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VERTEBRATE FAUNAS O F THE MIDDLE PLEISTOCENE O F HUNGARY The initial phases of the Middle Pleistocene The upper phase of the Middle Pleistocene Tarkoian.. If we review the sporadical fin

PLEISTOCENE VERTEBRATE FAUNAS OF HUNGARY

1 A J Boucot

EVOLUTION AND EXTINCTION RATE CONTROLS

2 W A Berggren and J A van Couvering

THE LATE NEOGENE-BIOSTRATIGRAPHY, GEOCHRONOLOGY AND PALEOCLIMATOLOGY OF THE LAST 15 MILLION YEARS IN MARINE AND CONTINENTAL SEQUENCES

Developments in Palaeontology and Stratigraphy, 8

PLEISTOCENE VERTEBRATE FAUNAS OF HUNGARY

AkadCrniai Kiad6, Budapest, 1979

A MAGYARORSZAGI PLEISZTOCEN TAGOLASA GERINCES FAUNAK ALAPJAN

Translated by

A Demeter

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Library of Congress Cataloging-in-Publication Data

Jiinossy, DCnes

Pleistocene vertebrate faunas of Hungary

(Developments in paleontology and stratigraphy; 8)

Rev translation of: A magyarorsdgi pleisztocen tagolasa gerinces faunak alapjan

Hi bliography : p

Includes indexes

I Geology, Stratigraphic-Pleistocene 2 Vertebrates, Fossil 3 Glacial epoch-Hungary

4 Geology-Hungary I Title 11 Series: Developments in palaeontology and stratigraphy; 8 QE697.J2613 1986 566’.09439 85-29182

ISBN 0-444-99526-9 (Vol 8)

lSBN 0-444-41 142-9 (Series)

(c) Akadeniiai Kiadb, Budapest, 1986

Printed in Hungary

CONTENTS

PREFACE

CHARACTERISTIC SEDIMENTARY ROCKS OF THE PLEISTOCENE OF HUNGARY

The filling of basins

Fiuvial sediments

Sediments deposited by wind

Freshwater limestone

Karstic forms

VERTEBRATE FAUNAS O F THE LOWER PLEISTOCENE OF HUNGARY

Boundary faunas between the Pliocene and the Pleistocene Vertebrate faunas of the Lower Pleistocene sensu stricfo Vertebrate faunas of the last phase of the Lower Pleistocene (Csarnbtasubstage)

(Beremend and Upper Villany substages)

(Betfian and Biharian substages)

VERTEBRATE FAUNAS O F THE MIDDLE PLEISTOCENE O F HUNGARY The initial phases of the Middle Pleistocene The upper phase of the Middle Pleistocene (Tarkoian Vertesszolosian and Upponyian substages) (Castellumian and Solymarian substages)

VERTEBRATE FAUNAS OF THE UPPER PLEISTOCENE O F HUNGARY

The initial phase of the Upper Pleistocene Lower Wiirrn” faunas of Hungary (Subalyukian and Tokodian substages) Faunas of the “Middle Wiirm” of Hungary Faunas of the “Upper Wiirm” (Pilisszantoian and Palankian substages) (“Riss-Wiirm” “pre.Wiirrn” Siittoian and Varboian substages)

(Ista116skoian substage)

7

9

9

9

12

14

16

18

19

27

46

70

71

102

114

115

133

144

149

A REVIEW OF THE PLEISTOCENE VERTEBRATE FAUNAS OF HUNGARY 168

Fauna and climate 168

Chronologyof thePleistoceneof Hungary based onvertebrate faunas 171

CONCLUSION 184

REFERENCES 191

INDEX OF LOCALITIES 199

INDEX OF VERTEBRATE TAXA 201

PREFACE

Nine-tenths of the surface or near-surface sediments of Hungary were deposited during the Pleistocene This fact alone is sufficiently significant to indicate that special attention should be paid to the biochronological problems of this era

Studies done in recent decades have demanded increasingly complex methods since we are striving to obtain knowledge in disproportionate detail what is of, in the palaeontological sense, a disproportionately short time period It has become increasingly clear that it is biostratigraphy which will give a solid foundation to the understanding of this era, as it has for the older ones The basis of an accurate

biostratigraphic record is the rapid, irreversible process of vertebrate evolution, complemented by relevant predominance phases Recent findings from malacology supplement the increasingly complex holism (Krolopp, 1982), as do the new physical methods which seem to come out every year, e.g absolute chronology, etc However, these results are informative only when incorporated into the overall microstratigraph-

ic picture drawn on the basis of vertebrate findings This is especially true for some of the short phases of the Pleistocene The stratigraphic picture of certain areas may be accurately drawn only when supported by relevant vertebrate findings The history of vertebrate palaeontological research has taken a very fortunate course in Hungary Jinos Salamon Pettnyi’s (1864) studies were ahead of those of his contemporaries and his work laid the foundations of modern Pleistocene research based on small-mammal finds This line of vertebrate palaeontological studies has been uninterrupted since then, some of the best known geologists having worked on Pleistocene vertebrates, even if sometimes only as “subsidiary” projects Antal Koch (1900) compiled a catalogue in which he summarized all the Pleistocene vertebrates known at that time Pleistocene vertebrate finds have also been reported by Halavits (from 1879 to 1914), Pethii (1901), KadiC (1911, 1916) and last, but not least, by SchrCter (from 1910 to 1953) Since the turn of the century, however, vertebrate palaeontology has been gaining increasing impetus I peed only mention the works

of Kormos, Ehik, Mthely, Mottl and GaB1 Whilst Ehik (1912, 1913, 1914, 1916,

1921, etc.), Mthely (1914) and Kormos (mainly between 1912 and 1937a) have con- tinued further on the small-mammal studies initiated by Petknyi, Gail (publishing between 1928 and 1954) and Mottl (from 1933 onwards, review in 1941) essentially based their works on larger mammals Finally, Mikl6s Kretzoi has been working in the field for over fifty years and his repeated reviews (1938, 1941a,b, 1953, 1961,

