdorf, Austria; de European Late Miocene vertebrate faunas. All of Bruijn, 1998) to MN13 (Lissieu, in France, the type these findings – including Rudabánya – are similar locality; Hugueney & Mein, 1965). Some Middle to the type material from Can Llobateres in tooth and Late Miocene occurrences include: Belchátow pattern and size. A and B (Kowalski, 1997), Anwil (Engesser, 1972), Suchomasty (Fejfar, 1989), Kohfidisch (Bachmayer & Wilson, 1983), Eichkogel (Daxner-Höck & de Bruijn, 1981), Amberieu 3 (Mein, 1984). Further Gliridae Thomas, 1897 Vallesian occurrences of Glirulus lissiensis in Dryomyinae de Bruijn, 1967 Austria are: Richardhof-Golfplatz, Richardhof- Glirulus Thomas, 1897 Wald and Schernham. Glirulus lissiensis (Hugueney & Mein, 1965) Glirulus lissiensis belongs to the same group as (Text-fig. 2/1-5; Tab. 3) the living and extinct species: G. japonicus (extant), G. pusillus (Pleistocene), G. conjunctus (Middle- Type locality Late Miocene / MN6-9), G. diremptus (Early-Middle Lissieu (France), MN13 Miocene / MN4-5) and G. ekremi (Early Miocene / MN3). The dental pattern and size of this group Measurements shows stability through time. Even G. ekremi, the See table 3 oldest known species, differs from the living G. 152 GUDRUN DAXNER-HÖCK

Text-fig. 2 - Teeth of glirids from Rudabánya (Hungary). Glirulus lissiensis (Hugueney & Mein, 1965): 1) *M2 right; 2) *m right; 3) *m 1 1 left; 4) *m left; 5) *m left. Paraglirulus werenfelsi Engesser, 1972; 6) *P4 right; 7) *M1 right; 8) *M2 left; 9)*p left; 10) °m left; 11) *m 2 3 4 1 1/2 right; 12) *m right. Muscardinus hispanicus de Bruijn, 1966: 13) *M1 left; 14) °M1 right; 15) *M2 left; 16) *m left. Muscardinus cf. valle- 1/2 2 siensis (Hartenberger, 1966): 17) *D4 left; 18) °m left. All right side teeth are figured as if they were from the left side. 2 EOMYIDAE AND GLIRIDAE FROM RUDABÁNYA 153

TABLE 4 - Measurements (in mm) of Paraglirulus werenfelsi from Rudabánya

P. werenfelsi Fig. P4 M1/2 p4 m1/2 m3 LWLWLWLWLW

*P4r 2/6 1.00 1.10 *M1r 2/7 1.25 1.30 °M1l 1.30 1.35 *M2l 2/8 1.25 1.40 °M2l 1.30 1.45 °M2r 1.35 1.50 *p4l 2/9 1.00 0.85 *m1/2r 2/11 1.30 1.25 *m1/2r 2/12 1.35 1.25 °m1/2l 2/10 1.30 1.25 °m1/2r 1.15 1.10 °m1/2r 1.30 1.25

