FRAGMENTA MINERALOGICA E T PALAEONTOLOGICA

18. BUDAPEST, 1996 p. 91-102

Lower Pleistocene vertebrate faunas from the localities 16 and 17 of Beremend (Southern Hungary)

by

D. Jánossy

JÁNOSSY, D. (1996): Lower Pleistocene vertebrate faunas from the localities 16 and 17 of Beremend (Southern Hungary). - Fragm. Min et Pal., 18: 91-102.

Abstract: A summary is given about the fauna of the Locality 16(17) of the great quarry system of Beremend, outstanding in the classical fauna! series of the Villány Mountains. The vertebrate fauna consists of several thousand bones belonging to more than one hundred species. The bird fauna is the richest in the series of the Lower Pleistocene ones of Hungary. The bats, due to their large number of species as individuals, give a whole picture of the environment and age, first time in the Villány Mountains. Other small mammal species make possible an exact stratigraphical evaluation of the fauna (characterized by Mimomys savini together with Allophaiomys pliocaenicus). Last but not least, very complete skeletal remains of large carnivore mammals and ungulates enrich the picture of this colourful vertebrate fauna.

INTRODUCTION

The Lower Cretaceous dark Requienian limestone of the flat Szőlőhegy (= Vineyard Hill) of Beremend, about 9 km south of Villány (southernmost part of Hungary) has been quarried for more than hundred years. Presently, intensive commercial quarrying is being carried out; karstic hollows, caves and fissures containing bones are opened nearly every year.

Already in 1847, Salamon János P E T É N Y I together with Ágoston K U B I N Y I made collections in three different localities in the quarry of that time (Beremend Loc. 1, 2 and 3). P E T É N Y I described 8 mammal species, new for science, and his descriptions meet even the correct, modern standards. Beginning with the turn of this century, T . K O R M O S , Gy. M É H E L Y and later J . N O S Z K Y collected further paleontological material in different localities in the hill (Loc. 4-5). In the 1970's, D. JÁNOSSY and G Y . T O P Á L

collected bone material from various (mainly scattered) localities (Loc.B/6-15). Shortly after the discovery of the "Crystal Cave" in the quarry system of Beremend

in 1984/85 it turned out that this locality is prominent not only by its dripstone and spheric hollow forms, but also as a vertebrate paleontological locality.

In the course of collecting in different parts of red clay from 1985 to 1987 in this locality (No. 16) of Beremend we marked our samples as B/16/1 to B/16/20 (the different numbers do not mean different topography but various collections in different

times). Thus, we give the lists of them summarized (details see in the inventory of the original material in the collection of Budapest) (Fig. 1).

In the quarry, before and above the (artificial) entrance of the cave, we found also a rich paleontological material, which seem to be contemporaneous with the former collected matter and we designated it as Locality 17, Beremend (B/17).

On the basis of all these, TAKÁCSNÉ BOLNER (1985) published a short interpretation of the fauna, according to determinations by D. JÁNOSSY and G Y . TOPÁL. She published the main Locality 16 Beremend under the designation "Passage of Bones".

We collected the large bones as well as a rich microfauna on the site and we obtained and washed about fifty kilogramm red clay, later picked out for bones too.

O 1 0 m

Fig. I . Sketch map of the horizontal plane of the "Crystal Cave" of Beremend at the level of the (artificial) entrance. Asterisks indicate places of paleontological localities of

Beremend 16

T H E FOSSIL REMAINS

In the following I give the list of the fossils, collected in the localities, with some systematical - stratigraphical remarks.

Celtis sp.; B/17: one whole and 89 fragments of the kernels of this Mediterranean tree, otherwise widespread in the Lower Pleistocene to present in the Villány-Mountains, were present among the bones. Celtis sp. is lacking in the Loc.B/16.

Osteichthyes ("Pisces"); B/16: Fragment of a pharyngeal tooth 5 vertebrae-fragm.; B/17: Fragm. of a pharyngeal tooth and two vertebrae-fragm. The remains originate presumably from Cyprinids.