1965, 1969) have all added to our knowledge I have followed this line of study for the past twenty-five years, with partial stratigraphic and zoogeographic syntheses

of various phases of the Pleistocene of Hungary (JAnossy, 1960a,b,c, 1963, 1965a,b, 1969a, 1970, 1973a,b) In these latter works the main approaches to the studies were partly faunal predominance of small vertebrates, partly microstratigraphy based on rnicroevolutionary, statistically significant changes A prerequisite to these studies

was the increasingly general use of the washing method in the field of palaeontology (Figs 10 and 11 in this work); for a detailed description of the method, see Jhnossy (1963)

Thus it became possible to reconstruct a microstratigraphic sequence valid ex- clusively for the Carpathian Basin, unique not only in Europe, but also worldwide During the last few years it has become possible, mainly by detailed analyses of avian finds, to reconstruct the faunal effects of the climatic waves of the early Pleisto- cene (Jhnossy, 1973b, 1976a,b, etc.)

A unified approach to the treatment of the whole material in question is made somewhat difficult by the variability, and, therefore, individual aspects of the various strata As mentioned above, Pleistocene sedimentary rocks are extensive in Hungary and the state of research on these sediments is uniquely advanced in international terms The basis for the stratigraphy was provided by the mass of small-mammal remains accumulated in statistically significant amounts in karstic hollows We may extrapolate from these findings and date regional sediments which also contain microfauna, which are unfortunately only individual finds

Since my work concurs a t several points with the data presented in VCrtes’ (1965) handbook, I will pay less attention to the localities of the Upper Pleistocene connected with archaeological finds and I refer the reader to the above-mentioned handbook,

in which all the relevant palaeontological data are also fully included Since the only archaeological locality older than the Upper Pleistocene is that of VCrtessziilos, which VCrtes treated only tangentially in his book, more attention is paid to that material here

In this compilation I have restricted my stratigraphic analyses primarily to animal communities rather rich in species (ten or more species) Since sporadic large-mammal finds do not fit into the overall picture developed in this book, I have included them only in very special cases; the large-mammal material has been reviewed elsewhere (Jhnossy and Voros, 1979)

I express my gratitude to the many persons who have helped me to complete this book by giving advice, conimentiFg on and checking the manuscript; my former academic advisers, the late Gyula Ehik, and especially Mikl6s Kretzoi, and my colleagues Endre Krolopp, Gyorgy Tophl, Lhszl6 Kordos and many others

Last, but not least I wish to thank academician J6zsef Fiilop, who suggested that

1 should write the book

D JANOSSY

Budapest

CHARACTERISTIC SEDIMENTARY ROCKS OF

THE PLEISTOCENE OF HUNGARY

Before considering in detail the vertebrate faunas of the Pleistocene of Hungary, the sedimentary rocks which have provided vertebrate finds are briefly described

The filling of basins*

Fluv ial sediments

Gravel, sand and other fluvial sediments cover large areas of Hungary There are a number of sporadic finds every year from these sediments, but these are only locally useful and, unfortunately, provide few data relevant to classical geomor- phological terrace studies

These sediments are the so-called “meridionalis gravels” which contain sporadic

remains, mainly of the “southern elephant” (Archidiskodon meridionalis) Classical

localities of these finds are, e.g., SzabadhidvCg (= ViroshidvCg), Ercsi (as type locality

in western Hungary), and Aszbd, southeast of Budapest At these localities, the overlying sediment is loess, whereas the underlying rock is Pannonian clay (Hala- vits, 1898; Schlesinger, 1922; SchrCter, 1958, etc.) Apart from these there have been similar finds in many other localities, but it is impossible to allocate them microstratigraphically in accordance with current practice, and therefore I will pass

over these localities, which are mainly of secondary importance anyway Reliable microstratigraphic dating was possible only i n the case of the Kislfing locality in FejCr county (Kretzoi, 1954b); the animal assemblage found there will be described

in detail Otherwise, the meridionalis gravels refer to the Lower Pleistocene only in a broad sense

The first pebble spots containing faunas not of a sporadic nature were found in the freshwater limestone underlying rocks of Buda Castle Hill (KadiC and Mottl,

1944; already containing Parelephas trogontherii)

The greatest number of remains are those from the fluvial sediments of the Upper Pleistocene, unequivocally proved by faunistics (“Wiirm terraces”) There are few literature data pertaining to this aspect (e.g KadiC and Mottl, 1944; Mottl, 1942), but many sporadic finds have been salvaged and deposited in public collections Na- turally, most of these finds are from the mammoth (which are most likely to catch the eyes of palaeontologists not working with vertebrates) and originate from either the beds of the rivers Danube and Tisza, or from their terrace sediments, although some

of the smaller rivers have also provided ample remains Especially noteworthy is the

* Sediments redeposited during solifluctional and other mass movements of the glacials of the Pleistocene are also included here

“bluish clay” of the bed of the Tisza, which, as is well known, is one of the richest locality series for mammoth and woolly rhinoceros in Europe (mainly between Tisza- fured and Szolnok) In addition, river terraces cutting into the northern mountains of Hungary are especially rich in such finds : mainly the river Zagyva, but also the Tarna,

the Saj6, the Hernhd and the Galga (Jinossy and Voros, 1979)