japonicus in small details only (de Bruijn, 1998: Discussion 117). Thus, species-determination on the basis of The first record of P. werenfelsi was thought to limited material is questionable. be from the Early Miocene (MN4) of Oberdorf in As a glider (Mein & Romaggi, 1991) G. lissiensis Austria (Daxner-Höck, 1998: 380). Since de Bruijn inhabited forested biotopes and lived in trees. (1998: 117) identified the Oberdorf-specimens as Glirulus (Paraglirulus) sp., the first record of P. werenfelsi is MN5, from the locality Obergänserndorf , Austria (Daxner-Höck, 1998: Paraglirulus Engesser, 1972 379). Further Middle to Late Miocene occurrences Paraglirulus werenfelsi Engesser, 1972 of P. werenfelsi are listed in Kowalski (1997: 187). (Text-fig. 2/6-12; Tab. 4) The last records are from the Late Miocene (Can 1974 Paraglirulus cf. lissiensis (Hugueney & Mein) - Kretzoi et Llobateres in Spain, Rudabánya in Hungary, al.; p. 357. Gritsiv in Ukraine, Belchatóv A in Poland, Richardhof-Golfplatz and Götzendorf in Austria: Type locality all MN9, and Richardhof-Wald in Austria: MN10). Anwil (Switzerland), MN8 Paraglirulus agelakisi (Van der Meulen & de Measurements Bruijn, 1982) from Aliveri in Greece (MN4) primari- See table 4 ly differs in having a shorter anterior centroloph and lacking lingual ornamentation of upper molars. Description The occlusal surface of the teeth is concave. The Glirinae Thomas, 1897 main ridges are higher than the extra ridges. The Muscardinus Kaup, 1829 main ridges connect the endoloph in upper molars 4 Muscardinus hispanicus de Bruijn, 1966 and the P . The lingual side of the upper teeth is (Text-fig. 2/13-16; Tab. 5) ornamented. In lower molars and the p the 4 endolophid is interrupted posteriorly to the cen- 1974 Muscardinus sp. - Kretzoi et al.: p. 375 trolophid. Maxillary P4 (Text-fig. 2/6) has a rounded Type locality occlusal outline. The anterior centroloph does not Pedregueras 2C (Spain), MN9 contact the endoloph. The roots are not preserved. Maxillary M1/2 (Text-fig. 2/7-8) are square in Measurements occlusal outline. There are five main ridges and See table 5 four extra ridges (posterior centroloph included). Except in one out of four M1/2’s the anterior cen- Description troloph connects the endoloph. The metaloph con- M1 (Text-fig. 2/13-14) is trapezoidal in occlusal nects the posteroloph buccally. There are no roots outline. Its buccal side is longer than the lingual preserved. one, and it is narrowed in its anterior part. There Mandibular p (Text-fig. 2/9) corresponds with are five main ridges, directed towards the postero- 4 the type material from Anwil (Engesser, 1972: Abb. lingual part of the tooth. The anteroloph and the 75/2). There are two roots in p . posteroloph end free on the lingual and buccal 4 Mandibular m (Text-fig. 2/ 10-12) is rectangu- sides. The oblique protoloph connects lingually 1/2 lar in occlusal outline. In addition to the four main with the anterior centroloph and the metaloph. ridges, there is a long centrolophid and there are There are two extra buccal ridges, a short anterior four to five extra ridges. The m has three roots: one, and the posterior centroloph of at least medi- 1 one anterior and two posterior. um length. Maxillary M1 has three roots, a lingual one and two buccal ones. 154 GUDRUN DAXNER-HÖCK

TABLE 5 - Measurements (in mm) of Muscardinus hispan- 1974 Muscardinus sp. - Kretzoi et al.: p. 375 icus from Rudabánya Type locality M. hispanicus Fig. M1 M2 m2 Can Llobateres (Spain), MN9. L WLWL W Measurements *M1l 1.40 1.15 See table 6 °M1l 2/13 1.40 1.10 °M1r 2/14 1.40 1.20 TABLE 6 - Measurements (in mm) of Muscardinus cf. *M2l 2/15 1.15 1.15 vallesiensis from Rudabánya *m2l 2/16 1.20 1.05 M. cf. vallesiensis Fig. D4 m2 LWL W

