Original article

A new interpretation of the Miocene rodent faunas fromComanesti 1 and Taut (W-Romania)§

Une nouvelle interprétation des faunes de rongeurs de Comanesti 1 et de Taut(Roumanie de l’ouest)

János Hír a,*, Jérôme Prieto b,c, Emanoil Stiuca d

a Municipal Museum of Pásztó, 3060 Pásztó, Pf. 15. Hungaryb Senckenberg Center for Human Evolution and Palaeoecology (HEP), Eberhard-Karls University Tübingen, Institute for Geoscience,

Sigwartstrasse 10, 72076 Tübingen, Germanyc Department for Earth and Environmental Sciences, Ludwig Maximilians University Munich & Bavarian State Collections for Palaeontology and Geology,

Richard Wagner Strasse 10, 80333 Munich, Germanyd Emil Racovita Institute of speleology, Romanian Academy, Str. Frumoasa nr. 11. C.P. 220-12, 78114, Bucharest, Romania

Received 30 April 2010; accepted 14 January 2011

Available online 24 February 2011

Abstract

The rodent faunas from the Comanesti 1 and Taut localities (Western Romania) are revised in light of the latest taxonomical, biostratigraphicaland palaeoenvironmental information. The main systematic results show that the two cricetid rodents Megacricetodon crisiensis andDemocricetodon iazygum are invalid, whereas Democricetodon zarandicus is retained. The original assigment of the cricetodontini remainsfrom Taut to Hispanomys is emended, as a relationship to Byzantinia is more likely. While the geological evidence suggests that the localities arelate Middle Miocene (Upper Volhynian-Bessarabian, late Sarmatian sensu stricto) in age, the association of Myoglis ucrainicus with Muscardinushispanicus rather argues for an MN9 correlation for Taut. Indeed, uncertainties and discontinuities in the Central and East European mammalianbiostratigraphy render any conclusion about the correlation of the localities to the MN ‘‘zonation’’ problematic. The high diversity of squirrels (fivegenera), as well as the presence of a pliopithecoid alongside, glirid and eomyid rodents, suggest a forest environment at the time of accumulation ofthe Taut fauna, which is in agreement with the indication of humid climate provided by ectothermic vertebrates.# 2011 Published by Elsevier Masson SAS.

Keywords: Biostratigraphy; Central Paratethys; Rodents; Miocene; paleoenvironment

Résumé

Une révision des faunes de rongeurs des localités Comanesti 1 et Taut (Roumanie de l’ouest) est réalisée à la lumière des dernièresconnaissances taxonomiques, biostratigraphiques et paléoenvironmentales. Les principaux résultats systématiques montrent que les deux rongeurscricetidés Megacricetodon crisiensis et Democricetodon iazygum ne peuvent être considérés comme valides, alors que Democricetodon zarandicusest conservé. L’attribution initiale des Cricetodontini de Taut au genre Hispanomys est corrigée, une relation avec Byzantinia étant plus probable.Bien que les informations géologiques suggèrent un âge Miocène moyen tardif (Volhynien supérieur-Bessarabien, Sarmatien tardif sensu stricto),l’association Myoglis ucrainicus + Muscardinus hispanicus est plus compatible avec une corrélation à MN9 pour Taut. Du fait des discontinuités etdes incertitudes liées à la biostratigraphie d’Europe centrale, toute conclusion est rendue fragile en ce qui concerne la correlation des sites avec la« zonation » MN. La grande diversité des sciuridés (cinq genres) ainsi que la présence d’un pliopithécoïde aux côtés de gliridés et d’éomyidés,peuvent être liées à une couverture végétale dense à l’époque de l’accumulation de la faune de Taut, une proposition en accord avec les donnéespluviométriques dérivées des vertébrés ectothermes.# 2011 Publié par Elsevier Masson SAS.

Mots clés : Biostratigraphie ; Parathéthys centrale ; Rongeurs ; Miocène ; Paléoenvironement

Geobios 44 (2011) 215–223

§ Invited editor: Jordi Agustí.* Corresponding author.

E-mail address: hirjanos@gmail.com (J. Hír).