Extensive overlying sedimentary rocks of fluvial origin of the Great Hungarian Plain are very poor in vertebrate finds Until the 1970s, when large-scale, accurately dated deep-boring schemes were embarked upon, there were very few data available Over thirty core-boring drills have been sunk in recent years, from 100 to 1550 m in

depth They have provided on the whole rich material for palaeontological study, containing vertebrate remains unique in a worldwide context, the vast majority of

CHARACTERISTIC SEDIMENTARY ROCKS 11

Fig 2 Thickness of the Quaternary sediments of the Great Hungarian Plain (mainly gravel and sand); Numbers indicate depths in meters below the level of the Adriatic Sea Black dots refer to the more important boring locations where vertebrate fossils were found (after Urbancsek, 1965 ; Kretzoi and Krolopp, 1972)

which are useful as microstratigraphic time-markers (along with very rich mollusc, pollen and ostracod material) A schematic review of this material is given here The sediments are composed of alternate strata of sand, clay and gravel, thus comprising an apparently uninterrupted cyclic series of strata This impression is also

supported by the rich pollen findings (R6nai et al., 1972) In contrast, the vertebrate

finds and the mollusc material show marked periodicity with the intermediate layers

showing signs of erosional discordance (Kretzoi and Krolopp, 1972) These discrep-

ancies continue to provoke heated debate over the reliability of analysis of this material

Obviously, I will attempt to give a review of this group of finds from the point of view of vertebrate fauna remains, as reported by Kretzoi and Krolopp Although the drillings carried out so far by the Central Office of Geology follow only a general north to south direction which traces (but cuts short) the course of the river Tisza, the results obtained so far are significant (Figs 1 and 2)

The upper, maximum 50 to 60 m, more or less evenly-wide sediment stratumis dated

as originating from the older Wiirm, as evidenced by the small-mammal finds Interestingly enough, the small-mammal remains agree well with the sub-surface large-mammal finds (from the bed of the Tisza), which indicate the same time period Downwards from about 60 to 100 m, there occurs a stratum aged in this work as having been deposited during the younger period of the Lower Pleistocene on the uneven Pannonic surface, sometimes in great width (a maximum of 500 m width has been proven by finds) This stratum is characterized primarily by Mimomys savini, Lagurodon pannonicus and Pliomys episcopalis small-mammal species This series of

layers is obviously identical with Kretzoi’s “Lower Biharian” (Templomhegy substage)

In the mollusc fauna this layer is in partial agreement with the stage marked by

“Viviparus bockhi”, which unit is almost always found in sediments devoid of ver- tebrates but containing molluscs

In the above-mentioned series of drillings one may easily follow the series of strata known as “Upper Villafranchian”-“Villhnyian”, as a thin layer corresponding to a

phase marked by Mimomyspliocaenicus, all lying a t various depths in accordance with

the undulating Pannonic surface

There is only one find, an Apodemus dominans from Mindszent, at a depth of 672 m,

which refers to remains of the Csarn6ta substage marking the boundary between the Pliocene and the Pleistocene

Finally, the downward series of finds is terminated by a new genus and species of

vole, Pannonicola brevidens Kretzoi, from 746 m of the JBszladhny boring, which,

incidentally, shows the most complete picture of all the drillings This vole is aged a t least Middle Pliocene

In every boring the vertebrate remains provide evidence for marked periodicity in sediment formation, the bulk of which is aged as Lower Biharian Sincc the complete description of small-mammal succession is based on the mass finds of small mammals

in karstic formations, the surprising results of the sinking of the Great Hungarian Plain would never have been fully understood without thorough knowledge of the Saunas of caves and hollows

Sediments deposited by wind

Large portion of the surface Pleistocene sediments of Hungary are covered by loess

or loess-like sediments which extend over the foothills up to a height of 400 m above sea level The loess and its genesis have received considerable attention in the literature (Bulla, 1938; Krivhn, 1955; PCcsi, 1965, etc.)

However, all these works deal with this type of rock only from the geomorphological point of view, and have developed a stratigraphy based on separate physical stratigraphy (lithology) and absolute dating

Approaching the problem from the faunistical and stratigraphic points of view, we find that although loess is rich in molluscs (“loess snails”; for recent literature, see Krolopp, 1973, 1982, etc.), it is very deficient in vertebrate fossils In contrast with some foreign loess localities, especially those from outside the Carpathian Basin, in Hungary there are very few “loess sites” and even those are mainly connected with archaeological remains (e.g., Shgvhr, Dunafoldvhr, N6grBdvero”ce, Erd, etc., in western Hungary)

The loess rocks may be fitted into the vertebrate stratigraphic series only because

of some sporadical large-mammal finds (mainly mammoth); localities which may be attributed to a more or less diverse small-mammal fauna are very few indeed (e.g Solt

CHARACTERISTIC SEDIMENTARY ROCKS 13

in Pest county: Kormos, 1911a; Szulimhn: Ptcsi, 1965; Szentlbszl6 in western Hungary: the collection of fossils from the latter locality have been deposited in the Hungarian Natural History Museum)

If we review the sporadical finds from regionally extending loess and loess-like sediments of Hungary, there are hardly any remains undoubtedly older than the Upper Pleistocene, merely a few fossils which could possibly be allocated to lower strata The so-called “hard loess” (known as “loess durci” in the French literature, contain-

ing Mimomys pliocaenicus, sometimes Archidiskodon meridionalis: Saint-Vallier,

Stranzendorf; Viret, 1954; Rabeder, 1971), which have been discovered over the past few decades are not readily identified by means of their vertebrate fauna in the regionally analogous loess of Hungary Isolated finds of faunistically supported