Maxillary M2 (Text-fig. 2/15) is almost square in *D4l 2/17 1.00 1.20 occlusal outline. There are six transverse ridges °m2l 2/18 1.50 1.40 which are connected lingually by an endoloph. Only the two posterior ridges have a buccal con- nection. The transverse lophs are separated by five valleys of decreasing distance from anterior to pos- Description terior. The roots are not preserved. Maxillary D4 (Text-fig. 2/17) is triangular in Mandibular m (Text-fig. 16) is rectangular in occlusal outline. The tooth has five lophs. The pro- 2 occlusal outline. The antero-buccal corner is tolph and the metaloph form a lingual V-shaped rounded. There are six transverse lophs, more or connection. The short anteroloph has an antero- less convex mesially. The anterior and the posterior buccal position. The anterior centroloph is short, pair of lophs are connected on the buccal and lin- the posterior centroloph slightly longer, forming an gual side. The roots are not preserved. U-shaped buccal connection with the metaloph. The roots are not preserved. Mandibular m (Text-fig. 2/18) has a rectangular Discussion 2 Five teeth of a small Muscardinus from occlusal outline. There are six transverse lophs, the Rudabánya were identified as M. hispanicus anterior ones are curved slightly mesially; the two because of resemblance in size and molar pattern. posterior ones are straight. The anterior pair of Muscardinus hispanicus ranges from MN9 to lophs is connected lingually and buccally. There are MN12 all over Europe. In Central Europe its range no roots preserved. is limited to the Vallesian. The species was distrib- uted from Southwestern Europe (Spain: Discussion Pedregueras 2C, Carrilanga 1, Can Llobateres, all For the time being the two isolated Muscardinus MN9; Peralejos, Masia del Barbo, MN10) to teeth are assigned to M. cf. vallesiensis because Western (France: Montredon, MN10; Castelnou 1, they differ from M. hispanicus in larger sizes and MN12) and across Central Europe (Germany: the more complex pattern of D4. The stratigraphic Marktl, Hammerschmiede; Poland: Belchatów A; range of M. vallesiensis is restricted to the Austria: Mariathal, Richardhof-Golfplatz and Vallesian: Can Llobateres (Hartenberger, 1966), Götzendorf, all MN9 and Richardhof-Wald, MN10) Castell de Barbera (Aguilar et al., 1979), to the East and Southeast (Romania: Comanesti Götzendorf (Daxner-Höck, 1997), Richardhof- 2b, MN9; Greece: Kastellios Hill, MN10). Golfplatz all MN9, and Richardhof-Wald MN10. Size and molar pattern set the Rudabánya pop- ulation apart from the Middle Miocene species M. thaleri and M. sansaniensis, and from all Late Glis Brisson, 1762 Miocene and younger species (M. vireti, M. davidi, Glis minor Kowalski, 1956 M. austriacus, M. pliocaenicus, M. dacicus, M. (Text-fig. 3/1-8, 4; Tab. 7) cyclopeus and M. helleri). M. heintzi from Montredon (MN10) differs by larger dimensions. 1974 Glis (s.l.) sp. - Kretzoi et al.; p. 375 There are some differences between M. topachevckii Nesin & Kowalski, 1997 from Gritsiv Type locality (Ukraine, MN9) and M. hispanicus mainly in the Podlesice (Poland), MN14 dental morphology of P4 and m , and in the num- 1 ber of roots. As equivalent teeth are not available Material 4 P4(°), 9 M1(°), 5 M2(°), 3 M3(°), 1 d (°), 2 p (°), 4 from Rudabánya, it is impossible to prove relations 4 4 m (°), 1 m (°), 1 P4(*), 2 M1(*), 1 M2(*), 3 M3(*), 2 d (*), in this respect. However, the teeth from Rudabánya 2 3 4 2 p (*), 2 m (*), 2 m (*), 3 m (*). differ from M. topachevskii by a small anterior 4 1 2 3 extra ridges and the longer posterior centroloph of M1, and by more and longer accessorial extra Measurements ridges of M2. See table 7 and Text-fig. 4. Description Muscardinus cf. vallesiensis (Hartenberger, 1966) The occlusal surface of is molars is slightly con- (Text-fig. 2/17, 18; Tab. 6) cave; the one of the premolars is flat. Upper molars EOMYIDAE AND GLIRIDAE FROM RUDABÁNYA 155

Text-fig. 3 - Teeth of glirids from Rudabánya (Hungary). Late Miocene (MN9). All right side teeth are figured as if they were from the left side. Glis minor Kowalski, 1963: 1) °P4 left; 2) °M1 left; 3) °M2 left; 4) *M3 left; 5/1) *d right; 5/2) *p left; 6) *m right; 7) *m left; 8) *m 4 4 1 2 3 left. Myoglis cf. ucrainicus Nesin & Kowalski, 1997: 9) *P4 right; 10) *M1 right; 11) *M2 right; 12) °M3 right; 13) *p left; 14) *m right; 15) 4 1 *m left; 16) *m right. 2 3 156 GUDRUN DAXNER-HÖCK

