Transcript
l Abbreviate bibliographic a Lynx (Praha), n. s., 32/2001
ISSN 0024-7774
MAMMALIOLOGICKÉ 2PRAVY NOVTTATES MAMMALIOLOGICAE NOVA SÉRIE, SVAZEK SERIES NOVA, FASC.
RIDi / EDITOR-IN-CHIEF PetrBENDA Narodni museum, 115 79 Praha 1 EDITORI CiSLA / EXECUTIVE EDITORS OF THE VOLUME Ivan HORACE« & Jiff MLlKOVSKY REDAKCNI R ADA / EDITORIAL BOARD Miloä ANDÉRA, Jaroslav ÖERVENY, Ludék J. DOBRORUKA, Alexander DUDICH, Jifi GAISLER, Vladimir HANAK, Vladimir HANZAL, Ludovit KOCIAN, Jaroslav PELIKAN, Antonin REITER, Marcel UHRIN
2001
NÄRODNI MUSEUM • PRAHA
Contents Obsah
HORACEK I. & ML!KOVSKY 3.: Focus on fossil mammals: life story of Oldfich Fejfar. -Vyzkumfosilnichsavcù:zivotnipfibèhOldrichaFejfara
9
BARYSHNIKOV G.: The Pleistocene black bear (Ursus thibetanus) from the Urals (Russia). - Pleistocénni medvëd usaty (Ursus thibetanus) z Uralu (Rusko)
33
BENDA P. & OBUCH J.: Notes on the distribution of hedgehogs (Insectivore: Erinaceidae) in Syria. - Poznamky k rozsîfenijezkû(Insectivora: Erinaceidae) v Syrii
45
ERBAJEVA M. A. & DAXNER-HOECK G: Paleogene and Neogene lagomorphs from the Valley of Lakes, Central Mongolia. - Paleogénni a neogénni zajicovci z Ûdoli jezer, stfedni Mongolsko
55
FORSTEN A. & TLEUBERDINA P.: Hipparions (Mammalia: Equidae) from the Mioand Pliocene of Kazakhstan, Central Asia. A review. - Pfehled hiparionû (Mammalia: Equidae) z miocénu a pliocénu Kazachstànu, stfedni Asie
67
HEINRICH W.-D.: Erster Nachweis von Lagurus lagurus (Pallas, 1773) (Mammalia, Rodentia, Arvicolidae) für das Jungpleistozän Norddeutschlands. - Prvni nâlez pestruäky piseêné, Lagurus lagurus (Pallas, 1773) (Mammalia, Rodentia, Arvicolidae), z pozdniho Pleistocénu sevemiho Nêmecka
89
HEISSIO K.: Anthracohyus (Artiodactyla, Mammalia), an Eurasian Achaenodontid. —Anthracohyus, evroasijsky pfislusnik celedi Achaenodontidae (Artiodactyla, Mammalia)
97
HlR J.: New Middle Miocene rodent faunas from northern Hungary. - Nova stfedomiocénni fauna hlodavcu ze severniho Mao"arska
107
HORACEK I. : On the early history of vespertilionid bats in Europe: the Lower Miocene record from the Bohemian Massif. - O rané historii netopyrù deledi Vespertilionidae v Evropé: spodnë miocénni nâlezy z Ceského masivu
123
KHENZYKHENOVA F. I.: Fossil lemmings (Rodentia: Cricetidae) in the Baikal region. - Fosilni lumici (Rodentia: Cricetidae) oblasti jezera Bajkal
155
KOENIGSWALD W. v.: Besonderheiten der Schmelzoberfläche bei Säugetierzähnen. - Zvlaätnosti povrchu skloviny savéich zubü
171
KOLFSCHOTEN T. v.: A fossil wolverine Gulo schlössen (Caraivora, Mustelidae) from Nieuwegein, The Netherlands. - Fosilni rosomâk Gulo schlössen (Carnivora, Mustelidae) z Nieuwegeinu, Nizozemi
183
KRETZOI M.: Trend und Phylum im Säugetiersystem. - Vyvojové trendy a taxony v systému savcû
193
LINDSAY E.: Asynchrony in mammalian biochronology. - Asynchronie v savci biochronologii