Amphibia-Reptilia; M. VENCZEL (Museum of Körösvidék, Nagyvárad, Transylvania) selected some species from the herpetological material of roughly assumed as 5000 fragments, their list is given here with his kind permission:

B/16 B/17 Triturus eristatus x Bufo viridis x x Pelobates fuscus x x Rana esculenta-group x x Lacerta cf. viridis x x Natrix tesselata x x Elaphe cf. longissima x x

Aves; remains of 45 species from the material of the two layers were published former (JÁNOSSY 1992). This bird material is the richest one among the hitherto known Lower Pleistocene ornithofaunas of Hungary. In the colourful material, the presence of an extinct form of storks (Ciconia stehlini Jánossy), as well as a small duck (Anas crecca percrecca Jánossy) is especially remarkable. Among Galliformes important osteological remains of Francolins (Francolinus [Lambrechtia] capeki Lambrecht) and of the extinct form of Black Grouses (Lyrurus partium Kretzoi) were found. The first proof of the presence of Little Crake (Porzana cf. parva), of the Waxwing (Bombicilla garrulus) and of the Calandra Lark (Melanocorypha sp.) in the Hungarian Lower Pleistocene ornithofaunas is given.

Erinaceus cf. ostramosi Jánossy; B/16: fragments of 4 molars and those of a premolar and of a (dist.) humerus show the size and morphology of a small Erinaceus europaeus. Stratigraphically we can count with the presence of E. ostramosi.

Talpa cf. fossilis Petényi; B/16: maxilla-fragm., two humeri, 3 phalanges manus; B/17: scapula- and ulna-fragm. The length of the intact humerus measures 16 mm (in recent T. europaea 15-18 mm) and falls, together with the other fragments, in the variation of the recent European species. Therefore we can count with the presence of the middle sized Lower Pleistocene form.

Desmana thermalis Kormos; B/16: I sup., mandibula fr., phalanx; B/17: I sup., mandibula-fragm., ulna fragm., 2 femora, calcaneus, metapodium and caudal vertebra. The only exactly measurable bone is the femur, the length of which measures 17.4 and 18.5 mm (recent 24 mm). This bone and all other remains are smaller and proportionally different from the characteristic features of this species (see JÁNOSSY

1965). Episoriculus gibberodon (Petényi); B/17: mandible, with the complete dentition,

agrees with the typical material in measurement and morphology. Beremendia fissidens (Petényi); B/16: 7 maxilla-fragm., 11 mandibula-fragm., 3 I

sup., 2 I inf., most of the material originates from B16/9. Al l fragments are typical. Sorex cf. runtonensis Kormos; B/16: 1 cranium-fr., 2 incisivus inf., 4 molar-fragm.,

30 mandible-fr. Sorex minutus Linné; B/16: 5 mandible-fr., 1 inc. inferior. Sorex aff. margaritodon Kormos; B/16: cranium-fr., 18 mandible-fragm. Crocidura obtusa Kretzoi; B/16: 3 maxilla-fragm., 10 mandibula-fragm.

Crocidura kornfeldi Kormos; B/16: 2 maxilla-fr. (viscerocrania); 43 mandible-fragm.; B/17: 1 maxilla-fr., 5 mandible-fr.

The above listed remains of Soricids were selected according to their size, within one morphological category. Al l remains are typical Lower Pleistocene forms. The climatical significance of some forms (e.g. the genus Crocidura) will be discussed in the conclusions of this article.

Chiroptera; the bat fauna is outstandingly rich in comparison to those found at the nearly 40 Lower Pleistocene localities of the Villány-Mountains. According to the investigations of G Y . T O P Á É it consists of some thousands of bones of about 20 species. The predominant element is an ancient form of the, today strictly Mediterranean, Mehely's Bat {Rhinolopus mehelyi Matschie) perhaps identical with the subspecies birzebbugensis Storch, described from Malta. The above mentioned climatical significance of some shrews and bats will be discussed later: Rhinolophus-Crocidura and Eptesicus nilssonii-Sorex, occurred together in some points of the locality. According to G Y . T O P Á L , the fossil material of bats helped to clear up the systematical position of some forms (Myotis bechsteini robustus = M. steiningeri; the validity of the species M. schaubi is proved, being previously known only by the type specimen, and now found in a great number in Beremend, etc.). T O P Á L speaks about a hibernating colony, which proves that an open cave were present during the Lower Pleistocene in the Villány Mountains (see T O P Á L in T A K Á C S N É B O L N E R 1985 and oral communication).

