Middle Miocene (Late Badenian) microvertebrates from Hidas, SW Hungary

Main Article Content

János Hír
Márton Venczel
Krisztina Sebe

Abstract

Up to the present, no terrestrial vertebrate fauna has been published from the pre-Pannonian Miocene of SW Hungary. In 2022 a microvertebrate assemblage was unearthed from a lime mud bed of the Middle Miocene Hidas Formation, in an abandoned coal mining field close to Hidas in the Mecsek Mts. The herpetofauna and the rodent material are described here. Fossil findings point to the Late Badenian MN 7+8 Zone, which, together with the earlier results based on the marine mollusc fauna, narrows the age of the unit to ~13.5–13.3 Ma. The amphibians and reptiles are aquatic, semiaquatic or periaquatic forms. Sedimentary features and the accompanying freshwater gastropod fauna are indicative of a shallow pond or a paludal depositional  environment. Crocodylian finds reported earlier from the Hidas Formation indicate a subtropical climate, just  before the end of the Miocene warm period in Central Europe. Among the rodents, glirids and flying squirrels as well as Democricetodon and Megacricetodon indicate the presence of humid arboreal vegetation around the  site. The rodent taxa are well known from the Middle Miocene faunas of northern Hungary, western Romania  and from the Upper Freshwater Molasse of southern Germany and Switzerland. The rodent material does not  show characteristics of an insular fauna, e.g. gigantism or endemism. Consequently, although the coeval  palaeogeography of the region has been described as an archipelago in the Central Paratethys, with the Mecsek  Mts. being one of the islands, the area must have had ecological connections towards the northern and eastern parts of the Pannonian Basin, and the marine areas within the archipelago did not form a barrier  against the distribution of microvertebrates. The corridor could have been located towards the NE from the Mecsek Mts., across the elevated basement blocks of central Hungary. 

Downloads

Download data is not yet available.

Article Details

Section
Original Scientific Papers

References

AGUILAR, J. (1982): Contributions à l’étude des micromammifères du gisement miocène supérieur de Montredon (Hérault-2). Les Rongeurs.− Palaeovertebrata, 12, 75–140.

BÁLDI, K., BENKOVICS, L. & SZTANÓ, O. (2002): Badenian (Middle Miocene) basin development in SW Hungary: subsidence history based on quantitative paleobathymetry of foraminifera.‒ International Journal of Earth Sciences, 91, 490–504. doi: 10.1007/s005310100226

BARANYAI, D. (2010): Hidasi középső-miocén otolithok összehasonlító vizsgálata [A comparative study of Middle Miocene otoliths from Hidas – in Hungarian].‒ Hungarian Student Scientific Competition paper, Eger, 42 p.

BAUDELOT, S. (1965): Complément à l’étude de la faune des rongeurs de Sansan: les Gliridés.‒ Bull. Soc. géol. France, 7, 758–764. doi: 10.2113/gssgfbull.S7-VII.5.758

BAUDELOT, S. (1972): Etude des Chiroptéres, Insectivores et Rongeurs du Miocéne de Sansan (Gers).‒ Thése, Toulouse, 496, 364 p.

BITON, R., GEFFEN, E., VENCES, M., COHEN, O., BAILON, S., RABINOVICH, R., MALKA, Y., ORON, T., BOISTEL, R., BRUMFELD, V. & GAFNY, S. (2013): The rediscovered Hula painted frog is a living fossil.‒ Nature Communications 4, e1959. doi: 10.1038/ncomms2959

BÖHME, M. (2003): The Miocene Climatic Optimum: evidence from ectothermic vertebrates of Central Europe.‒ Palaeogeography, Palaeoclimatology, Palaeoecology, 195, 389–401. doi: 10.1016/S0031-0182(03)00367-5

BOHNNÉ HAVAS, M. (1973): A keleti Mecsek torton Mollusca faunája [Tortonian mollusc fauna of the Mecsek Mountains].‒ Annals of the Geological Institute of Hungary, 53/4, 218 p.

CSEPREGHYNÉ-MEZNERICS, I. (1950): A hidasi (Baranya m.) tortonai fauna [The Tortonian fauna of Hidas (Baranya County) – in Hungarian].‒ Annals of the Geological Institute of Hungary, 39/2, 3–114.

CUENCA-BESCOS, G. (1988): Revision de los Sciuridae del Aragoniense y del Rambliense en la fossa de Calatayud-Montalban.− Scripta Geologica, 87, 1–115.