“older loess” have, however, been reported

The oldest loess-like sediments referable to the Lower Pleistocene have been found

in Hungary at locality no 5 at Villhny, locality no 4 imbedded in the freshwater limestone of Dunaalmhs and locality no 2 at Somssich Hill * These rocks, however, cannot be considered as typical loess because originally they were situated in cracks and crevices or deposited on freshwater limestone They thus show a somewhat inter- mediate character, similar to cave loess Similarly, we may refer the Lower Pleistocene material of the yellow layers of locality no 8 at Villhny to cave loesses (Kretzoi, 1956),

in which the fauna elements indicate a cool, dry environment at the time of deposition The oldest fauna, undoubtedly Middle Pleistocene, is that of VtrtessziiEs, which is characterized by rich “Mindel”, Upper Biharian vertebrate fauna (Kretzoi and Vtrtes, 1965b, and especially Jhnossy, manuscript) ; this fauna was uncovered from strongly sandy loess, which was closely connected with the underlying freshwater limestone The loess containing the Uppony fauna is only slightly younger in the geological sense than the above-mentioned rocks Although it may be referred to the group of

“loess durci“ (hard substance that may be picked only with difficulty), since it belongs

to the internal material of karstic hollows it is genetically unrelated to the regional loesses

The loess profile of locality no 6 at Siittii shows a sandy facies and contains in its

upper stratum an Upper Pleistocene “interglacial” fauna (“Riss- Wiirm”), with a

pronouncedly tundra small-mammal fauna 5 m below (Riss-loess ?, Jhnossy and Krolopp, 1981)

Naturally, the above faunas will be described in detail

In spite of the fact that data from the above-mentioned locality no 6 at Siittii were the first to be obtained, some further remarks are necessary on its faunistic and physical characters since they provide data which may be generalized to the process of loess formation The profile of locality no 6 is partly composed of fossil soil which contains not woodland but rather a woodland steppe-steppe mosaic fauna The 5 m wide series of strata in the underlying rock are, however, completely uniform, they are light yellow in colour and the fauna indicates a transition from a tundra environ- ment to a warmer (though in the present sense by no means “Mediterranean”) environ- ment Thus, we have further evidence to prove that the biological changes associated with changes in climate are much more rapid and pronounced than the physical changes shown by the sediments (Brunnacker et al., 1980)

* In all cases the sediments from Somssich Hill are yellowish in colour, the physical appearance,

etc all indicating loess By courtesy of Laszl6 Kordos, grain distribution and chemical tests have been carried out in the laboratory of the National Geological Institute According to theze results, the size fraction 0.01-0.02-0.05 mm constitutes 55.20% and the CaC03 content is very high (50.51%), both values corresponding to those of loess

primarily due to examination of the loose sediments which accompany the freshwater limestone formation by means of the washing method

0 1 2 k m

u

Fig 3a Distribution of the freshwater limestone deposits (shaded in black) around the edge of the Gerecse Mountains Patches given designated by names have provided vertebrate faunas (after Schreter, 1953); for details, see the text

CHARACTERISTIC SEDIMENTARY ROCKS 15

Fig 3b Distribution of the freshwater limestone deposits (shaded in black) in the Buda Hills Patches designated by names have provided vertebrate faunas (after Schrkter, 1953); for details,

see the text

The increasingly refined stratigraphy of freshwater limestones by means of geo- morphology, absolute dating, etc., is supplemented by the following mosaic-like vertebrate palaeontological data

Because of the lack of vertebrate fossils, the most problematic limestones from the faunistical-stratigraphical points of view are the patches situated highest on the

Szabadsig Hill (Csillebtrc) in Budapest Based on their mollusc fauna, they are un- doubtedly from the Pliocene A fragmentary Hipparion metatarsus and other vertebrate

palaeontological finds recently discovered have also confirmed this view

The next locality by age is the upper layer at locality no 4 of the Dunaalmhs

assemblage, which is readily allocated from its small-mammal finds: its age is un- questionably Lower Pleistocene (the upper part of the Villhny biotic zone, according

to obsolete nomenclature, “Giinz”, “Middle Villafranchian”, etc.) A freshwater limestone complex at Siitt6 has proved to be somewhat older than this, according to

a recent revision of the large-mammal fauna (see below: Jhnossy and Krolopp, 1981)

Similarly, small-mammal material of other collections (Krolopp, 1961) have helped

to determine the age of the limestones of Uromhegy; these can be allocated to Temp- lomhegy biotic zone of the Biharian phase of faunal genesis (according to older ter- minology, “Gum-Mindel”, “Upper Cromer”, etc.) The limestones of Budakalhsz are probably of similar age (Jhnossy, 1961a)

Collecting activities aimed at obtaining small mammals have failed at the freshwater limestone deposits of Kiscell, on the outskirts of Budapest, but in view of the large- mammal finds of older acquisition, an age of older (Lower-Middle?) Pleistocene may

be estimated (Schrtter, 1953, pp 120-121)

With the rich small-mammal material available, we may confidently say that Vtrtessz6lBs and the Virhegy (Castle Hill in Budapest) originate from the younger part of the Middle Pleistocene, from the TarkB substage of the Biharian (previously:

“Mindel” - “Mindel-Riss”) Vtrtesszoliis has been shown to be somewhat older than Virhegy (Kretzoi and Vtrtes, 1965b, Jinossy, 1969a, and manuscript)

Finally, there are two Upper Pleistocene localities with rich fauna, one at Tokod- HegyeskG and the other at Tata, both in western Hungary In a recent revision,