Hypolagus beremendensis (Petényi); the about 70 remains represent nearly all anatomical units of a hare, as follows: B/16: molar sup., I sup., mandible with P 3-P 4-M! and the same with all teeth, molar inf., scapula-fragm., 2 ulnae fr., femur prox. fragm. and a complete one, tibia dist. fr., calcaneus, 4 metapodii, phalanx; B/17: I sup., P 3, 6 molar-fr., 5 mandible-fr., humerus dist. fr., 4 ulna-fragm., femur dist. fr., 7 metapodial-fi \ , 17 phalanges, 5 vertebral-fragm. Each present P 3 show the enamel-pattern characteristic for Hypolagus and not for Lepus, which fact agrees with the stratigraphical position of the remains. According to KRETZOI (1956) the valid name is Hypolagus beremendensis and not H. brachygnathus.

Hystrix (sp.?); B/16: complete femur and a morphologically-systematically uncertain phalanx I . The morphological features in the proximal and distal epiphysis of the femur speak unambiguously for the remain of a porcupine. The length of the bone (including the trochanter major) measures 107 mm, the width in the middle is 14 mm, the distal width is 28 mm. For comparison the geologically contemporaneous and anatomically identical bone from Osztramos 8 is the most convenient ( JÁNOSSY 1972); its adequate measurements are: 103/12/24 mm. Thus, the differences in the measurements and the view of the proportions show that the Beremend-specimen is more robust than the Osztramos one. This slight difference does not seem to be enough to decide whether there are two different species or not.

Citellus cf. primigenius Kormos; B/16: 7 fragments of molars, about 20-30% larger than those in my recent comparative material of Citellus citellus. According to their stratigraphical position we have to count with the presence of the above mentioned form.

Apodemus sylvaticus Linné; B/16: 3 maxilla-fragm., 12 mandible-fr.; B/17: 8 maxilla-fragm., 13 mandible-fragm., molar. The remains of this banal form of our Pleistocene do not differ, according to our present knowledge, from the recent species.

Cricetus cricetus nanus Schaub; B/16: 36 maxilla-fr., 16 mandible-fr., 34 molar-fr., humerus-fr., femur-fr.; B/17: 31 maxilla-fr., 31 mandible-fr. The material, represented by about 150 fragments of teeth and bones, falls within the range of variation of the subspecies C. nanus. The length of teeth-row of some few measurable specimens is about 6 mm. The variation of the same measurement in Hungarian Allocricetus material of the Lower and Middle Pleistocene ranges between 4.1 - 5.2 mm, in Cricetus cricetus of the same age between 7.6 - 10.0 mm. C. cricetus nanus seems to be fairly characteristic for the Allophaiomys-faums of our territory (JÁNOSSY 1986).

Mimomys savini Hinton; B/16: 25 Mj and 10 mandibles (only two with the whole tooth-row); B/17: 6 M | (2 in the mandible). The remains of this stratigraphically so important water vole (the size [length of M | 2 .9-3 .3 mm] and enamel pattern of Arvicola [only in one case a Mimomys-\s\et], with strongly hypselodont roots, few cement and triangles with thinner enamel in_front than in caudal side) are very typical (see JÁNOSSY 1961). Only a few of the known faunas with Mimomys savini (more than one hundred but growing in number from year to year) can be characterised by the contemporaneous occurrence of this species with Allophaiomys, which seems to have a stratigraphical significance.