DAAMS, R. (1981): The dental pattern of the dormice Dryomys, Myomimus, Microdyromys and Peridyromys.‒ Utrecht Micropaleontological Bulletins, Special Publications, 3, 115 p.

DAAMS, R. (1985): Glirinae (Gliridae, Rodentia) from the type area of the Aragonian and adjacent areas (provinces of Teruel and Zaragoza, Spain).‒ Scripta Geologica, 77, 1–20.

DAMS, R., FREUDENTHAL, M. & VAN DER MEULEN, A. (1988): Ecostratigraphy of micromammal faunas from the Neogene of Spain.‒ Scripta Geologica, Special Issue, 1, 287–302.

DAXNER-HÖCK, G. (2004): Flying Squirrels (Pteromyinae, Mammalia) from the Upper Miocene of Austria.‒ Annalen des Naturhistorischen Museums in Wien, Serie A, 106, 387–423.

DAXNER-HÖCK, G. (2010): Sciuridae, Gliridae and Eomyidae (Rodentia, Mammalia) from the Middle Miocene of St. Stefan in the Gratkorn Basin (Styria, Austria).‒ Annalen des Naturhistorischen Museums in Wien, 112A, 507–536.

DAXNER-HÖCK, G. & HÖCK, E. (2015): Catalogus fossilium Austriae. Band 4: Rodentia neogenica.‒ Verlag der Österreichischen Akademie der Wissenschaften, 158 p. doi: 10.2307/j.ctt1vw0qvk

DE BRUIJN, H. (1999): Superfamily Sciuroidea.‒ In: Rössner, G. & Heissig, K. (eds.): The Miocene Land Mammals of Europe, Verlag Dr. Friedrich Pfeil, München, 271–280 p.

DE BRUIJN H., MEULEN VAN DER, A. & KATSIKATSOS, G. (1980): The mammals from the lower Miocene of Aliveri (Island of Evia, Greece). 1. The Sciuridae.‒ In: Proceedings of the Koninklijke Nederlandse Akademie van Wetenschappen, Ser. B, 83, 241–261.

DE BRUIJN, H., VAN DEN HOEK OSTENDE, L., KRISTKOIZ-BOON, E., RUMMEL, M. THEOCHAROPOULOS, C. & ÜNAY, E. (2003): Rodents, lagomorphs and insectivores, from the middle Miocene hominoid locality of Çandir (Turkey).‒ Courier Forschungs-Institut Senckenberg, 240, 51–87.

ENGESSER, B., MATTER, A. & WEIDMANN, M. (1981): Stratigraphie und Säugetierfaunen des mittleren Miozäns von Vermes (Kt. Jura).‒ Eclogae geologicae Helvetiae, 74/3, 893–952.

ESTES, R. (1981): Encyclopedia of Paleoherpetology, Gymnophiona, Caudata.‒ In: WELLNHOFER, P. (ed.), Handbuch der Paläoherpetologie (Encyclopedia of Paleoherpetology), Part 2. Gustav Fischer Verlag, Stuttgart and New York, 115 p.

EVANS, S.E. & MILNER, A.R. (1993): Frogs ans salamanders from the Upper Jurassic Morrison Formation (Quarry nine, Como Bluff) of North America.‒ Journal of Vertebrate Paleontology, 13, 24–30.

FAHLBUSCH, V. (1964): Die Cricetiden (Mamm.) der Oberen Süsswasser-Molasse Bayerns.− Bayerische Akademie der Wissenschaften, matematisch-naturwissenschaftliche Klasse, Abhandlungen, Neue Folge, 118, 1–136.

FEJFAR, O., HEINRICH, W. & LINDSAY, E. (1997): Updating the European Neogene rodent biochronology.‒ In: AGUILAR J., LEGENDRE S. & MICHAUX J. (eds.): Actes de Congrès BiochroM’97, Mémoires et Travaux École Pratique des Hautes Études de l’Institut de Montpellier, 21, 563–565.

GEORGALIS, G. L., VILLA, A., IVANOV, M., VASILYAN, D. & DELFINO, M. (2019): Fossil amphibians and reptiles from the Neogene locality of Maramena (Greece), the most diverse European herpetofauna at the Miocene/Pliocene transition boundary.‒ Palaeontologia Electronica, 22.3.68, 1–99. doi: 10.26879/908

HÁMOR, G. (2001): Miocene palaeogeography of the Carpathian Basin. Explanatory notes to the palaeogeographic maps of the Carpathian Basin 1: 3 000.000.‒ Hungarian Geological Institute, 71 p.