I have shown that though both are Upper Pleistocene (“Lower Wiirm”, Subalyuk- Tokod substages), the “principal fauna” of Tata is in the geological sense older than the one at Tokod

Karstic forms

As is well known, the karstic forms ofthe limestone mountains of Hungary developed

during the Pliocene and the Pleistocene, and the filling up, multiple secondary limestone deposition and high lime content render them superbly suited for fossilization of bone remains Especially important are the smaller caves, fissures and hollows, since these are the karstic formations which have preserved the vertebrate remains of the geohistorical phase we are most interested in, and these are the finds on which a microstratigraphical series unique in Europe could b2 based

As has already been mentioned and will be again in more detail, two karstic systems

of fissures serendipitously supplemmt each other as regards geographical-climatic (facies) properties These are the Villiny mountains and Osztramos, each showing the Mediterranean and the northern Carpathian nature of the Middle and Upper Pliocene, and the Lower Pleistocene

Just as with the other sediment assemblages, the Middle Pleistocene is represented

by only sporadic localities that have survived only in special karstic formations

(Tarko-Uppony-Hilton-Solymhr-Siitt6, etc.)

In contrast, finds from the filling material of most of the caves of varying lengths and hollows, right from the Aggtelek-Jbsvaf6 mountains through the Biikk, the Pilis,

CHARACTERISTIC SEDIMENTARY ROCKS 17

the Gerecse and the Bakony, then southwards to the Mecsek, show a varied combina- tion of the Upper Pleistocene and the Holocene Most ofthe localities are supplemented

by archaeological sites, and are characterized just as well in the variable facies corre- sponding to their original geographical positions, as in the clefts and crevices preserving the older Pleistocene (Fig 4)

Buda Hills

r‘

Fig 4 Geographical distribution of highlands in Hungary, where karstic formations

vertebrate palaeontological remains are known

containing

There is no published review of the faunas of cave sediments and karstic formations

of Hungary apart from the fauna lists presented by Mottl(1941), the Upper Pleistocene

cave finds given in Vtrtes’s (1965) book and some smaller treatises on the faunas of local regions (VillBny mountains; Kormos, 1937b; Kretzoi, 1956; Osztramos locality complex: JBnossy and Kordos, 1976b)

I have already referred to the paradoxical position of the faunas of the sediments of karstic formations : these are local, but nevertheless may be of decisive important? when estimating the age of regional sediments

VERTEBRATE FAUNAS OF THE LOWER PLEISTOCENE OF HUNGARY

The series of strata beginning with marine remains are replaced from the second half of the Pliocene onwards by a stratigraphical succession of remains of brackish, freshwater and finally pronouncedly terrestrial sediments Naturally, because of isola- tion, the faunas of a sea broken into an increasing number of small lakes are unlikely

to be as uniform as those the sediments of extensive seas Since, as indicated above, it

is important to refine the stratigraphy of this period, espscially that of the later Pleistocene, it is natural that terrestrial vertebrates (i.e small vertebrates) showing rapid evolution and varying dominance phases should become the focus of attention

It follows from the above as a matter of course that identification of marine or terrestrial strata is highly uncertain Also our knowledge is very sketchy regarding the fauna in between the vertebrate faunas of the classical Lower Pliocene (“Pannonian”,

“Hipparion faunas” in the narrow sense) and the well known animal communities of

the uppermost Pliocene (Csarnbta) The first pieces of the mosaic came from recent discoveries of “Middle Pliocene” localities (nos 1 and 9) at Osztramos Hill, northern Hungary These findings, however, represent only very short phases of this long period

In the Hungarian literature, this uncertain period used to be called “Levantine” after the Slavonian series, at least as far as the regional sediments are concerned As

stated in the introduction, the stratum marked by Viviparus bockhi, which reaches

great widths because of the sinking of the Great Hungarian Plain, has been referred to

as Levantine ever since HalavBts’s work (1888), but the vertebrate finds allocate it to

the Pleistocene (Mimomys suvini, Lower Biharian, Lower or-according to other opinion-Middle Pleistocene) Thus, the former designation of Levantine is highly uncertain and reference to the Pliocene is at best valid for the lowermost layers There have also been attempts to draw a comparison between the Upper Pliocene and the Mediterranean Astian transgression, since the sediments, several hundred meters in depth in the South of France (sand at Montpellier, clay in the vicinity of Perpignan), yield vertebrate finds every year which can be used for comparison (Kretzoi, 1969, etc., Michaux, in litt.)

If Rbth’s (1879) too loose definition of the Pannonian (see Bartha, 1971, p 29) is applied only to the Lower Pliocene, then we may call the period in question post- Pannonian Pliocene Bartha’s work (1973) employs the designation Pannonian for the whole of the Pliocene and coins the term “Upper Part of the Late Pannonian” for the phase best described by the Baltaviir-Estramontium-Ruscinium and Csarnbta terrestrial vertebrate palaeontological “biozones”

Irrespective of these problems of nomenclature this phase is significant from the biochronological aspect, and is marked by repeated climatic changes Following Kretzoi (1962, 1969, etc.) and JBnossy (1972a, 1974), the sequence of terrestrial vertebrate faunas may be characterized as follows

FAUNAS OF THE LOWER PLEISTOCENE 19

The BaltavBr vertebrate fauna, which, unfortunately has not yet been analysed by modern methods, indicates an extensive grassland environment on a continental scale and may be characterized by “Pannonian” (Lower Pliocene) Hipparion fauna