TABLE 7 - Measurements (in mm) of Glis minor from buccal connection of the protoloph and anterior Rudabánya centroloph, and in two other specimens there are small extra ridges anterior and posterior to the G. minor min mean max n stdev anterior centroloph. An extra ridge (sometimes very small) is always present between anteroloph P4 L 1.25 1.28 1.30 3 0.02887 and protoloph. P4 W 1.35 1.42 1.50 3 0.07638 Maxillary M3 (Text-fig. 3/4) is trapezoidal in M1 L 1.55 1.63 1.75 11 0.06431 occlusal outline. The tooth narrows posteriorly. M1 W 1.65 1.76 1.85 10 0.06433 The lingual connection of protoloph, metaloph and M2 L 1.55 1.61 1.65 7 0.03780 posteroloph is characteristic. In two out of six M2 W 1.80 1.85 1.90 7 0.05000 specimens the anteroloph connects this endoloph M3 L 1.25 1.30 1.35 6 0.03162 also. Extra ridges are situated posteriorly to the M3 W 1.50 1.54 1.60 6 0.03764 anteroloph and anteriorly to the posteroloph, respectively. The valley between the protoloph and d4 L 1.15 1.17 1.20 3 0.02887 the metaloph is variably ornamented by relicts of d4 W 1.05 1.13 1.20 3 0.07638 the anterior and the posterior centrolophs. p4 L 1.15 1.21 1.30 4 0.07500 The general arrangement of ridges in the lower p4 W 1.10 1.20 1.25 4 0.07071 molars is: the four main transverse ridges m1 L 1.75 1.90 2 (anterolophid, metalophid, mesolophid and pos- m1 W 1.70 1.80 2 terolophid) are thickened at their buccal part. m2 L 1.70 1.75 1.80 6 0.04472 Between them there are three ridges which are m2 W 1.70 1.83 1.95 6 0.08756 shorter, lower, not as thick as the main ridges, and m3 L 1.60 1.66 1.70 4 0.04787 which do not extend to the buccal border. The long m3 W 1.45 1.56 1.65 4 0.08539 mesolophid and the short centrolophid always end free on both sides. Lingual connections of ridges are variable. There are no buccal connections. Mandibular p (Text-fig. 3/5/2) is almost triangu- 4 lar in occlusal outline. According the molar pat- tern, p has four main ridges but only two extra have three roots, a long lingual one and two round- 4 ed buccal ones. All lower molars have two wide ridges that are situated anterior to the mesolophid roots: an anterior and a posterior one. Lower pre- and to the posterolophid. The posterior extra ridge molars have one root. One upper premolar shows, connects the posterolophid lingually. There are that the originally three roots are fused. both, lingual and buccal connections of the The general arrangement of ridges in upper anterolophid and of the metalophid. Mandibular d (Text-fig. 3/5/1) includes three molars is: the four main transverse ridges 4 (anteroloph, protoloph, metaloph and posteroloph) teeth thought to be deciduous, or embryonic per- are thickened at their lingual ends. There is an manent premolars: The occlusal outline is not tri- anterior centroloph, but no posterior centroloph in angular but rounded. No roots are preserved. The P4-M2. Two extra ridges are present: a short extra enamel is very thin. When present, the extra ridges ridge posterior to the anteroloph and a second one are very short and thin, and even the main ridges of medium length which is situated anterior to the are not as thick as in permanent premolars. Mandibular m (Text-fig. 3/6) is rectangular in posteroloph. The four main lophs have no lingual 1 3 occlusal outline. It is longer than the m . Other connection except for the M . Buccal connections 2 than the m , the m is narrow in its anterior part are variable, but never present between the anteri- 2 1 or centroloph and the metaloph. and wide in its posterior part. The centrolophid is Maxillary P4 (Text-fig. 3/1) is rounded in the shortest of the seven ridges, and is the only one occlusal outline. The figured specimen is the small- that does not reach the lingual border. It is situated est of three P4’s. In addition to the four main in the center of the tooth. Mandibular m (Text-fig. 3/7) is almost square, ridges, there is an anterior centroloph of short or 2 medium length. In two out of three teeth there is a sometimes rectangular in occlusal outline. The posterior extra ridge between the metaloph and the number of ridges is equivalent to m1. There are posteroloph. The anteroloph protoloph and the some lingual connections between ridges: The anterior centroloph are connected buccally. anterolophid connects the metalophid in four out Likewise, there is a buccal connection of the met- of six specimens (two are fragmentary). In four aloph and the posteroloph. specimens the anterior extra ridge reaches this lin- Maxillary M1 (Text-fig. 3/2) is square in occlusal gual connection of anterolophid and metalophid. In three out of six m ’s the posterior extra ridge and outline. There is a very short extra ridge posterior 2 to the anteroloph, only in one specimen the extra the posterolophid connect lingually. Mandibular m (Text-fig. 3/8) is narrow posteri- ridge is of medium length. The posterior extra 3 orly, but similar to the m in its occlusal mor- ridge is generally longer. It connects the pos- 2 teroloph buccally in six out of eleven specimens. In phololgy. The centrolophid, although partly thin six M1’s there is a buccal connection of the pos- and short, is present in all four specimens. teroloph and metaloph. Maxillary M2 (Text-fig. 3/3) is slightly wider than Discussion the M1. Its postero-buccal edge is rounded. The The genus Glis ranges from the Oligocene G. dental pattern is similar to the M1 except for two guerbuezi Ünay, 1989 to the living G. glis (Linnaeus, characters: in two out of seven specimens there is a 1766). Glis shows a remarkable stability in tooth EOMYIDAE AND GLIRIDAE FROM RUDABÁNYA 157