HEISSIG, K. (1995): Die Entwicklung der grossen Democricetodon -arten und die gattung Collimys (Cricetidae, Mamalia) im späten Mittelmiozän.‒ Mitteilungen der Bayerischen Staatsasammlung für Paläontologie und historische Geologie, 35, 87–108.

HEISSIG, K. (1997): Mammal faunas intermediate between the reference faunas of MN4 and MN6 from the Upper Freshwater Molasse of Bavaria.− Actes du Congrès BiochroM’ 97, Mém. Trav. E.P.H.E. Inst. Montpellier, 21, 537–546.

HERRE, W. (1955): Die Fauna der miozänen Spaltenfüllung von Neudorf a.d. March (SCR).‒ Sitzber. Akad. Wiss. Math. Naturwiss. Kl., Abt A, 164, 783–803.

HÍR, J. (2001): New Middle Miocene rodent faunas from Northern Hungary.− Lynx (Praha) n. s., 32, 107–122.

HÍR, J. (2019): The Albanensia finds from Hungary and Romania.‒ Fragmenta Palaeontologica Hungarica, 36, 1–24. doi:10.17111/FragmPalHung.2019.36.1

HÍR, J. (2020): The Rodent faunas of the middle Miocene palaeovertebrate localities in the Pannonian Basin (Mammalia, Rodentia).‒ Manuscript, thesis, Municipal Museum of Pásztó, 1–285.

HÍR, J. & KÓKAY, J. (2004): Middle Miocene molluscs and rodents from Mátraszőlős (Mátra Mountains, Hungary).− Fragmenta Palaeontologica Hungarica, 22, 83–97.

HÍR, J. & KÓKAY, J. (2011): Late Badenian (MN 7/8) molluscs and rodents from Mátraszőlős 3 (Northern Hungary).− Fragmenta Palaeontologica Hungarica, 29, 69–78.

HÍR, J., CODREA, V. & PRIETO, J. (2019): Two new early Sarmatian s.str. (latest Middle Miocene) rodent faunas from the Carpathian Basin.– Palaeobiodiversity and Palaeoenvironments, 99/3, 527–543. doi:10.1007/s12549-019-00399-y PBPE-D-19-00003R1

HÍR, J. & VENCZEL, M. (2005): New Middle Miocene vertebrate localities from Subpiatră (Bihor District, Romania).‒ Acta Palaeontologica Romaniae, 5, 211–221.

HÍR, J. & VENCZEL, M. (2018): A preliminary report on the first results of the reexcavation of the middle Miocene palaeovertebrate locality Szentendre, Cseresznyésárok (Hungary, Pest County).‒ Nymphaea, Folia Naturae Bihariae, 45, 35–80.

HÍR, J., VENCZEL, M., CODREA, V., ANGELONE, CH., VAN DEN HOEK OSTENDE, L., KIRSCHER, U. & PRIETO, J. (2016): Badenian and Sarmatian s. str. from Carpathian area: Overview and ongoing research on Hungarian and Romanian small vertebrate evolution.‒ Comptes Rendus Palevol., 15, 863–875. doi:10.1016/j.crpv.2016.08.001

HÍR, J., VENCZEL M., CODREA V., RÖSSNER G., ANGELONE CH., VAN DEN HOEK OSTENDE, L., ROSINA, V., KIRSCHER, U. & PRIETO, J. (2017): Badenian and Sarmatian s.str. from the Carpathian area: Taxonomic notes concerning the Hungarian and Romanian small vertebrates and report on the Ruminants from the primate bearing Felsőtárkány Basin.− Comptes Rendus Palevol., 16, 312–332. doi: 10.1016/j.crpv.2016.11.006

HORVÁTH, F., BADA, G., SZAFIÁN, P., TARI, G., ÁDÁM, A. & CLOETHING, S. (2006): Formation and deformation of the Pannonian basin: Constraints from observational data.– In: GEE, D.G. & STEPHENSON, R.A. (eds.): European Lithosphere Dynamics, Geological Society, London, Memoirs 32, 191–206. doi: 10.1144/GSL.MEM.2006.032.01.11