0016-6995/$ – see front matter # 2011 Published by Elsevier Masson SAS.doi:10.1016/j.geobios.2011.01.003

J. Hír et al. / Geobios 44 (2011) 215–223216

1. Introduction

Late Middle Miocene to earliest Late Miocene smallmammal localities have rarely been reported from CentralEurope, and thus the faunal evolution during this period of timeremains poorly understood in this area. During the 5thSymposium of the Romanian Palaeontologists in Bucharest(2005), the first author and Márton Venczel therefore took theopportunity to re-study the original fossil material from Tautand Comanesti 1 that is kept in the Speleological Institute of theRomanian Scientific Academy (with the kind permission of thedirectorship). It became clear already after a first overview, thata detailed and thorough revision was necessary. At the sametime, an elaboration of the herpetofauna was completed(Venczel and Stiuca, 2008; excluding chelonians). In this paper,we present a taxonomical revision of the rodent faunas, anddiscuss the consequences of this revision on biostratigraphicaland palaeoenvironmental interpretations.

2. Historical and geological settings

2.1. Historical background

The faunas from Comanesti 1 and Taut have an importantposition in the history of Middle Miocene micromammalianresearch Europe. Indeed, the original descriptions of fossilsfrom these two localities represent a starting point for theHungarian and Romanian research in this region (Feru et al.,1979, 1980). During the following three decades, intensive fieldwork was conducted, which provided large collections ofMiocene micromammals and small vertebrates in general, thatform the basis for the present work (Hír and Kókay, 2004, 2009,2010; Hír and Venczel, 2005; Hír, 2006; Venczel and Stiuca,2008).

2.2. The localities

2.2.1. Comanesti 1This vertebrate locality is located at the eastern margin of the

village Comanesti, on the right side of the rivulet Hasmas. Thesediments of the eleven meter-high outcrop crop out in a privateyard (GPS: N 468 30.568’, E 228 02.807; section figured byIstocescu, 1971 and Feru et al., 1980), and are extremelyenriched in molluscs. From layer 21, which containscontinental vertebrates, Feru et al. (1980) listed Cardiumvindobonense vindobonense, C. latisulcum latisulcum, C.gracilicostatum, Ervilia dissita podolica and Trochus pictus.This layer belongs to the top of the lower lithological complexof the Sarmatian in the Crisul Alb Basin (sensu Istocescu, 1971;Istocescu and Istocescu, 1974) and correlates to the lateVolhynian/earliest Bessarabian (Feru et al., 1980) or to theentire Ervilia Zone of the Central Paratethys, late Volhynian(Harzhauser and Piller, 2004).

The continental vertebrate fossils collected from the lensesof tufa were initially described by Feru et al. (1980), andsubsequently followed by a study of the cricetid rodents withthe description of two new species (Radulescu and Samson,

1988). These authors correlate the fossils to the MN8 (latestMiddle Miocene). More recently, Grigorescu and Kazár (2006)described the odontocetes from this locality.

2.2.2. TautThis vertebrate locality is located south of the small village

Taut, in the Migiesului Valley (Arad District, WesternRomania). The tufaceous green clay containing Helix andPlanorbis shells was originally reported by Istocescu (1971)and Istocescu and Istocescu (1974). From the microvertebratematerial originally described by Feru et al. (1979), the cricetidrodents were emended by Radulescu and Samson (1988), withthe description of Democricetodon zarandicus nov. sp.McNulty et al. (1999) reported the lower deciduous premolarof a small-sized Pliopithecoid primate (Crouzelidae gen. andsp. indet.) from the Taut fauna. The herpetofauna (excludingchelonians) has been described by Venczel and Stiuca (2008).During the last decade, Márton Venczel and János Hír tried torediscover the locality in the field, but have been unsuccessfulso far.

Taut belongs to the middle lithological complex (sensuIstocescu, 1971; Istocescu and Istocescu, 1974). Feru et al.(1980) noticed that this complex corresponds to a distallacustrine facies that was accumulated, at least in its upper part,synchronously with the deposition of the lower lithologicalcomplex. Based on the evolutionary stage of the micromam-malian fauna, these authors suggest a lower Bessarabiancorrelation for this locality.

3. Material and methods

The nomenclature of the dental elements follows: Sciuridae:Cuenca-Bescos (1988); Gliridae: Daams (1981); Eomyidae:Engesser (1999); Cricetidae: Mein and Freudenthal (1971); andRummel (1998) for the special terminology of Cricetodontiniand Eumyarion teeth. Measurements were taken through ameasuring ocular mounted on a MBS 9 stereomicroscope andare given in mm. The micrographs were captured with a CanonEos 400D digital camera equipped with a Canon MP-E65 mmmacro-objective and a Canon MR-14EX macro ring light.Retouches were made by the first author using Photoshop 2 andPhotoshop 6.0. Drawings were made using a Zeiss drawingmicroscope.