Mimomys (small species); B/16: Maxilla-fragm. with M 2 , M 3 , 6 M j , 2 mandibles with M j ; B/17: Mol. sup., 3 mandibles with M j . I listed under the designation "Mimomys small sp." all rooted molars smaller than those of M. savini. 9 Mj represented the Mimomys pusillus type, one matches M. reidi. It would be desirable to revise these two "morphotypes", which are present from the beginning of the classical Pleistocene, because their evolutionary changes are unknown.

Allophaiomyspliocaenicus Kormos; B/16: 2 Maxilla-fr., 149 M j ; B/17: maxilla-fr., 2 mol, 14 M [ . This is the richest occurrence of this species in Hungary. A. pliocaenicus (the most important "index fossil" among voles, besides Mimomys) is characterised by rootless molars and few cement in the enamel re-entrants of the molars. The morphological variation of Mj is very insignificant, chiefly Arvicola-like (length of M , ranges between 2.5 - 3.0 mm), some specimens are similar to "praenivaloides Chaline", but a typical "Mcrote-pattern" is not observable. Very. typical is the non-differentiated, thin enamel (personally seen in the material from several localities on the occasion of the Neogene Vertebrate Conference in Krakow, 1994). In the localities where samples were taken for absolute chronological determinations, the radiometric ages range around 1-1.2 million years.

Lagurus arankae Kretzoi; B/16: 2 maxilla-fr., molaris, 422 Mj and mandibles; B/17: 53 maxilla-fr., 4 molars, 580 M , and mandibles. This very rich material (more than 1200 specimens) shows, as in the case of Allophaiomys, an insignificant morphological variation of the M j . The morphotype arankae is absolutely predominant and the "pannonicus-type" appears in less than 1 percent. This very homogeneous material of Lagurus underlines the stratigraphical significance of the above mentioned, from this point of view important, voles.

Mustela palerminea Petényi; B/16: cranium-fr., 2 canini, P 4, 2 humeri dist. fr., femur prox. fr., tibia (?= praenivalis), tibia dist. fr.; B/17: two maxilla-fr. with toothrows, mandible with teeth, 3 canini, os penis.

Mustela praenivalis Kormos; B/17: cranium fr., 4 maxilla-fr., M 1 , two mandibula-fr., femur.

The determination of small Mustelids was carried out by the comparison of the corresponding anatomical units with the plus-variants of recent. M. nivalis viz. the minus-variants of the recent M. erminea of the comparative collection. The only complete extremity-bone, the tibia falls in size within the variation of the two recent species.

Pannonictis sp.; A phalanx 1 in B/16 aroused the suspicion that Pannonictis is also present in the fauna. In 1994, an amateur collector, S. PONGRÁCZ (Györújfalu) found a complete mandible with the whole tooth-row of a smaller carnivore in the blasted material before the entrance of the cave. The measurements (length of the tooth-row together with the canine = 48 mm, height of the mand. = 17 mm) as well as the morphology of the M 1 (broad, cuplike excavated talonid, length of the tooth = 14.5 mm) compared with the type material of KORMOS (1931) strengthens the presence of the genus Pannonictis. A detailed description must remain for the future.

Canis cf. etruscus Falconer; B/17: two phalanges represent the size category of a smaller wolf, for which I designate the given stratigraphical name: etruscus.

Vulpes sp.; B/16: mc 2. The bone agrees, in morphology as well as in size, with the same anatomical unit of Vulpes vulpes.

Alopex praeglacialis Kormos; B/16: right and left mandible with whole tooth row; B/17: lower canine. The morphology as well as the measurements of the mandible (length of the toothrow, together with the canine = 56 mm, length of the M ! = 13.5 mm) speaks unambiguosly for the presence of the Arctic fox and not that of the Corsac. Later on we shall discuss the climatic significance of the Arctic fox in the (Mediterranean) Lower Pleistocene.

Canis sp. (large form); B/17: a morphologically typical canid phalanx I digiti 1 is much more larger, as the corresponding bone of the recent wolf in my comparative material. Whether this originates from a large lycaonine (Xenocyon gigas), described by KRETZOI (1938) or from a large canide, remains an open question.