201
LOPEZ-MARTINEZ N.: Paleobiogeographical history ofProlagus, an European ochotonid (Lagomorpha). - Paleobiogeograficka historie rodu Prolagus, evropské piät'uchy (Lagomorpha: Ochotonidae)
215
MARKOVA A. K., SMIRNOV N. G., KOZINCEV P. A., KHENZYKHENOVA F. L, SIMAKOVA A. N., ALEKSEEVA N. V, KITAEV L. M. & K.OZARINOV A. V.: Zoogeography of Holocene mammals in northern Eurasia. - Zoogeografie holocénnich savcu severni Evroasie. ...233 MAUL L. C.: The transition from hypsodonty to hypselodonty in the Mimomys savini - Arvicola lineage. - Pfechod od hypsodoncie k hypselodoncii u hrabosi linie Mimomys savini - Arvicola
247
van der MEULEN A. J. & DOUKAS C. S.: The Early Biharian rodent fauna from Tourkoubounia 2 (Athens, Greece). - Fauna hlodavcû casného Biharia z Turkubunia 2 (Athény, Recko)
255
MLIKOVSKY J.: Late Cenozoic biostratigraphy of Europe: mammal zones and the fossil record of birds. - Pozdnê kenozoickâ biostratigrafie Evropy: savci zóny a fosilni nâlezy ptakù
279
PEVZNER M., VANGENGEJM E. & TESAKOV A.: The age of the Ruscinian lower boundary. - Stâri spodni hranice Ruscinia
295
PIKSA K. & WOLOSZYN B. W.: Postglacial bat remains from the Polish Tatra Caves. - Postglaciâlni pozûstatky netopyrù z jeskyni polskych Tater
301
REUMER J. W. F.: Plio-Pleistocene Insectivore Diversity in Central Europe and the Eastern Mediterranean, a Comparison (Mammalia, Lipotyphla). - Srovnàni diversity plio-pleistocénnich hmyzozravcû stfedni Evropy a vychodniho Stfedomofi (Mammalia, Lipotyphla}
313
RÖSSNER G. E. & RUMMEL M.: Pomelomeryx gracilis (Pomel, 1853) (Mammalia, Artiodactyla, Moschidae) from the Lower Miocene karstic fissure filling complex Rothenstein 10/14 (Germany, Bavaria). — Pomelomeryx gracilis (Pomel, 1853) (Mammalia, Artiodactyla, Moschidae) ze spodnëmiocénniho krasového komplexu Rothenstein 10/14 (Nemecko, Bavorsko)
323
STORCH G.: Fossil record of "edentates" outside South America. — Nâlezy fosilnich "chudozubych" mimo Jizni Ameriku
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TESAKOV A., VANGENGEJM E. & PEVZNER M.: Position of reference mammal localities of the Lower Villafranchian in the magnetochronological scale. - Posice referencnich savéich lokalit spodniho Villafranchianu v magnetochorologické Skâle
363
WESSELS W., THEOCHAROPOULOS K. D., DE BRUIJNH. & ÜNAY E.: Myocricetodontinae and Megacricetodontini (Rodentia) from the lower Miocene of NW Anatolia. Fauna podceledi Myocricetodotinae a tribu Megacricetodontini (Rodentia) ze spodnîho miocénu severozâpadni AnatoHe (Turecko)
371
WITHALM G.: Die Pathologien des Höhlenbärenschädels aus dem Heimatmuseum in Golling an der Salzach (Salzburg, Österreich). - Patologie lebky jeskynniho medvéda z Vlastivëdného musea v Gollingu nad Salzachem (Solnohradsko, Rakousko)
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Lynx (Praha), n. s., 32/2001: 183-191.