4. Systematic palaeontology

Full paleontological descriptions of all identified taxa, aswell as the corresponding references, are given in Appendix A.A synoptical overview of the taxon occurrences resulting fromthe systematic analysis is given in Table 1. In addition, Figs. 1–

4 present rodent specimens from the localities Comanesti 1 andTaut.

The main results of our study show that:

� because of the small sample size from Comanesti 1, thecricetid rodent Democricetodon iazygum is considered asnomen dubium;

Table 1Overview of the taxon occurrences and synonymies resulting from the systematic analysis of the rodents from Taut and Comanesti 1.

Feru et al. (1979) Feru et al. (1980) Radulescu and Samson (1988) This paper

COMANESTI 1Peridyromys gr. hamadryas Miodyromys aff. grycivensisMyoglis meiniHispanomys cf. lavocati cf. Byzantinia sp. or div. sp.Hispanomys cf. bijugatus cf. Byzantinia sp. or div. sp.Democricetodon gr. gaillardi Democricetodon iazygum nov. sp. Democricetodon cf. freisingensis

vel. cf. gaillardiMegacricetodon gr. similis Megacricetodon crisiensis nov. sp. Megacricetodon cf. minutus

TAUvSpermophilinus bredai Spermophilinus bredaiMiopetaurista sp. Miopetaurista sp.Forsithya sp.Blackia sp. Blackia miocaenicaLeptodontomys gr. catalaunicus Eomyops oppligeriEomuscardinus gr. sansaniensis Muscardinus hispanicusGlirid gen. and sp. indet. Glirulus lissiensisMyoglis meini Myoglis ucrainicusDemocricetodon gr. gaillardi Democricetodon gr. gaillardi-brevis Democricetodon zarandicus nov. sp. Democricetodon zarandicusEumyarion cf. bifidus Eumyarion sp.

Glirudinus sp.?Albanensia sp.Neopetes hoeckarum

J. Hír et al. / Geobios 44 (2011) 215–223 217

� the diagnosis of Democricetodon zarandicus is emended;� for similar reasons, we provisionaly assign the specimens

from Comanesti 1 to Megacricetodon cf. minutus, consider-ing Megacricetodon crisiensis as a junior synonym of theAustrian species;� the largest cricetid molars from Comanesti 1 are morpho-

logically similar to Byzantinia, but the lack of sufficientmaterial neither permits any confident assignment of thesample nor valuation of its taxonomical homogeneity;� the dormouse Miodyromys aff. grycivensis may have been

present in Comanesti 1;� Muscardinus hispanicus, Glirudinus sp., Myoglis ucrainicus

and Glirulus lissiensis as well as the eomyid Eomyopsoppligeri occur in Taut.

5. Discussion

5.1. Biostratigraphic implications

The taxonomic revision of the fossils from Comanesti 1 andTaut allows us to re-evaluate the biostratigraphic correlation ofthe localities.

5.1.1. Comanesti 1Because of the small number of specimens, the taxonomic

determinations, and thus the resulting biostratigraphic implica-tions are vague. Megacricetodon disappears during MN9 (e.g.,Kälin, 1999). The earliest occurrence of a form related toMegacricetodon minutus has been reported from Gratkorn ataround 12–12.5 My (Prieto et al., 2010; Gross et al., in press).The oldest Byzantinia specimens have been recorded for thelatest Middle Miocene of Anatolia (MN7+8; Ünay et al., 2003),

whereas a Cricetodon-Byzantinia transitional species also livedin SE-Europe at this time (localities not indicated; Rummel,1999). With regard to the mollusk fauna, the deposits of thelocality are assigned to the upper part of the Volhynian, thusclearly indicating a late Middle Miocene age for the fauna.