Text-fig. 4 - Scatter diagrams of tooth measurements of Glis minor Kowalski, 1963 from Rudabánya (Hungary). L =length, W = width, P4 = upper premolar, M1-3 = upper molars, p4 = lower premolars, m1- 3 = lower molars.

pattern for more than 25 million years, i.e. from de Bruijn & Rümke, 1974 , G. tryolsi Daams, 1976 the Oligocene to modern times. Thus, species-dif- and G. minor complicatus de Bruijn, 1998. ferentiation was based in some cases on size-differ- Glis minor was originally described from the ences, rather than on significant morphological Pliocene localities Podlesice (MN14), Weze (MN15) features. Size and molar pattern set the Rudabánya and Rebielice (MN16) in Poland. However, some population apart from the Oligocene and Early Austrian Vallesian and Turolian occurrences Miocene species: G. apertus Mayer, 1979, G. gali- include: Richardhof-Golfplatz (MN9), Götzendorf topouli Van der Meulen & de Bruijn, 1982, G. major (NM9), Richardhof-Wald (MN10) and Kohfidisch 158 GUDRUN DAXNER-HÖCK

Text-fig. 5 - Scatter diagrams of tooth measurements of Myoglis cf. ucrainus Nesin & Kowalski, 1977 from Ru- dabánya (Hunga- ry). L =length, W = width, P4 = upper premolar, M1-3 = upper molars, d4/ p4 = lower premo- lars, m1-3 = lower molars.

(MN11). The Late Miocene (Vallesian) G. vallesiensis ical features: no extra ridge between anteroloph and is known by a few teeth only, from three localities, protoloph of M2; no centrolophid in m3; more or Ballestar, Can Petit and La Bastida in Spain. less square outline of m2 (Agusti, 1981). According to Agusti (1981), G. vallesiensis differs Glis from Rudabánya is intermediate in occlusal from G. minor and from G. sackdillingensis morphology and size between the smaller G. minor (Pleistocene) by its larger size, and from G. major by Kowalski, 1965 and the larger G. vallesiensis its smaller size only. G. vallesiensis is almost as large Agusti, 1981, but definitely resembles G. minor as the extant G. glis, but differs in minor morpholog- more. EOMYIDAE AND GLIRIDAE FROM RUDABÁNYA 159