Felis magna Schmerling; B/?17: a fragment of an upper canine of a small fel id, somewhat stronger as my recent Felis silvestris, may be designated by this stratigraphical name.

Epimachairodus latidens Owen; the hitherto richest sabre-tooth material from Hungary was determined, except the dentition, with an "exclusional method", i.e.: large felid-bones, which seem to be different from the recent lion-bones of the comparative material in the Budapest-Collection were determined preliminary as machairodontid. The material originating at least from two individuals, merits in the future a monographical elaboration. We determined provisionally, with the collaboration of I . VÖRÖS (archeozoologist, Hungarian National Museum, Budapest), the following anatomical units: B/16: neurocranium-fragment, two (left and right) unworn upper caninus (Pl. I : 1) and a damaged one, lower caninus, I 3 ,P 4 , mandibula-fragm. with the P3-P4 (Pl. I : 2 ) , scapula-fr., humerus, prox.fr., ulna-fr., pelvis-fr., femur, dist.fr.,

patella, tibia prox. and dist.fr., calcaneus, 6-8 metapodial-fr., 5 phalanx-fr. A detailed

comparison of different anatomical units with those of the skeleton of Homotherium crenatidens from Senéze in the Museum of Basel, which I examined by the courtesy of B. ENGESSER, convinced me that this is an absolutely different, bear-like form, with robust extremity-bones. The Beremend-material originates from a slender-boned form. Thus, the determination as " Epimac hair o dus latidens" seems to be established.

Leo sp.; We could not decide on a series of bones of large felids, whether they originate from machairodontids or from a lion, but the presence of the latter is highly probable (fragments of humeri, ulnae, pelvis etc.), all from B/16.

Hyaena sp.; In the course of selection of the fragments of bones of different mammals I found a distal part of humerus of a larger carnivore in B/16. The shape and size of the bone (distal width about 42 mm, the overall picture of the trochlea, of the epicondyli and of the foramen obturatum) speaks for the presence of a hyena. The size and the shape of the bone fragments agrees well with the corresponding anatomical unit of Crocuta spelaea from an Upper Pleistocene specimen of the Collection of the Budapest Museum (Igric-Cave, Transylvania, dist. width about 52 mm). Taking into consideration that about extremity-bones of hyenas almost nothing is published in the literature, we can hardly tell more about the Beremend-specimen.

Ursus sp.; Among the great number of bones we found an M , and M 1 , a metapodial-fragm. and a phalanx in B/16 originating from a larger bear. The bones refer perhaps to Ursus deningeri or an allied form.

Bovidae; One of the greatest problems of identification of the large-mammalian material of Beremend 16 - 17 is posed by the more than two hundred cranial and postcranial elements of bones and bone-fragments, which originate from antilopes or other bovids. I made drawings and took photocopies and we measured of the entire bones of this group from the locality. Furthermore, by courtesy of B. ENGESSER (Basel), I had the possibility to compare the data of the Hungarian material with the specimens of Leptobos (Seneze, Saint Valuer), Megalovis (Seneze), Nemorhoedus (Gallogoral, skeleton of the neotype from Seneze), and literature data.

Together with I . VÖRÖS we could elaborate the following provisional picture about the bovid material of Beremend: most of the bones speak for the presence of a large Nemorhoedus (= Gallogoral) species (with cca 75 extremity-bones and fragments)(Pl. I : 6). One of the metacarpals is so broad that its measurements (length = 176 mm, proximal width = 55 mm) correspond to those of Ovibos (Pl. I : 7). During the Conference in Krakow (1994) we consulted with A. N. TlKHONOW (Sankt Petersburg), the best expert in the osteology of Ovibos, and in his opinion the mentioned metacarpal bone does not match that of the musk ox.

The next one would be the uncertainly determined Megalovis (Pl. I : 3-4) (about 90 bones and fragments) and a slender built antilope (cca 50 pieces). Scattered bones of the size and proportions of Leptobos (6 pieces) and those of a small bovid (? Ovis sp., 10 pieces) close the row. Unfortunately the horn-cores, so important for the determination, are not present for examination, because most of them get to a private collector (I . MAKOVNYIK) and are not available.