HOSSINI, S. (1993): A new species of Latonia (Anura, Discoglossidae) from the Lower Miocene of France.‒ Amphibia-Reptilia, 14, 237–245. doi: 10.1163/156853893X00435

IVANOV, M. (2008): Early Miocene Amphibians (Caudata, Salientia) from the Mokrá – Western Quarry (Czech Republic) with comments on the evolution of Early Miocene amphibian assemblages in Central Europe.‒ Geobios, 41, 465–492. doi: 10.1016/j.geobios.2007.11.004

JONIAK, P. (2016): Upper Miocene rodents from Pezinok in the Danube Basin, Slovakia.− Acta Geologica Slovaca, 8/1, 2–17.

KÄLIN, D. (1997): Eomyops hebeiseni n. sp., a new large Eomyidae (Rodentia, Mammalia) of the Upper Freshwater Molasse of Switzerland.− Eclogae Geologicae Helvetiae, 90/3, 629–637.

KÄLIN, D. & KEMPF, O. (2009): High-resolution stratigraphy from the continental record of the Middle Miocene Northern Alpine Foreland Basin of Switzerland.− Neues Jahrbuch für Geologie und Paläontologie, 254/1–2, 177–235. doi: 10.1127/0077-7749/2009/0010

KORDOS, L. (1985): A magyarországi eggenburgi – szarmata képződmények szárazföldi gerinces maradványai, biozonációja és rétegtani korrelációja [Terrestrial vertebrate remains from the Eggenburgian to Sarmatian of Hungary: Biozonation and stratigraphic correlation].‒ Annual Report of the Geological Institute of Hungary on 1983, 157–166.

KORDOS, L. & SOLT, P. (1984): A magyarországi miocén tengeri gerinces faunaszintek vázlata (An outline of Hungary’s Miocene marine vertebrate faunal horizons).‒ Annual Report of the Geological Institute of Hungary on 1982, 347–353.

KOVÁČ, M., ANDREYEVA-GRIGOROVICH, A., BAJRAKTAREVIĆ, Z., BRZOBOHATÝ, R., FILIPESCU, S., FODOR, L., HARZHAUSER, M., NAGYMAROSY, A., OSZCZYPKO, N., PAVELIĆ, D., RÖGL, F., SAFTIĆ, B., SLIVA, L. & STUDENCKA, B. (2007): Badenian evolution of the Central Paratethys sea: paleogeography, climate and eustatic sea-level changes.‒ Geologica Carpathica, 58/6, 579–606.

KOVÁČ, M., HUDAČKOVÁ, N., HALÁSOVÁ, E., KOVÁČOVÁ, M., HOLCOVÁ, K., OSZCZYPKO-CLOWES, M., BÁLDI, K., LESS, GY., NAGYMAROSY, A., RUMAN, A., KLUČIAR, T. & JAMRICH, M. (2017): The Central Paratethys Palaeoceanography: a water circulation model based on microfossil proxies, climate, and changes of depositional environment.‒ Acta Geologica Slovaca, 9/2, 75–114.

KOVÁČ, M., HALÁSOVÁ, E., HUDÁČKOVÁ, N., HOLCOVÁ, K., HYŽNÝ, M., JAMRICH, M. & RUMAN, A. (2018): Towards better correlation of the Central Paratethys regional time scale with the standard geological time scale of the Miocene Epoch.– Geologica Carpathica, 69/3, 283–300. doi: 10.1515/geoca-2018-0017

KRETZOI, M. & FEJFAR, O. (2005): Sciurids and Cricetids (Mammalia, Rodentia) from Rudabánya.− Palaeontographia Italica, 90, 113–148.

KRIJGSMAN, W., LANGEREIS, C., DAAMS, R. & MEULEN VAN DER, A. (1994): Magnetostratigraphic dating of middler Miocene climate change in the continental deposits of the Aragonian type area in the Calatayud-Teruel basin (Central Spain).‒ Earth and Planetary Science Letters, 128, 513–526.

KRIJGSMAN, W., GARCÉS, M., LANGEREIS, C., DAAMS, R., MEULEN VAN DER, A., AGUSTÍ, J. & CABRERA, L. (1996): A new chronology for the middle to late Miocene continental record in Spain.‒ Earth and Planetary Science Letters, 142, 367–380. 10.1016/0012-821X(96)00109-4

MARIDET, O. (2003): Révision du genre Democricetodon (Mammalia, Rodentia, Cricetinae) et dynamique des faunes de rongeurs du Néogène d’Europe occidentale: évolution, paléobiodiversité et paléobiogéographie.‒ Thèse Université Claude Bernard, 252 p.