Two further occurrences of middle sized Glis of lower molars are anterolophid, anterior extra have been reported from Belchatóv A in Poland ridge, mesolophid, metalophid and posterolophid. (MN9) and from Gritsiv in Ukraine (MN9) The valleys are V-shaped in cross section. The P4 (Kowalski, 1997; Nesin & Kowalski, 1997). Both and the M3 have three roots: one lingual and two populations come close to G. minor from buccal. The M1/2’s have four roots. The p has one 4 Rudabánya in morphology and size. They were root, the lower molars (m ) have three roots: two 1-3 identified as G. vallesiensis, and the smaller dimen- anterior and a wide one posteriorly. sions were interpreted as being geographical varia- Maxillary P4 (Text-fig. 3/9) is large and of vari- tions (Kowalski, 1997). Potential relations to G. able size and shape. There are five to six lophs. The minor were not discussed. protoloph, the metaloph and the posteroloph are lingually connected in five out of ten specimens. The anterior centroloph varies from absent to Myoglis Baudelot, 1965 almost long. Myoglis cf. ucrainicus Nesin & Kowalski, 1997 Maxillary M1 (Text-fig. 3/10) has a rectangular (Text-fig. 3/9-16, 5; Tab. 8) occlusal outline. The ridges are strongly oblique. In seven out of fifteen specimens the long main 1974 Pentaglis meini de Bruijn - Kretzoi et al.: p. 375 ridges, other than the anteroloph, connect in the postero-lingual corner of the crown. In eight out of Type locality fifteen specimens only the metaloph and the pos- Gritsiv (Ukraine), Vallesian (MN9). teroloph are connected, but the protoloph and the anteroloph end free. As a rule there is an extra Material ridge on either side of the centroloph. There is a 5 P4(°), 10 M1(°), 6 M2(°), 2 M3(°), 8 p (°), 6 m (°), 9 4 1 very small third extra ridge in the posterior valley m (°), 2 m (°), 5 P4(*), 5 M1(*), 6 M2(*), 2 M3(*), 5 p (*), 2 3 4 in a few specimens. 10 m (*), 5 m (*), 3 m (*). 2 1 2 3 Maxillary M (Text-fig. 3/11) is square in occlusal outline. The number and orientation of lophs is simi- Measurements lar to the M1, but the centroloph and the extra ridges See table 8 and Text-fig. 5. are generally longer. The three main ridges are always confluent in the postero-lingual corner. TABLE 8 - Measurements (in mm) of Myoglis cf. ucraini- Maxillary M3 (Text-fig. 3/12) is strongly narrow cus from Rudabánya posteriorly. All lophs are curved distally and partly confluent with the posteroloph. The number of G. minor min mean max n stdev ridges is highest in M3, there are seven to eight ridges of varying length. P4 L 1.25 1.28 1.30 3 0.02887 Mandibular p (Text-fig. 3/13) is triangular in 4 P4 W 1.35 1.42 1.50 3 0.07638 occlusal outline, and rounded. There are four M1 L 1.55 1.63 1.75 11 0.06431 ridges ending free on both sides of the crown. Only P4 L 1.50 1.63 1.90 10 0.12748 in three out of thirteen specimens the mesolophid P4 W 1.60 1.77 1.90 10 0.11068 and the posterolopid are connected lingually. If M1 L 1.85 1.93 2.05 15 0.06172 present, the centrolophid is short and low. M1 W 1.85 1.93 2.05 15 0.06172 Mandibular m (Text-fig. 3/14) is rectangular in 1 M2 L 1.70 1.83 2.00 12 0.08908 occlusal outline. The m is most variable in size. 1 M2 W 1.95 2.08 2.30 11 0.09318 The anterior extra ridge is high, but medium M3 L 1.75 1.78 1.80 4 0.02887 length. As a rule there are two low ridges the cen- M3 W 1.85 1.93 2.00 3 0.07638 trolophid and the posterior extra ridge. In six out of sixteen specimens there is a weak extra ridge p4 L 1.25 1.42 1.50 13 0.07742 posteriorly to the posterolophid. All the lophs end p4 W 1.15 1.33 1.50 13 0.09216 free on both sides of the tooth. m1 L 1.45 1.90 2.05 16 0.13292 Mandibular m (Text-fig. 3/15) is rectangular or 2 m1 W 1.60 1.72 1.85 16 0.07042 almost square in occlusal outline. The molar pat- m2 L 1.85 1.98 2.05 14 0.07003 tern is similar to m . 1 m2 W 1.90 2.01 2.10 14 0.06157 Mandibular m (Text-fig. 3/16) is similar to m , 3 2 m3 L 2.00 2.06 2.10 5 0.05477 but narrow posteriorly, and wide anteriorly. m3 W 1.90 2.00 2.15 5 0.09354 Discussion The dental pattern of Myoglis is characterized Description by flat crowns with four main ridges oblique to the The teeth are large and have a flat occlusal sur- longitudinal axis and by a varaiable number of face. The ridges are directed distally in the upper extra ridges. From the Early to the Late Miocene teeth and mesially in the lower teeth. In the upper (M. tryolsi, MN2 - M. houlezi, MN3 - M. antecedens, teeth there are four long ridges (anteroloph, pro- MN 3-5 - M. meini, MN 5-10? - M. ucrainicus, toloph, metaloph and posteroloph) and a well MN9-10) there is a trend towards increasing size developed anterior centroloph which may extend and towards a more complex dental pattern halfway across the crown. These five ridges are of (Aguilar, 1997; Daams, 1976; Werner, 1994; Mayr, equal elevation in upper and lower teeth. The extra 1979; Daams & de Bruijn, 1995; Nesin & Kowalski, ridges are lower than the main ridges and vary in 1997; Kowalski, 1979). length and number. The corresponding five ridges Myoglis meini ranges from the Middle Miocene 160 GUDRUN DAXNER-HÖCK