5.1.2. TautSquirrels are not normally useful for fine biostratigraphy, but

the relatively large-sized Spermophilinus molars from Taut arein agreement with data available for the late Middle Miocene to(?)earliest Late Miocene localities from Central Europe(Fig. 5). The cricetid rodents do not provide precise informationbecause of taxonomic uncertainty (Eumyarion), or limitedgeographic range (Democricetodon zarandicus). The presenceof Eomyops oppligeri might argue for an MN8 correlation, butthe species has also been reported from localities traditionallycorrelated to MN9 (early Late Miocene) such as Bełchatów A(Garapich, 2002) and Nebelbergweg (Kälin and Engesser,2001). Similarly, E. oppligeri co-occurs at the top of theFelsötárkány succession with Microtocricetus molassicus (Hírand Kókay, 2010). The importance of Microtocricetus as acharacteric taxon for the MN9 has been challenged by Prietoet al. (2010), and the presence of species related to the Collimyshiri-C. dobosi lineage in both Bełchatów A and Felsötárkány3/10 also probably argues for a latest Middle Miocenecorrelation of the two localities (Prieto, pers. opinion). Theglirid Muscardinus hispanicus first occurs in Spain, i.e. in theupper part of the zone G3 of the Calatayud-Daroca and Teruelbasins (MN8, Casanovas-Vilar et al., 2008). Myoglis ucrainicushas originally been described from Grytsiv, a locality thatbelongs to the MN9 because of the presence of hipparioninhorses, and has also been recorded for the Austrian LateMiocene (Daxner-Höck and Höck, 2009). On the other hand,

[()TD$FIG]

Fig. 1. Sciuroid molars from Taut. 1. Albanensia sp. d4 (Tt 0132, reversed). 2. Neopetes hoeckarum p4 (Tt 0144). 3. Albanensia sp. D4 (Tt 0132). 4. Miopetauristasp. M1-2 (Tt 0158). 5. Spermophilinus bredai M1-2 (Tt 0142/3, reversed). 6. S. bredai M3 (Tt 0142/6, reversed). 7. Blackia miocaenica M1-2 (Tt 0162/13). 8.Neopetes hoeckarum m1-2 (Tt 0162/19). 9. N. hoeckarum m3 (Tt 0147). 10. S. bredai m3 (Tt 0162/11, reversed). 11. S. bredai m1-2 (Tt 0162/18).

J. Hír et al. / Geobios 44 (2011) 215–223218

[()TD$FIG]

Fig. 2. Eomyid and glirid molars from Taut and Comanesti 1. 1. Eomyops oppligeri m2 (Tt 0118, reversed). 2. Glirudinus sp. m2 (Tt 0200/1, reversed). 3. Gliruluslissiensis M1-2 (Tt 0201/1). 4. Miodyromys aff. grycivensis M1-2 (Com1-0115). 5. Muscardinus hispanicus M1 (Tt 0203/1). 6. M. hispanicus M1 (Tt 0203/2). 7.Myoglis ucrainicus p4 (Tt 0164). 8. M. ucrainicus M1 (Tt 0164, occlusal view). 9. M. ucrainicus M1 (Tt 0164, radical view). 10. M. ucrainicus m1 (Tt 0164). 11. M.ucrainicus m2 (Tt 0164).

J. Hír et al. / Geobios 44 (2011) 215–223 219

the fissure filling sediments from Grytsiv contain Sarmatimac-tra vitaliana (Topachevsky and Scorik, 1992; Topachevskiet al., 1996; Rzebik-Kowalska and Topachevsky, 1997;Vangegeim et al., 2006), a characteristic bivalve from theLate Sarmatian s. str. in the Central Parathethys (Piller et al.,

2007). The presence of M. ucrainicus in Taut indicates that itwas already present in the latest Middle Miocene. The presenceof Glirudinus, which is absent from the Late Miocene ofAustria (Daxner-Höck and Höck, 2009), could possibly be usedto confirm the correlation, but its presence in Jujurieux 1 (MN9;

[()TD$FIG]

Fig. 3. cf. Byzantinia sp. or div. sp. molars from Comanesti 1. 1. m1 (Cm1-0132/1). 2. m1 (Cm1-0131/1 reversed, occlusal view). 3. m1 (Cm1-0131/1 reversed, labialview). 4. M2 (Cm1-0134/1 reversed, occlusal view). 5. M3 (Cm1-0135/1 reversed). 6. m3 (cm1-0133/1). 7. M2 (Cm1-0134/1 reversed, lingual view). 8. m3 (Cm1-0136/1 reversed).