As we have seen above, even a scanty impression of the vertebrate faunas of the phases subsequent to this one is difficult to obtain However, inferring from the faunas around the edges of the Carpathian Basin (Ivan6cz, AjnBcskB, Bar6t-Kopec) and other localities in Europe (Wolfersheim, Montpellier, etc.), it is becoming increas- ingly clear that here, too, there must have been a phase abounding in forests, which gradually replaced the general grassland-puszta (steppe) vegetation Characteristic

of this phase was the influx of southern, southeastern and eastern Asian faunal ele- ments to the area Amongst others, characteristic elements are the Upper Pliocene

mastodons (Anuncus urvernensis and Zygolophodon borsoni, which, however, appeared

a t a later stage and became extinct by the Lower Pleistocene and are therefore of little value as time-markers), tapirs, swine and bullock species related to Indian forms

(Propotamochoerus and Purabos), and of the carnivores, cat bears (Puruilurus) presently

restricted in their distribution to the sub-Himalayan region Murids and cricetids occurred in large numbers among the rodent fauna and of the varied insectivore fauna relatives of present-day east-Asian forms dominated

In Hungary, the Osztramos microfaunas representing shorter phases of this period are noteworthy Although not important for an overall reconstruction of the period, they are significant in two respects

(1) Murids dominate in both faunas rich in small-mammal species (Osztramos, nos

1 and 9), but cricetids are completely absent from locality no 9 This striking difference

is indicative of the fairly regular climatic fluctuations of the Upper Pliocene

(2) Although voles, which play an important role in the stratigraphy of the Pleisto-

cene (Arvicolids, the “foraminifera of the Pleistocene”) are very ancient forms - little

divergent from the cricetid stage going back to the Pannonian of Hungary (Punnonicolu

brevidens Kretzoi, 1965) - the first “true voles” of Europe, obviously Asiatic new-

comers, appeared in locality no 9 of Osztramos (Mimomys silasensis JBnossy, 1974;

Promimomys microdon JBnossy, 1974), but always as individual, rare specimens

The next phase is marked by an explosive diversification in species and increase in the numbers of voles, and may be considered as the dawn of the Pleistocene

Boundary faunas between the Pliocene and the Pleistocene

(Csamota substage)

Three of the four vertebrate fossil localities ca 1.5 km south of the village of

Csamota are undoubtedly of the Csarn6ta layer This locality lies in the westernmost part of the VillBny mountains in southern Hungary (Fig 5), on the flat ridge of Cserhegy, and comprises pillars of red clay formed in limestone of the Middle Triassic (Anisian) which had been left by quarrymen as dead rock in small dispersed quarries PBlfy was the first to draw attention to the localities and it was he who encouraged Kormos, who collected between 1910 and the 1930s, mainly at the classical locality no

2 Finally, Kretzoi excavated in the company of the author, amongst others, in the very same locality, bringing material to the surface from 25 strata and a depth of

about 3 m

In the following section I present a reconstructed faunal list of locality no 2, based

mainly on Kretzoi’s monographic treatises (1959, 1962) and my own revision of the avian finds (Jhnossy, 1976b) (As Kretzoi notes, the old lists of Kormos often do not

Fig 5 Upper Pliocene and Lower Pleistocene vertebrate fossil localities in the Villany Mountains (after Kretzoi, 1956)

indicate the exact site of the locality from which the material has been obtained, a common practice of the time.) The pillar excavated by Kormos and the pit 1 X 1.5 m in size worked by Kretzoi contained the following (as I have mentioned, reconstructed) fauna, with the number of specimens also indicated (species with unknown number

of specimens are marked by asterisks)

Anura, mainly Bufo sp.-ca 20,000

Lacerta ruscinensis Deptret-several thousand

Lacerta aff agilis L.-few

Anguis sp.-several thousand

Ophisaurus intermedius Bolkay-common

Varanus deserticolus Bolkay-few

Ophidia indet.-abundant

Testudo cf lambrechti Szalai-rare

Tetrao macropus JBnossy *-few

1977)

* The two Pliogallus species described by Gaillard are to be deleted from the list (see Janossy,

FAUNAS OF THE LOWER PLEISTOCENE 21

Francolinus capeki wezensis JBnossy-sporadic

Aves div indet +c

Talpa csarnotana Kretzoi ca 100

Talpa cf fossilis PetCnyi-rare

Desmana sp indet I-11-rare

Beremendia Jissidens (Petbnyi) ca 400

Blarinoides mariae Sulimski-few

Petenyia hungarica Kormos-ca 100

Episoriculus gibberodon (Petinyi) ca 100

Petenyiella gracilis (Petinyi)%

Pliopetes hungaricus Kretzoi-few

Pliopetaurista pliocaenica (DepCret)-2

Glirurus gemmula Kretzoi-1

Glis minor Kowalski-few

Muscardinus sp.-few

Prospalax priscus (Nehring)-ca 600

Cricetinus europaeus Kretzoi-few

Baranomys loczyi Kormos-few

Promimomys cor Kretzoi-1

Cseria gracilis Kretzoi common

Dolomys nehringi Kretzoi common

Propliomys hungaricus (Kormos)-abundant

Micromys praeminutus Kretzoi-few

Rhagapodemus frequens Kretzoi-common

Canidae 1-111 indet.-few

Ursus s 1 sp indet.-few

Mustela aff palerminea (Pet6nyi)-few

Baranogale beremendensis (Pet6nyi)-few

Xenictis pilgrimi (Kormos)-few

Machairodontida indet +

Felis s 1 indet (size of catus)-1

L y n x (?) sp indet.%

Cervus (?Rusa) sp indet.X

(Megaceros cf dupuisi Stehlin-unlikely, according to Kretzoi)