(MN5) to the Late Miocene (MN10): Hambach 6C TABLE 9 - Stratigraphic ranges of glirids and eomyids in Germany (Nemetschek & Mörs, 2003), from Rudabánya Neudorf/Spalte in Slowakia (Fejfar, 1989), Manchones in Spain (de Bruijn, 1966), Sansan in FOD LOD France (Baudelot, 1965), Anwil in Switzerland and Can Llobateres in Spain (Engesser, 1972), Castell Giridae: Glirulus lissiensis MN4 MN13 de Barbera (Aguilar et al., 1979), Comanesti in Paraglirulus werenfelsi MN5 MN10 Romania (Feru et al., 1980), Douvre and Soblay in Glis minor MN4 MN16 France (Mein, 1984) and other localities (see: Myoglis cf. ucrainicus MN9 MN10 Nemetschek & Mörs 2003, Tab.3). Muscardinus hispanicus MN9 MN10 Myoglis ucrainicus so far was only described Muscardinus cf. vallesiensis MN9 MN10 from the Late Miocene (MN9) of Gritsiv in Ukraine Eomyidae: Eomyops catalaunicus MN9 MN10 (Nesin & Kowalski, 1997). It differs from M. meini by higher numbers of roots in the upper and lower teeth, and by the tendency to lose two morphologi- cal characters: the lingual connection of protoloph Rudabánya can be biochronologically character- and metaloph in M , and the small loph posterior 1-2 ized as follows: only three glirid species are known to the posterolophid in lower molars (still present to range from the Early to the Late Miocene and in about half of the Gritsiv specimens). Myoglis the Pliocene, respectively (Paraglirulus werenfelsi, ucrainicus may have descended from M. meini. Glis minor and Glirulus lissiensis). The others The Rudabánya-specimens are largest, they (Myoglis cf. ucrainicus, Muscardinus hispanicus, cover varying morphological features of M. meini Muscardinus cf. vallesiensis) are limited to the and M. ucrainicus. Concerning root numbers they Vallesian (Table 9). However, in Central Europe the are identical with M. ucrainicus. However, the latter were replaced by Myomimus dehmi, Rudabánya-specimens and some other Vallesian Vasseuromys pannonicus and more advanced occurrences from Austria (Richardhof-Golfplatz, species of Muscardinus during the Turolian. MN9; Götzendorf, MN9; Richardhof-Wald, MN10; Schernham, MN10) are identified as M. cf. ucraini- cus. ACKNOWLEDGEMENTS