J. Hír et al. / Geobios 44 (2011) 215–223220

[()TD$FIG]

Fig. 4. Democricetodon and Eumyarion molars from Taut. 1–14. D. zarandicus. 1. m1 (Tt 0149). 2. m1 (Tt 0152, reversed). 3. m1 (Tt 0155). 4. m1 (Tt 0148,reversed). 5. m1 (Tt 0154). 6. M1 (Tt 0175). 7. M1 (Tt 0176, reversed). 8. M1 (Tt 0178). 9. M1 (Tt 0188, reversed). 10. M2 (Tt 0173, reversed). 11. M3 (Tt 0185/1). 12.M3 (Tt 0183/1, reversed). 13. M3 (Tt 0185/2). 14. M3 (Tt 0183/2, reversed). 15, 16. Eumyarion sp. mediusi. 15. M2 (Tt 0193, reversed). 16. (Tt 0192, reversed).

J. Hír et al. / Geobios 44 (2011) 215–223 221

[()TD$FIG]

Fig. 5. Scatter diagram showing the size variation of the two first upper molars of Spermophilinus.

J. Hír et al. / Geobios 44 (2011) 215–223222

Aguilar et al., 2004) would argue against this. Based on thepreceding considerations, a correlation of Taut with both MN8and MN9 can be justified. However, this result is partially indisagreement with the chronostratigraphic position of thelocality (Middle Miocene), if a correlation to MN9 (LateMiocene) is proposed. On the other hand, the concept of theMN biozonation is based on three criteria (Mein, 1999):

� characterising species with short chronological extension ofwell established lineages;� large mammal ‘‘association’’ wich coexistence is reduced to

one unit;� apparition of taxa (first occurrence) at the generic level.

With regard to this definition, it has to be pointed out that ourcorrelation does not meet the requirements:

� for taphonomic reasons, we deal only with small mammals, inthis case rodents;� due to the lack of sufficient localities around the Middle to

Late Miocene transition in Central and Eastern Europe, thelineages are not well defined, and similarly, the firstoccurrences of taxa are based on an unconfident amount ofinformation.

For these reasons, it is not illogical to have rodent faunaswith MN9-like evolutionary stage in the Middle Miocenedeposits.

5.2. Environmental implications

The small rodent sample from Comanesti 1 is not sufficientfor environmental reconstructions, whereas the mammalassociation from Taut offers a deeper insight into the habitat

in which these animals lived. The sciurids qualitativelydominate the sample (five genera) with the presence of theflying squirrels, which are indicators of a dense canopy(Daxner-Höck and Höck, 2009). Based on the methodpresented by Böhme et al. (2006), Venczel and Stiuca (2008)calculated the mean annual precipitation (709 mm � 250–

280 mm) derived from the herpetofauna from Taut. Thiscalculation is indicative of a humid climate, with precipitationmore or less similar to present-day conditions (note that theauthors do not include chelonians in their analysis, therefore thevalues can be underestimated M. Böhme; pers. communica-tion). Böhme et al. (2008) estimated the volume of thepalaeoprecipitation during the Late Middle and Late Miocenefor Central and Eastern Europe. During the period 12.0–11.4My, palaeoprecipitation values were similar to modern ones. Incontrast, between 10.7–9.7 My (Early Pannonian) the rainfallexceeded present-day values by more than 200% (a so-called‘‘wash house climate’’). This means that the precipitationsignificantly increased around the Astaracian-Vallesian bound-ary. Clearly, the composition and the palaeoecologicalcharacter of the Taut assemblage resemble those seen in earlyPannonian faunas from the Carpathian Basin representingforested environments (e.g., Richardhof-Golfplatz, BorskySväty Jur, Götzendorf, Rudabánya) favorable for primates,as evidenced by the discovery of a Crouzelidae remain in theTaut deposits (Begun et al., 2006; McNulty et al., 1999).

Acknowledgements

The authors would like to express there sincere thanks toMárton Venczel (Oradea) for his kind help. Madelaine Böhme(Tübingen) and Peter Joniak (Bratislava) are thanked for theirvery useful comments on an earlier draft of the manuscript. Thisstudy was supported by the Deutsche Forschungsgemeinschaft

J. Hír et al. / Geobios 44 (2011) 215–223 223

grant BO1550/16-1 and the project no. T046719 of theHungarian Scientific Research Fund (OTKA).

Appendix A. Supplementary data

Supplementary text and data (Tables S1 to S13 and Figs. S1and S2) associated with this article can be found, in the onlineversion, at doi:10.1016/j.geobios.2011.01.003.