MARIDET, O. & SEN, S. (2012): Les Cricetidae (Rodentia) de Sansan.– In: PEIGNÉ S. & SEN S. (eds.): Mammiféres de Sansan.− Muséum national d’Histoire naturelle, 203, 26–95.

MARKOVIĆ, Z. & MILIVOJEVIĆ, M. (2010): The Neogene small mamals of Serbia –collection methods and results.‒ Bulletin of the Natural History Museum in Beograd, 3, 105–114.

MARKWICK, P.J. (1998): Fossil crocodilians as indicators of Late Cretaceous and Cenozoic climates: implications for using palaeontological data in reconstructing palaeoclimate.‒ Palaeogeography, Palaeoclimatology, Palaeoecology, 137, 205–271. doi: 10.1016/S0031-0182(97)00108-9

MCKENA, M. & BELL, S. (1997): Classification of mammals: above the species level.− Columbia University Press, New York, 1–631 p.

MEIN, P. & FREUDENTHAL, M. (1971): Les Cricetidae (Mammalia, Rodentia) du Néogène Moyen de Vieux- Collonges. Partie 1: Le genre Cricetodon Lartet, 1851.‒ Scripta Geologica, 5, 1–51.

MIKLAS, P. (2002): Die Amphibienfauna (Amphibia: Caudata, Anura) der obermiozänen Fundstelle Götzendorf an der Leitha (südliches Wiener Becken, Niederösterreich). ‒Annalen des Naturhistorischen Museums in Wien, 103 (A), 161–211.

NAGYMAROSY, A. & HÁMOR, G. (2012): Genesis and Evolution of the Pannonian Basin.- In: HAAS J. (ed.) 2012: Geology of Hungary. Springer, Heidelberg, 149–198.

PRIETO, J. (2007): Kleinsäuger-Biostratigraphie und Paläoökologie des höheren Mittelmiozäns (MN8) Bayerns: Spaltenfüllungen der Frankischen Alb und Lokalitäten der Oberen Süβwassermolasse im Vergleich.− Dissertation zur Erlangung des Doktorgrades (Dr. rer. nat.) an der Fakultät für Geowissenschaften der Ludwig-Maximilians-
Universität München, 1–213 p.

PRIETO, J. (2012): Comments on the morphologic and metric variability in the cricetid rodent Deperetomys hagni (Fahlbusch, 1964) from the Middle Miocene of South Germany.‒ Zitteliana, A52, 71–77.

PRIETO, J. & RUMMEL, M. (2016): Some considerations on small mammal evolution in Southern Germany, with emphasis on late Burdigalian-Earliest Tortonian (Miocene) cricetid rodents.− Comptes Rendus Palevol, 15, 837–854. doi: 10.1016/j.crpv.2016.08.002

RAGE, J.-C. & HOSSINI, S. (2000): Les Amphibiens du Miocène moyen de Sansan.– In : GINSBURG, L. (ed.): La faune miocène de Sansan et son environnement. Mémoires du Muséum national d’histoire naturelle, 183, 177–217.

ROČEK, Z. (1994): Taxonomy and distribution of Tertiary discoglossids (Anura) of the genus Latonia v. Meyer, 1843.‒ Geobios, 27, 717–751. doi: 10.1016/S0016-6995(94)80058-8

ROČEK, Z., RAGE, J.C. & VENCZEL, M. (2021): Fossil frogs of the genus Palaeobatrachus (Amphibia: Anura).‒ Abhandlungen der Senckenberg Gesellschaft für Naturforschung, 575, 1–151.

SANCHÍZ, B. (1998a): Vertebrates from the Early Miocene lignite deposits of the opencast mine Oberdorf (Western Styrian Basin, Austria): 2. Amphibia.‒ Annalen Des Naturhistorischen Museums in Wien 99A, 13–29.

SANCHÍZ, B. (1998b): Salientia.– In: WELLNHOFER, P. (ed.): Handbuch der Paläoherpetologie. (Encyclopedia of Paleoherpetology), Part 4. Friedrich Pfeil, Munich, 1–275 p.