Cervus cf. acoronatus Beninde (Pl. I : 5); Some fragments (about them the distal fragm. of mt.) matches those of a middle-sized Cervus elaphus. The systematical designation is of course a "stratigraphical name".

Equus (lAllohippus) sp.; Only the distal part of a mt 3 (distal width = 50 mm) speaks for the presence of a slender horse in the material of Beremend 16.

Archidiskodon meridionalis (Nesti); A fragment of a molar with two lamellae is the proof for the presence of this stratigraphically important element.

E C O L O G I C A L AND S T R A T I G R A P H I C A L CONCLUSIONS

From ecological point of view the whole animal assemblage, similarly to other ones in the Villány-Mountains, has a steppe character, with a great quantity of snakes (Ophidia), with the presence of bustards (Otis), steppe lemmings (Lagurus), porcupine (Hystrix) from among smaller vertebrates and antilopes in the large mammal-fauna. The proportion of the members of forest-environment (perhaps Epimachairodus and the first appearance of "true deer": Cervus in our territory) and the water elements (fishes, desmans) is subordinate.

The appearance of some "boreal" elements (e.g. Bombicilla garrulus from birds and Alopex from mammals) in this Mediterranean environment speaks for a change in the climatical demands of these forms during their evolution. On the other hand, it seems to be remarkable that, considering shrews, in some places of the localities Crocidura appears to be predominant (B/16/9) together with Apodemus and the strictly Mediterranean bats: Rhinolophus mehelyi, whereas in other places chiefly Sorex (with northern relatives B/16/5/8/19) was found. This facts show how cautious we must be with paleoclimatical conclusions (always individual and chiefly on generic, rather than specific level!).

As the faunal lists show, the localities B/16 and B/17 are, as mentioned above, geologically contemporaneous. They give a colourful and a many-sided picture about the contemporaneous life of vertebrates. They differ from the hitherto known more than forty vertebrate localities of the Villány region in two features:

(1) The special richness of the bat-fauna, which refer to the presence of a cave at that times.

(2) The frequency of large mammals, which may give a basis for supposing the presence of a natural trap (shaft) which functioned as a gatherer of large ungulates and carnivores as well.

At the same time the small mammal bones fossilized in owl-pellets are present also in considerable quantity.

From stratigraphical point of view the appearance of the vole Allophaiomys, present in all parts of the localities, is decisive. Due to the "inundation of information" also in the science, as mentioned above, we know at present about 120 localities in Europe (from Spain, through Middle-Eastern Europe to the Caspian Sea, cf. M A U L 1990), in Middle Asia and in North America (Dakota: MARTIN, oral comm. 1994) characterized by the appearance of this vole. In the rich series of localities in the Villány Mountains this is the fifth occurrence of Allophaiomys (former known from Villány 5/, Nagyharsányhegy 21 /mixed/ and Somssich-hegy 1/ 121, see JÁNOSSY 1986) together with Lagurus arankae. The only Arvicola-Wke pattern of the enamel of M j of Allophaiomys speaks for an earlier stage of the Betfia-Phase.

The contemporaneous occurrence of Allophaiomys with Mimomys savini in B/16, noticed in some localities in Europe ( M A U L 1990), is to be examined in respect of

stratigraphical significance in the future. In any case, the facts mentioned in the review of different systematic units of the locality (e.g. the evolutionary stage of Desmana thermalis, the exclusive appearance of Cricetus cr. nanus, the last 111 appearance of Pannonictis and Episoriculus within a modern carnivore and insectivore fauna, etc.), prove definitely the geological contemporaneity of the whole locality-complex of Beremend 16/17, with a local colorit (E-Europe, Mediterranean).