ISSN 0024-7774
A fossil wolverine Gulo schlossert (Carnivora, Mustelidae) from Nieuwegein, The Netherlands Fosibii rosomäk Gulo schlössen (Carnivora, Mustelidae) z Nieuwegeinu, Nizozemi
Thijs van KOLFSCHOTEN Faculty of Archaeology, Leiden University, P.O. Box 9515, NL-2300 RA Leiden, The Netherlands
[email protected] Received 8 September 2001 Abstract. A fragment of a mandible, assigned to a wolverine, has been collected from sediments dredged up near Nieuwegein, The Netherlands. The specimen is assigned to the Early or early Middle Pleistocene Gulo schlosseri because of its small size and the morphological characters of the p3 and p4. Fossil remains of this species are very rare in the European fossil record. INTRODUCTION Pleistocene mammalian remains have been and still are collected at many localities in the Netherlands, in particular there where sands and gravels are dredged up for example along the main rivers Rhine, Ussel, Waal and Maas. One of the areas, exploited for commercial purposes between 1970 and 1989, is located southwest of the city of Utrecht, near the village of Nieuwegein (Fig. 1). The artificial pit with the coordinates 132.575/449.300 where the sediments have been dredged up is called "The put van Weber". The sands and gravels came from a depth of up to 26 meters according to OFFERMAN-HEYKENS & BROUWERGROENEVELD (1990); however other local collectors mention a maximal depth of about 40 metres. The latter option is more likely because of the age of the mammal fossils, which have been dredged up. The age of the sediments range from Holocene to late Early Pleistocene (RuEQG, pers. comm. 1990). The base of the Holocene deposits varies from -5 to -8 metres; the base of the late Saalian, early Eemian and Weichselian Kreftenheye Formation is at a depth of about 25-30 metres. The Kreftenheye deposits cover sediments with a thickness of 6-7 metres referred to the Urk/Veghel Formation with a Middle Pleistocene age. These sediments lie on top of deposits of the Sterksel Formation dating to the late Early and early Middle Pleistocene. A number of collectors (among which C. BROUWER-GROENEVELD, J. OFFERMAN-HEYKENS, P. STOEL and P. SOUVEREIN) searched for archaeological and fossil remains in the sediments dredged up and gathered a large collection of Middle Palaeolithic artefacts (OFFERMANHEYKENS & BROUWER-GROENEVELD 1990) and Pleistocene and Holocene fossils. The fossil record represent a wide range of species (see Tab. 1); some of them (Mammuthus (Archi-
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diskodon) meridionalis and Stephanorhinus cf. etruscus) date from the Early or early Middle Pleistocene. The vertebrate remains are very fragmentary, only a few complete bones have been collected. One of the most spectacular specimen, described in this paper, is the fragment of a mandible of a wolverine or gluton Gulo schlössen. DESCRIPTION Order Camivora Family Mustelidae Genus Gulo Storr, 1780 Gulo schlössen Kormos, 1914 MATERIAL, fragment of a right mandible with P5 and P4 and the alveoli of C, PI and P, and the anterior root of the M, (Fig. 2 a-c) (collection: C. BROUWER-GROENEVELD; coll. nr. 1). MEASUREMENTS. Tab. 2.
The mandible is incomplete, the anterior and posterior part are broken off. The only elements present are the P5 and the P4 and both are only slightly worn. The shape and size of the alveole indicate the presence of a strong, well-developed canine. The P, had only one root; the PI two roots. The alveoli of these premolars indicate that the P2 was orientated obliquely to the axis of the mandible. The Pj and P4 are conical, their long axis are about parallel to the mandible axis. A well-developed ridge at the lingual side of the crown as well as a lingual and posterior cingulum at the base of the crown can be observed at both premolars. The posterior part of the P4 is well-developed and about 0.4 mm wider than the anterior part. Two foramina mentale are present; one is located below the anterior root of the Pj, the second just anterior of the anterior root of the P4.
Fig. I. Geographical position of the locality Nieuwegein (The Netherlands). 184
Tab. I. List of the vertebrate species identified in the mixed (Early/early Middle Pleistocene - Late Pleistocene - Holocene) faunal association from Nieuwegein (The Netherlands) Esox lucius Pisces indet. Anatidae indet. Aves indet. Emys orbicularis Castor fiber Trogontherium cuvieri Arvicola terrestris Lagomorpha indet. Ursus sp. Gulo schlössen Mammuthus (A.) méridionales
Mammuthus primigenius Elephas (P.) antiquus Coelodonta antiquitatis Stephanorhinus cf. etruscus Equus sp. Sus scrofa Cervus elaphus Rangifer tarandus Capreolus capreolus Cervidae indet. (large deer) Bos / Bison Ovis aries / Capra hircus
The Nieuwegein mandible, although incomplete, shows a number of clear morphological features which indicate that the specimen cannot be referred to ad one of the larger Mustelids such as Aonyx antiquus or Lutra lutra. Aonyx, as known from the Maasvlakte fauna association (VERVOORT-KERKHOFF & van KOLFSCHOTEN 1988, van KOLFSCHOTEN 2001, WILLEMSEN 1992) has compared to the Nieuwegein specimen, a less robust mandible. The location of the posterior foramen mentale differs as well as the shape of the premolars. P3 and P., are smaller and both premolars have an undulated and well-developed cingulum at the lateral base of the crown, a feature which cannot be traced at the Nieuwegein specimen. The mandibles of Lutra lutra are also less robust and the premolars are more slender.