SEBE, K., SELMECZI, I., SZUROMI-KORECZ, A., HABLY, L., KOVÁCS, Á. & BENKÓ, ZS. (2019): Miocene syn-rift lacustrine sediments in the Mecsek Mts. (SW Hungary).‒ Swiss Journal of Geosciences, 112, 83–100. doi: 10.1007/s00015-018-0336-1

SELMECZI, I., KÓKAY, J., HÁMOR, G., VETŐ, I. (2023): Hidasi Formáció [Hidas Fm.].‒ In: BABINSZKI, E., PIROS, O., CSILLAG, G., FODOR, L., GYALOG, L., KERCSMÁR, ZS., LESS, GY., LUKÁCS, R., SEBE, K., SELMECZI, I., SZEPESI, J. & SZTANÓ, O. (eds.): Magyarország litosztratigráfiai egységeinek leírása II. Kainozoos képződmények [Cenozoic Lithostratigraphic units of Hungary II. Cenozoic formations – in Hungarian]. Supervising Authority of Regulated Activities, Budapest, 92–93.

SEN, S. (1997): Magnetostratigraphic Calibration of the European Neogene Mammal Chronology.− Palaeogeography, Palaeoclimatology, Palaeoecology, 133/4, 181–204. doi: 10.1016/S0031-0182(97)00079-5

STEININGER, F. (1999): Chronostratigraphy, Geochronology and Biochronology of the Miocene „European Land Mammal Mega-Zones (ELMMZ) and Miocene „Mammal-Zones (MN-Zones)”.− In: RÖSSNER, G. & HEISSIG, K. (eds.): The Miocene Land Mammals of Europe. Verlag Dr. Friedrich Pfeil, München., 9–24.

SZABÓ, M., KOCSIS, L., TÓTH, E., SZABÓ, P., NÉMETH, T. & SEBE, K. (2022): Chondrichthyan (Holocephali, Squalomorphii and Batomorphii) remains from the Badenian of southern Hungary (Tekeres, Mecsek Mountains): the first deepwater cartilaginous fishes from the Middle Miocene of the Central Paratethys.‒ Papers in Palaeontology, e1471, 50 p. doi: 10.1002/spp2.1471

VADÁSZ, E. (1935): A Mecsekhegység [The Mecsek Mountains – in Hungarian].‒ Hungarian Royal Geological Institute, Budapest, 180 p.

VAN DAM, J. & WELTJE, G. (1999): Reconstruction of the Late Miocene climate of Spain using rodent palaeocommunity successions: an application of end-member modelling.‒ Palaeogeography, Palaeoclimatology, Palaeoecology, 151/4, 267–305. doi: 10.1016/S0031-0182

VAN DER MEULEN, A. & DE BRUIJN, H. (1982): The mammals from the Lower Miocene of Aliveri (Island of Evia, Greece). Part 2. The Gliridae.‒ Proceedings of the Koninkljke Nederlandse Akademie van Wetenschappen, Ser B., 85/4, 485–524.

VENCZEL, M. (2004): Middle Miocene anurans from the Carpathian Basin.‒ Palaeontographica Abteilung A 271, 151–174. doi: 10.1127/pala/271/2004/151

VENCZEL, M. & HÍR, J. (2013): Amphibians and Squamates from the Miocene of Felsőtárkány Basin, N-Hungary.‒ Palaeontographica Abteilung A 300, 117–158. doi: 10.1127/pala/300/2013/117

VENCZEL, M. & HÍR, J. (2015): Lissamphibians and squamate reptiles from the early middle Miocene of Litke, northern Hungary.‒ Geobios, 48, 491–504. doi: 10.1016/j.geobios.2015.09.001

WEERD VAN DE, A. & DAAMS, R. (1978): Quantitative composition of rodent faunas in the Spanish Neogene and paleoecological implications.‒ In: Proceedings of the Koninklijke Nederlandse Akademie van Wetenschappen, Ser. B, 81/4, 448-473.

ZACHOS, J., PAGANI, M., SLOAN, S., THOMAS, E., BILLUPS, K. (2001): Trends, rhythms, and aberrations in global climate 65 Ma to present.‒ Science, 292, 686–693. doi: 10.1126/science.1059412

ZIEGLER, R. & FAHLBUSCH, V. (1986): Kleinsäuger-Faunen aus der basalen Oberen Süsswasser-Molasse Niederbayerns.− Zitteliana, 14, 3–80.