On the level of our knowledge we can establish, based on the proposal at the description of the fauna of Osztramos 8, the following fine-stratigraphical arrangement for the material of richer faunas of Hungary (Betfia-phase):

(1) AI'lophaiomys-iauna with different Microtus species, presence of some small Mimomys, already with Lagurus pannonicus: Osztramos 12;

(2) AllophaiomysA^mm. with small Mimomys and the large Mimomys savini as well as Lagurus arankae: Beremend 16/17;

(3) Allophaiomys-fauna with small Mimomys and with Lagurus arankae: Osztramos 8.

All in all, we can state that independently from the fine-stratigraphical position of the locality Beremend 16/17, its fauna (together with birds and bats) gives the most colourful picture about this period of Pleistocene, not only in Hungary and the Carpathian Basin, but in the whole range of the animal assemblages of Allophaiomys.

SUMMARIZED LIST OF V E R T E B R A T E S O F T H E L O C A L I T I E S B E R E M E N D 16 AND 17

Osteichthyes (Pisces) Perdix sp. Triturus cristatus Otis kalmani Bufo viridis Otis cf. lambrechti Pelobates fuscus Cr ex sp. Rana esculenta-group Porzana cf. porzana Lacerta cf viridis Porzana cf. parva Natrix tesselata Gallinula chloropus Elaphe cf. longissima Tringa sp. Ciconia stehlini Limosa cf. limosa Anas crecca percrecca Numenius cf. arquatus Anas cf acuta Larus cf ridibundus cf. Spatula clypeata Columba cf livia Anas cf sir eper a Strix intermedia Falco tinnunculus atavus Asio sp. Lyrurus partium Athene veta Tetrastes praebonasia Alauda arvenis Francolinus capeki Galerida sp. Francolinus subfrancolinus Melanocorypha sp. Coturnix coturnix Hirundo sp. Bombycilla cf garrulus Emberiza sp. Turdus aff. iliacus Carduelis sp. Turdus cf. philomelos Pyrrhula sp. Erithacus sp. Serinus sp.

Sturmis cf. vulgaris Hystrix sp. Pyrrhocorax graculus vetus Citellus primigenius Garrulus cf. glandarius Apodemus sylvaticus Corvus janossyi Cricetus cricetus nanus Erinaceus ostramosi Mimomys savini Talpa fossilis Mimomys sp. (small) Desmana thermalis Allophaiomys pliocaenicus Episoriculus gibberodon Lagurus arankae Beremendia fissidens Mustela praenivalis Sorex margaritodon Mustela palerminea Sorex runtonensis Pannonictis sp. Sorex minutus Canis cf etruscus Crocidura ob tusa Canis (cf. Xenocyorí) sp. Crocidura kornfeldi Alopex praeglacialis Rhinolophus mehely birzebbugensis Vulpes sp. Rhinolophus macrorhinus anomalidens Ursus sp. My Otis brandti Felis magna My Otis äff natter er i Epimachairodus latidens Myotis steiningeri Leo sp. Myotis schaubi Hyaena sp. Myotis blythi Megalovis sp. Myotis dasycneme Antilope sp. Myotis äff. frater Gallogoral sp. Myotis sp. lövibos sp. Eptesicus äff. nilssoni Leptobos sp. Plecotus äff. auritus Cervus acoronatus Miniopterus äff. schreibersi Equus (Allohippus) sp. Hypolagus beremendensis A rchidiskodon meridional is

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Author's address:Dr. Dénes JÁNOSSY Department of Geology and Paleontology Hungarian Natural History Museum Budapest, P. O. B. 137. H - 1431 HUNGARY

EXPLANATION TO P L A T E I.

Some bones of larger mammals from Beremend, Loc. 16 (Figures 1-3 approx. 1/2 of natural size, Figs. 4-8 approx. 1/3 of natural size)

1-2: Epimachairodus latidens (1: caninus superior, 2: mandible fragment) 3-4: ?Megalovis sp. (3: mandible fragm., 4: humerus) 5: Cervus acoronatus (metacarpale) 6: Gallogoral sp. (metatarsal) 7: Ovibos-like form (metacarpale) 8: Equus (Allohippus) sp. (metatarsale fragm.)

P L A T E I.