References

Begun, D.R., Nargolwalla, M.C., Hutchison, M.P., 2006. Primate diversity inthe Pannonian Basin: In situ evolution dispersal or both? Beiträge zurPaläontologie 30, 43–56.

Böhme, M., Ilg, A., Ossig, A., Küchenhoff, H., 2006. New method to estimatepaleoprecipitation using fossil amphibians and reptiles and the middle andlate Miocene precipitation gradients in Europe. Geology 34, 425–428.

Böhme, M., Ilg, A., Winklhofer, M., 2008. Late Miocene ‘‘washouse’’ climatein Europe. Earth and Planetary Science Letters 275, 393–401.

Casanovas-Vilar, I., Alba, D.M., Moya-Sola, S., Galindo, J., Cabrera, L.,Garces, M., Fùrio, M., Robles, J.M., Köhler, M., Angelone, M., 2008.Biochronological, taphonomical and paleoenvironmental background of thefossil great ape Pierolapithecus catalaunicus (Primate hominidae). Journalof Human Evolution 55, 589–603.

Cuenca-Bescos, G, 1988. Revisión de los Sciuridae del Aragoniense y delRambliense en la fosa de Calatayud – Montalban. Scripta Geologica 87, 2–

115.Daams, R., 1981. The dental pattern of the dormice Dryomys, Myomimus,

Microdyromys and Peridyromys. Utrecht Micropaleontological Bulletins.Special Publication 3, 1–115.

Daxner-Höck, G., Höck, E., 2009. New data on Eomyidae and Gliridae(Rodentia, Mammalia) from the Late Miocene of Austria. Annalen desNaturhistorisches Museums Wien 111A, 375–444.

Engesser, B., 1999. Family Eomyidae. In: Rössner, F.G., Heissig, F.K. (Eds.),The Miocene land Mammals of Europe. Verlag Dr. Friedrich Pfeil,München, pp. 319–335.

Feru, M., Radulescu, C., Samson, P., 1979. La faune de Micromammifères duMiocène de Taut (département d’Arad). Travaux de l’Institut de Spéologie« Emil Racovitza » 18, 185–190.

Feru, M., Radulescu, C., Samson, P., 1980. La faune de Micromammifères duMiocène de Comanesti (département d’Arad). Travaux de l’Institut deSpéologie « Emil Racovitza » 19, 171–190.

Garapich, A., 2002. An overview of Miocene rodents from Bełchatów (Poland).Folia Zoologica 51 (Suppl. 1), 59–66.

Grigorescu, D., Kazár, E., 2006. A new Middle Miocene odontocete (Mam-malia, Cetacea) locality and the Sarmatian Marine Mammal Event in theCentral Paratethys. Oryctos 6, 53–67.

Gross, M., Böhme, M., Prieto, J., in press. Gratkorn – a benchmark locality forthe continental Sarmatian s.str. of the Central Paratethys. InternationalJournal of Earth Sciences (doi:10.1007/s00531-010-0615-1).

Harzhauser, M., Piller, W., 2004. Integrated stratigraphy of the Sarmatian(Upper Middle Miocene) in the western Central Paratethys. Stratigraphy1, 1–22.

Hír, J., 2006. Late Astaracian (Late Sarmatian) Lagomorphs and Rodents fromFelsotárkány – Felnémet (Northern Hungary). Beiträge zur Paläontologie30, 155–173.

Hír, J., Kókay, J., 2004. Middle Miocene molluscs and rodents from Mátraszo-

los (Mátra Mountains, Hungary). Fragmenta Palaeontologica Hungarica 22,83–97.

Hír, J., Kókay, J., 2009. Middle Miocene Molluscs, Lagomorphs and Rodents ofFelsotárkány 1 and Felsotárkány 2. Fragmenta Palaeontologica Hungarica27, 81–89.

Hír, J., Venczel, M., 2005. New Middle Miocene vertebrate localities fromSubpiatra (Bihor County, Romania). Acta Palaeontologica Romaniae 5,211–221.

Hír, J., Kókay, J., 2010. A systematic study of the middle-late Miocene rodentsand lagomorphs (Mammalia) of Felsotárkány 3/8 and 3/10 (NorthernHungary). Geodiversitas 32, 307–329.