2cm Fig. 2 Gulo schlössen: fragment of a right mandible with P3 and P< and the alveoli of C, P, and P2 and the anterior root of the M, (Fig. 1 a-c) (collection: C. BROUWER-UROENEVELD; coll. nr. 1). a: occlusal view; b: buccal view; c: lingual view.
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The Nieuwegein specimen is a fragment of a rather robust mandible with a obliquely orientated P2 and a slightly expanded posterior part of the P4, characters which can be observed in mandibles of the genus Gulo (BONIFAY 1971, ANDERSON 1977). The size of the mandible and the premolars (see Tab. 2) as well as the morphological features correspond in detail to type specimen of Gulo schlössen Kormos, 1914 (= Gulo Schlössen") from Betfia (= Püspökfürdö) (Romania). Other (incomplete) mandibles of Gulo schlössen, which closely resemble the Nieuwegein specimen, are known from Mundesley (Great Britain) described and figured by NEWTON (1880) as Gulo luscus (= Gulo gulo) and the mandible from Mosbach (Germany) figured and described by TOBIEN (1957). A left branch of a mandible from Krestovka (Kolyma River basin, north-eastern part of the Russian Federation) referred to Gulo cf. schlössen by SOTNIKOVA (1982) shows similar characters as the one described in this paper. An identification as Gulo gulo (Linnaeus, 1758) has been excluded because of the small size (see Tab. 2), the absence of a prominent, well-developed posterior cingulum at the P4, and the fact that the posterior part of the P4 is only slightly wider than the anterior one. Gulo minor Sotnikova, 1982, known from the Lower Adycha River (north-eastern part of the Russian Federation) is smaller than Gulo schlössen. The mandible of Gulo minor is characterized furthermore by a weaker curvature of the tooth row-axis, the direction of the main axes of the P4 and the M,, and the narrow and elongated P3 and P4 (SOTNIKOVA 1982). HABITAT AND EVOLUTIONARY HISTORY OF THE GENUS GULO The present-day wolverine Gulo gulo has a Holarctic distribution ranging from Scandinavia, the Russian Federation to North America and Canada. It inhabits the forests and treeless highlands of the taiga but in summer it also penetrates into the southern part of the tundra (CORBET 1966, BANCI & HARESTAD 1990). Wolverines are not restricted to any single biotope; with a long coarse coat and in wintertime a dense under-fur (even the soles of the feet are furred), they are well adapted to living under conditions of snow and extreme cold. The wolverine diet consists of carrion, small birds and mammals, fruit and berries. The wolverine feeds mainly by scavenging, but during the winter, when the snow conditions are unfavourable for its prey, it is also a capable hunter able to kill a reindeer Rangifer tarandus and even a subadult elk Alces alces. The present distribution of the wolverine coincides with that of Rangifer tarandus, which comprises the major part of its diet. Wolverines are rarely represented in the fossil record, a record that indicates that the genus Gulo evolved in the Old World and that the migration of the genus to North America most probably took place during the later part of the Middle Pleistocene (during the Saalian-Illinoian (KURTEN 1963). The origin of the genus Gulo is very unclear but it has generally been accepted that the Mio-PHocene Plesiogulo should be regarded as the direct ancestor of the Pleistocene Gw/o-lineage (G. minor - G. schlossert — G. gulo) (KURTEN 1963, SOTNIKOVA 1982). The lineage shows a number of evolutionary changes in the lower jaw: a general increase of size (an increase that shows an interruption and a change to a decrease during the Late Pleistocene), a more stronger curvature of the mandible, the development of more robust and wider P3 and P4> and a relative increase of the size of the Pj (SOTNIKOVA 1982). The Pliocene Plesiogulo was replaced by the smaller Gulo minor at the end of the Pliocene; a replacement that might be related to the first major cooling about 2.5 Ma ago
186
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l (BoNiFAY 1971 ). Our knowledge of G. minor so far is restricted to the lower jaw from the Lower Adycha River described by SOTNIKOVA ( 1982). The age of the specimen is most probably Early Pleistocene although a late Pliocene age cannot be excluded (SOTNIKOVA 1982). The next representative of the Guto lineage, Gulo schlössen, is better known; remains referred to as this species are known from several Early as well as Middle Pleistocene localities. Its stratigraphical range will be discussed in more detail in the next paragraph. The living Gulo guh occurred during the late Middle Pleistocene (e.g. Tornewton Cave - Glutton Stratum) as well as the Late Pleistocene. During the late Weichselian, the species was widespread; fossil remains, although rare, have been found (mainly in cave deposits) as far south as the Pyrenees in northern Spain and northern Italy (SUTCLIFFE et al. 1985) as well as in the Caucasus (KURTEN 1968). The G. gulo from Tornewton Cave is about the same size as modern Fennoscandian specimens but about 10% smaller than those dating from the Last Cold Stage (KURTEN 1973). HOLOCENE
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Fig. 3. Stratigraphical range of the different Gulo species and the possible stratigraphical position of the Nieuwegein Gulo schlössen related to the Dutch chronoslratigraphic subdivision of the Quaternary and the palaeomagnctic scale.