For financial, logistic and scientific support the following PALEOECOLOGY AND BIOSTRATIGRAPHY institutions and persons are gratefully acknowledged: M. Kretzoi, L. Kordos and R. Bernor provided me with fossils from Rudabánya. A. Englert and F. Topka screened the pre-washed Extant dormice inhabit the Palearctic region, samples in the laboratory of the Natural History e.g. from England to the Mediterranean, Northern Museum/Vienna and sorted the washing remains. The SEM- Africa, Asia Minor to Russian Asia. Only one genus, photos were taken by D. Gruber (Biozentrum/University Graphiurus, lives in Africa and a second one, Vienna), and the prints were made by A. Schumacher (Museum Glirulus, is known from Japan only. All these of Natural History/Vienna). E. Höck assisted with the computer- graphics. These investigations were granted by the National dormice live in trees and bushes, gardens and Geographic Society, by the LSB Leaky Foundation and by the rocks. Typical dormice are active at night. They are Austrian Science Fund: FWF-project P-15724-N06. squirrel-like in some of their habits. They are climbing animals, shelter in hollow trees, on the branches of trees or shrubs, among rocks and in REFERENCES deserted burrows of other animals, generally in a nest built of plant materials. The diet of extant AGUILAR J.-P. (1974): Les rongeurs du Miocène inférieur en Bas- dormice includes nuts, berries, seeds, soft fruits, Languedoc et les corrélations entre échelles stratigraphiques insects and occasionally the nestlings and eggs of marine et continentale. Géobios, 7 (4), 345-398. birds. Hibernation is known from all living AGUILAR J.-P., AGUSTI J. & GIBERT J. (1979): Rongeurs Miocenes dans le Valles-Penedes. 2 – Les Rongeurs de Castell de dormice, although Muscardinus remains active at Barbera. Palaeovertebrata, 9 (I), 17-31. temperatures above 16°C. For hibernation they use AGUSTI J. (1981): Glis vallesiensis n. sp., Nouveau Gliridae nests in hollow trees, beneath debris or under (Rodentia, Mammalia) du Neogene de Seu d`Urgell ground (Walker, 1964: 974-975). (Catalogne, Espagne). Géobios, 14 (4), 543-547. In that some dormice from Rudabánya belong BACHMAYER F. & WILSON R. W. (1983): Tertiary Gliridae to the living genera Glis, Muscardinus and Glirulus, (Dormice) of Austria. Ann. Naturhist. Mus. Wien, 85A, 129- we assume comparable paleoecological conditions 134. in Late Miocene times. Thus, the glirids from BAUDELOT S. (1965): Complément à l`étude de la faune des Rudabánya most probably lived in arboreal habi- rongeurs de Sansan: les Gliridés. Bull. Soc. Géol. France, 7, tats, and some were probably gliders. They inhabit- 758-764. ed forests surrounded by shrubs, like the eomyid, BERNOR, R.L., KORDOS, L., ROOK, L., AGUSTÍ , J., ANDREWS, P., Eomyops, and the flying squirrels did. Others ARMOUR-CHELU, M., BEGUN, D.R., CAMERON, D.W., DAMUTH, J., DAXNER-HÖCK, G., DE BONIS, L., FEJFAR, O., FESSAHA, N., might have inhabited open woodlands more dis- FORTELIUS, M., FRANZEN, J., GASPARIK, M., GENTRY, A., tant from swamps and water bodies. HEISSIG, K., HERNYAK, N., KAISER, T., KOUFOS, G.D., KROLOPP, According to Daams & de Bruijn (1995: 14, Fig. E., JÁNOSSY, D., LLENAS, M., MESZÁROS, L., MÜLLER, P., 2) in the Early Miocene the glirids were very abun- RENNE, P., RO_EK, Z., SEN, S., SCOTT, R., SZYNDLAR, Z., TOPÁL, dant all over Europe. The largest species-numbers GY., UNGAR, P.S., UTESCHER, T., VAN DAM J.A., WERDELIN & L., ZIEGLER, R. (2003): Recent advances on multidisciplinary were present during MN4-5. It gradually decreased research at Rudabánya, Late Miocene (MN9), Hungary: a to the end of the Miocene. The glirids from compendium. Palaeontographia Italica, 89 (2002), 3-36. EOMYIDAE AND GLIRIDAE FROM RUDABÁNYA 161

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