“Alces” sp indet.+

Gazellospira cf torticornis Aymard +

Procamptoceras cf brivatense Schaub X

Hemitragus cf bonali Harlt et StehIinX

Ochotonoides csarnotanus Kretzoi-1-2

Hypolagus cf beremendensis (Pet6nyi)-few

Picture 1 The cave of locality no 4 of Csarn6ta (photo by Fejfar)

Inspection of this substantial faunal list reveals that in general the faunal elements showing a relationship with contemporary eastern and southeastern Asian forms, as mentioned above, are very characteristic Such elements are the giant salamander

(Megalobatrachida), certain birds (Francolinus), shrews (Petenyia, Episoriculus), flying squirrels (Pliopetes, Pilopetaurista) and certain dormice (Glirurus, Dryomimus) At the same time, the mass influx of voles (Promimomys, Cseria, Dolomys, Propliomys), besides murids (Apodemus, Rhagapodemus) is clearly indicative of the forthcoming

Pleistocene Specimens from each of the strata showed that there had been a very slow and gradual change through time from a forest animal community to a grassland- steppe faunal assemblage

The mstsrial from sites nos 1 and 3 at Csamota is negligible in comparison with that of no 2, and it is not readily suited to revision because of the older collections and identifications However, as regards age in the geological sense, the material from all

three localities represents the same stratum, whereas locality no 4 is of different age,

as shown by the first pilot collections (Picture 1)

Locality no 7 of Osztramos, a crevice thought to represent the oldest Pleistocene locality of Hungary, because of its similarities with the Csarnbta locality is also described

Osztramos Hill (Pictures 2, 3 and 4), the various localities of which arise repeatedly

in the present discussion (Fig 6), rises insularly above the basin of the Upper Bbdva

between Tornaszentandris and Bbdvarhkb, as the northernmost member of the Rudabinya mountains (in the very north of Hungary)

Pant6 and Kretzoi were the first to collect, in 1955, fossils from a site of this locality complex (Kretzoi, 1956) With my colleagues I carried out excavations from 1965 to

FAUNAS OF THE LOWER PLEISTOCENE 23

Picture 2 Location of the north-northeastern localities (indicated by numbers) of the quarry system of Osztramos (photo by L Kordos, 1972)

1975 at various localities in the enormous quarry of the 380 m a.s.1 hill, made up of Wetterstein limestone from the Triassic-Ladinian (Jhnossy and Kordos, 1976b) The “classical” tectonic fissure, which we named locality no 7, was opened up in

1969 in the course of commercial quarrying operations in the western part of section

XI1 of the quarry This crevice is about 1-1.5 m wide and as high as 30 m The upper part of the karstic formation was filled with dark red clay, the lower with more yellow- ish clay, occasionally cemented with calcite spots (Picture 5)

Picture 3 Location of the south-southwestern localities (indicated by numbers)

of the quarry system of Osztramos (photo by L Kordos, 1972)

Picture 4 Part of the southwestern wall of locality no 12 of the quarry of Osztramos (notice the thin vertical fissure on the left), and locality no 1 (the wider dark fissure-filling material on the right) (photo by L Kordos, 1972)

Picture 5 Locality no 7 of Osztramos (photo by L Kordos, 1972)

FAUNAS O F THE LOWER PLEISTOCENE 25

Fig 6 Pliocene and Pleistocene vertebrate fossil localities excavated up to 1972

Osztramos

in the quarries of

Unfortunately, because of commercial quarrying and the narrow width of the cleft,

collection by the layer was not possible The locality has been found to contain the following taxa, taken from Jhnossy (1 973a), and subsequent identifications of material collected between 1969 and 1977

Francolinus capeki wezensis Jhnossy

Francolinus minor Jhnossy

Bubo sp

Athene veta Jhnossy

Surnia robusta Jhnossy

Passerformes indet

Talpa sp

Desmana sp

Beremendia Jissidens ( PetCnyi)

Blarinoicles marine Sulimski

Petenyia hungarica Kormos

Petenyiella c f gracilis ( Pettnyi)

Episoriculus gibberodon ( PetCnyi)

Chiroptera indet

Pliopetaurista dehneli (Sulimski)

Pliopetes hungaricus Kretzoi

Sminthozapus janossyi SulimTki

Glis minor Kowalski

Miiscardinus sp

Glirulus pusillus (Heller)

Prospalax priscus (Nehring)

Apodemus sp 1-11

Cricetinus sp 1-11

Germanomys cf weileri Heller

Mimomys sp div (incl Cseria sp.)

Lemmus sp.-2 M I

Hystrix cf major Gervais

“Hypolagus beremendensis (Pet6nyi)”-abundant

Canis aff arnensis Del Campana

Vulpes sp

Felis cf lunensis Martelli

Ursus cf minimus Devkze et Boillet

aff Pannonictis janossyi Rabeder

Mustela cf praenivalis Kormos

Mustela aff plioerminea Stach

Putorius stromeri Kormos

Dicerorhinus megarhinus-jeanvireti group

Cervus s 1 sp I (= ?Cervodama pontoborealis Flerow et Pidoplicsko)

Cervus s 1 sp I1 (= cf Cervus philisi Schaub)

I t is instructive to compare the fauna lists of locality no 2 of Csarn6ta and locality

no 7 of Osztramos, since the former lies a t a latitude of 45’53’ and the latter a t 48’31‘

Csarn6ta being in the sub-Mediterranean and Osztramos in the Carpathian climatic zone

Similarly to the fauna of Csarn6ta, francolins, shrews, Bying squirrels and dormice