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THE STRATIGRAPHICAL RANGE OF GULO SCHLOSSER! The number of fossils referred to as Gulo schlössen is very restricted; in particular if one takes into account the stratigraphical range of the species. The type specimen of Gulo schlössen comes from Betfia VII/1 (Romania); from a fauna with an Early Pleistocene (Waalian-Early Biharian) age according to TERZEA (1995). The remains from Gombazok referred to as Gulo schlosseri by KRHTZOI (1941) are much younger; they have a Late Biharian age. The specimen from the Cramer Forest Bed deposits at Mundesley (Great Britain) might have about the same age as the finds from Gombazok. Although precise stratigraphical data of the mandible from Mundesley are lacking it is generally assumed that the record dates from the Cromerian s. 1. (STUART 1982). The locality Mosbach (Germany) yielded a mandible fragment that has been assigned to Gulo schlosseri as well as remains of a larger wolverine referred to as Gulo gulo (TOBIEN 1957). The smaller Gulo schlosseri is part of the main Mosbach fauna (BRUNING 1980); a fauna with a late Cromerian (early Toringian) age. Tobien (1957) assumed that the laiger Gulo remains derived from the upper deposits referred to the Mindel (= Elsterian) glaciation. Therefore, it has been stated that Gulo schlosseri became extinct at the end of the Cromerian Complex and has been replaced by Gulo gulo during the Elsterian (TOBIEN 1957). However, the cranial and post-cranial Gulo remains from L'Escale (France), referred to as Gulo gulo schlossert by BONIFAY (1971), are also supposed to have a post-Cromerian (Elsterian) age and the dentition as well as the bones are still small in comparison with the dimensions of the living wolverines. BONIFAY (1971), therefore, assumes that the increase in size that marks the transition of Gulo schlosseri to Gulo gulo took place during the later part of the Middle Pleistocene, after the Elsterian. The presence in the fauna of l'Escale indicates, however, that the adaptation to a cold climate took place earlier in time (BONIFAY 1971). Based on the stratigraphical data of the remains listed above it is clear that the occurrence of Gulo schlosseri in Europe ranges from the Waalian to the Elsterian, a time span of about 0.8-1 Ma. CONCLUSIONS The mandible fragment from Nieuwegein, assigned to Gulo schlosseri, is a remarkable find because of the scarcity of the species in the Pleistocene record of Eurasia. Taking into account the long stratigraphical range and its geographically widespread occurrence finds of Gulo schlosseri are extremely rare. The specimen from Nieuwegein most probably date from the later part of the Early Pleistocene or the early Middle Pleistocene, a conclusion based on lithostratigraphical data and accompanying faunal elements such as Mammuthus (Archidiskodon) meridionaiis and Stephanorhinus cf. etruscus. SOUHRN V ulo/cnindch vytëzenych u Niewegeinu (Nizozemi) byl nalezen lilomck spodni delisti rosomâka. Kost byla na zâkladé malych rozmërù a rnorfblogickych znaku na premolarech (p3 a p4) urtena jako £ast jedince rosomâka druhu Gulo schlosseri, pochazejici z raného äi ranê stfedniho pleistocénu. Pozùstatky tohoto rosomâka jsou ve evropském fosilnîm zàznamu velmi vzâcné.