Istocescu, D., 1971. Geologisches Studium der West Zone des Crisul Alb-Beckens und ihre Umrahmung im Codru – und Highis – Gebirge. Studiitechnice si Economice, Institutul de Geologie si Geofizica, Bucuresti. SeriesJ8 1–201.

Istocescu, D., Istocescu, F., 1974. Geological considerations on the Neogenedeposits from Crisurilor Basin. Studii si Cercetari Geologice, Geofizice,Geografice, seria Geologie Bucuresti 19, 115–128.

Kälin, D., 1999. Tribe Cricetini. In: Rössner, G., Heissig, K. (Eds.), TheMiocene land Mammals of Europe. Verlag Dr. Friedrich Pfeil, München,pp. 373–387.

Kälin, D., Engesser, B., 2001. Die jungmiozäne Säugetierfauna vom Nebel-bergweg bei Nunningen (Kanton Solothurn, Schweiz). SchweizerischePaläontologische Abhandlungen 121, 1–61.

McNulty, K., Radulescu, C., Samson, P., Feru, M., Delson, E., 1999. Morphol-ogy and identification of a previously undescribed catarrhine tooth from themiddle Miocene of Romania. American Journal of Physical Anthropology28, 199.

Mein, P., 1999. European Miocene Mammal Biochronology. In: Rössner,G.E., Heissig, K. (Eds.), The Miocene land mammals of Europe. VerlagDr. Pfeil, Munich, pp. 25–38.

Mein, P., Freudenthal, M., 1971. Une nouvelle classification des Cricetidae(Mammalia, Rodentia) du Tertiaire de l’Europe. Scripta Geologica 2, 1–37.

Piller, W.E., Harzhauser, M., Mandic, O., 2007. The Middle and UpperOttnangian of the Simsee area (SE Germany): Miocene Central Para-tethys stratigraphy-current status and future directions. Stratigraphy 4,151–168.

Prieto, J., Böhme, M., Gross, M., 2010. The cricetid rodent fauna from Gratkorn(Austria, Styria): a benchmark locality for the continental Sarmatian s. str.(late Middle Miocene) in the central Paratehys. Geologica Carpathica 61,419–436.

Radulescu, C., Samson, P., 1988. Les Cricétides (Rodentia, Mammalia) duMiocène (Astarcien Supérieur) du Roumanie. Travaux de l’Institut deSpéologie « Emil Racovitza » 37, 67–78.

Rummel, M., 1998. Die Cricetiden aus dem Mittel- und Obermiozän der Türkeiunter besonderer Berückschtigung der Sickenberg/Tobien’schen Aufsamm-lung (1968–1970). Documenta naturae 123, 1–300.

Rummel, M., 1999. Tribe Cricetodontini. In: Rössner, G., Heissig, K. (Eds.),The Miocene land Mammals of Europe. Verlag Dr. Friedrich Pfeil,München, pp. 359–364.

Rzebik-Kowalska, B., Topachevsky, V.O., 1997. Insectivora (Mammalia) fromthe Miocene of Grytsiv in Ukraine. 1. Heterosoricidae Viret and Zapfe,1951. Acta Zoologica Cracoviensia 40, 237–247.

Topachevski, V.A., Nesin, V.A., Topachevsky, V., Semenov, Y.A., 1996. Theoldest locality of the Middle Sarmatian microtheriofauna (Insectivora,Lagomorpha, Rodentia) in Eastern Europe. Dopovidi NAN Ukrainy 2,107–110.

Topachevsky, V.A., Scorik, A.F., 1992. Neogene and Pleistocene primitivecricetoids of Southeastern Europe. Naukova Dumka Press, Kiev, pp. 1–242.

Ünay, E., Bruijn, H., de, Saraç, G., 2003. A preliminary zonation of thecontinental Neogene of Anatolia based on rodents. In: Reumer, J.W.F.,Wessels, W. (Eds.), Distribution and migration of Tertiary mammals inEurasia, 10. A volume in honour of Hans de Bruijn, DEINSEA, pp. 539–

547.Vangegeim, E.A., Lungu, A.N., Tesakov, A.S., 2006. Age of the Vallesian lower

boundary (Continental Miocene of Europe). Stratigraphy and GeologicalCorrelation 14, 655–667.

Venczel, M., Stiuca, E., 2008. Late middle Miocene amphibians and squamatereptiles from Taut, Romania. Geodiversitas 30, 731–763.