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ACKNOWLEDGEMENTS The author would like to express his gratitude to several persons who contributed to the realization of this paper in various ways. The owner of the specimen, Ms C. BROUWER-GROENEVELD, gave permission to study the mandible fragment. She, together with J. OFFERMANS-HEYKENS, P. STOEL and P. SOUVEREIN, collected the archaeological and palaeontological finds at the locality. Dr. G. H. J. RUEOO (former Geological Survey of the Netherlands) provided the lithostratigraphical data and Dr. J. HR (Hungary) send me additional information on the find from Betfia. W. den HARTOG and J. LUTEYN (Faculty of Earthsciences, Utrecht University) made Fig. 2 and Ms O. YATES corrected the English. This paper is written in honour of Oldfich FEJFAR, a colleague who is just as unique as the specimen described in this paper. A colleague who inspired me because of the scientific and personal interest he showed since the first time we met. I appreciated the fact that in spite of his vast knowledge of the Neogene and Quaternary fossil record he remained a modest and cheerful person. A personality in the field of mammalian palaeontology whom I regard both as a colleague and as a friend.
REFERENCES ANDERSON E., 1977: Pleistocene Mustelidae (Mammalia, Carnivora) from Fairbanks, Alaska. Bull. Mus. Comp. Zool., 148(1): 1-21. BANCI V. & HARESTAD A. S., 1990: Home range and habitat use of wolverines Gulo gulo in Yukon, Canada. Holar. Eco/., 13: 195-200. BONEFAY M.-F., 1971: Carnivores quarternaires du Sud-Est de la France. Mém. Mus. Hist. Nat. C, 21(2): 43-377. BRUNNING H., 1980: Die eiszeitliche Tierwelt von Mosbach: Ihre Umwelt, Ihre Zeit. Museumfîihrer Naturhist. Mus. Mainz, 6: 1-60. CORBET G. B., 1966: The terrestrial mammals of western Europe. Foulis & Co., London, 264 pp. KOLFSCHOTEN T. v., 2001: Pleistocene Mammals from the Netherlands. Pp.: 209-215. In: ROOK L. & TORRE D. (eds.): Neogene and Quaternary continental stratigraphy and mammal evolution Papers in honour of Augusta Azzaroh's outstanding contribution in Geology and Paleontology. Boll. Soc. Paleontol. Ilal., 40(2): 115-313. KORMOS T., 1914: Drei neue Raubtiere aus den Präglazial-Schichten des Somlyóhegy bei Püspökfiirdö. MM. Jahrb. Ung. Geol. Reichsansi., 22: 223-247. KRETZOI M., 1941 : Weitere Beiträge zur Kenntnis der Fauna von Gombaszög^n/i. Mus. Naln. Hung., 24:105-139. KURTEN B., 1963: Notes on some pleistocene mammal migrations from the Palaearctic to the Nearctic. Eiszeitalter und Gegenwart, 14: 96-103. KURTEN B., 1968: Pleistocene mammals of Europe. Weidenfeld and Nicolson, London, 317 pp. KURTEN B., 1973: Fossil Glutton (Gulo gulo (L.)) from Tornewton Cave, South Devon.Comm. Bio!., 66.3-8. NEWTON E.T., 1880: On the occurrence of the glutton, Gulo luscus, in the "Forest Bed" of Mundesley, Norfolk. Geol. Mag., 2, 424-427. OFFERMAN-HEYKENS J. & BROUWER-GROENEVELD C., 1990: Het Midden-Palaeolithicum van Nieuwegein (Utrecht). Westerheem, 39(3): 98-109. SOTNIKOVA M. V., 1982: The History of the genus Gulo in Eurasia. Proc. Zool. Inst. AS USSR, 3: 65-73. STUART A. J., 1982: Pleistocene vertebrates in the British Isles. Longman, London, 212 pp. SUTCLIFFE A. J., LORD T. C., HARMON R. S., IVANOVICH M., RAE A. & HESS J. W., 1985: Wolverine in Northern England at About 83,000 yr B. P.: Faunal Evidence for Climatic Change during Isotope Stage 5. Quat. Res., 24: 73-86. TERZEA E., 1995: Mammalian events in the Quaternary of Romania and correlations with the climatic chronology of Western Europe. Acla Zool. Cracov., 38(1): 109-120.
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ToBiEN H-, 1957: Cuon Hodg. und Culo Frisch (Camivora, Mammalia) aus den altpleistozänen Sanden von Mosbach bei Wiesbaden. Ada Zool. Cracov., 2: 433-445. VERVOORT-KERKHOFF Y. & KOLFSCHOTEN T. v., 1988: Pleistocene and Holocene mammalian faunas from the Maasvlakte near Rotterdam (The Netherlands). Meded. Werkgr. Terl.Kwart. Geol.,25(\): 87-98 WiLLEMSEN G. F., 1992: A revision of the Pliocene and Quaternary Lutnnae from Europe.Ser. Geol., 101: 1-115.
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