FAUNAS OF THE LOWER PLEISTOCENE 21

with contemporary relatives in southeastern and eastern Asia (Francolinus, Pliopetau-

rista, Pliopetes, Glirulus) were present a t Osztramos However, instead of the forms of

voles (Dolomys, Propliomys), north-Atlantic elements (Mimomys stehlini), and what

is especially outstanding, lemmings (Lemmus sp.), which are today restricted to the boreal biotic zone, also appeared in Europe for the first time True mice (Apodemus

sp.) were represented only by a few species and in small numbers, in contrast with Csarn6ta

It is noteworthy that considerable large-mammal finds have been recovered from locality no 7 of Osztramos, which, although rather fragmentary, are easily identi- fiable Thus, of the canids of American origin, the presence of a large wolf (Cunis aff

urnensis) was shown by its fragmentary extremities, whereas the presence of the New

World horse, “Equus robustus”, was betrayed by its teeth and limb bones These

immigrants have been considered, even since Haug (1 900), to provide decisive evidence for the boundary between the Pliocene and the Pleistocene In the Hungarian series,

especially important stratigraphically is the large rhinoceros (Dicerorhinus megarhinus-

jeunvireti) of the Upper Pliocene and the lowermost Pleistocene, since it indicates a

substantial difference in the large-mammal fauna compared with that of KislLng where

the smaller D etruscus is the equivalent species and survives right to the Middle

Pleistocene

It is to zoogeographical reasons that we must attribute the complete absence of

antilopes (Gazellospira, Tragospira, Procamptoceras) at Osztramos-though they occurred in similar-aged faunas of the VillLny mountains-and the frequent presence

of cervids (mainly the smaller species: the Cervus “philisi“ group) Last, but not least,

there are the finds of bear related to the Tibetan bear Complete mandibles and maxilla and bones of the appendages are found, unusually complete, all over Europe

(Ursus cf minimus Dev6ze et Bouil.) I have had the opportunity of comparing the finds with the original Perpignanian (Ruscinian) specimens deposited in Lyon, and found slight differences in size of the dentition, which finding is in accordance with the difference in age between the two faunas

In summary, we may say that locality no 7 of Osztramos contains the key to the boundary faunas between the Pliocene and the Pleistocene of Europe

There is hardly any concrete evidence available for this period, which from the biochronological point of view is also important in the regional sediments of Hungary

Only one find of Apodemus cf dominans from a depth of 672 m of the Mindszent boring of the series of drills in the Great Hungarian Plain (Kretzoi and Krolopp, 1972) indicates that parts of the layer in question are also to be found in the depth of the sunken plain

Vertebrate faunas of the Lower Pleistocene sensu strict0

(Beremend and Upper Villiny substages)

We shall begin with the fauna of the karstic clefts of the Sziiliihegy of Beremend, since this is estimated as having originated from the beginning of the Pleistocene (Fig 6a)

The flat Szolohegy of Beremend, 174 m a s l., situated about 9 km south of the

VillAny mountains and the village of Villhny, is composed of Lower Cretaceous dark Requienian limestone and is covered by loess The limestone has been quarried for more than hundred years and presently there is intensive commercial quarrying being carried out; karstic hollows and clefts containing bones are opened u p every year

Fig 6a Pleistocene localities (nos 1-15) in Beremend (situation in 1982)

Kretzoi (1956) described ten localities and since then their number has risen to

sixteen Only the most important ones will be related here, that is, the group oflocalities nos 1-3 of Kretzoi, PetCnyi’s classical sites, and the most recently excavated localities nos 5 and 11

Of historical importance is the work of Jinos Salamon PetCnyi, who together with

Agoston Kubinyi, collected material in 1847 from three clefts (the above-mentioned

nos 1-3), from which he described 8 species as new to science in a manner far

preceding his time; his descriptions meet even correct modern standards (PetCnyi,

1864) The localities have since been demolished but I will list the names of the new taxa (with the current names in parentheses), which are important not only for the history of science but are still of taxonomical-stratigraphical significance The following

is a list of the taxa from localities nos 1-3, with the revised nomenclature of Kretzoi (1956)

Talpa vulgaris fossilis Pettnyi (= T fossilis Pet.)

Crossopus fissidens Pettnyi (= Beremendia fissidens Pet.)

Sorex gracilis PetCnyi (= Petenyiella gracilis Pet.)

Crocidura gibberodon PetCnyi (= Episoriculus gibberodon Pet.)

Lepus beremendensis Pettnyi (= Hyoplagus beremendensis Pet.)

Mustela martelina PetCnyi (= Canis [ s 1.1 martelinus Pet.)

Mustela beremendensis PetCnyi (= Baranogale beremendensis Pet.)

Foetorius palermineus PetCnyi (= Mustela palerminea Pet.)

FAUNAS OF THE LOWER PLEISTOCENE 29

Apart from the above taxa, Nehring (1898) described from one of the three localities

Dolomys milleri Nehring, an “index fossil” of Beremend-aged faunas to this day

Locality no 4 of Beremend could not be identified by Kretzoi, who found that the

list given by Kormos (1937b) contained heterogeneous, differently aged members Noszky salvaged some finds in the large, western quarry of the hill when he was mapping the area in the autumn of 1952 This is the only homogeneous material in the chronological sense of the older acquisitions Kretzoi (1956), applying the name

locality no 5 of Beremend, listed the following taxa (number of specimens in paren-

Desmana kormosi Schreuder-I

Soriculus gibberodon (Pettnyi)-1 1

Petenyiu hungarica Kormos-1

Beremendia jissidens (PetCnyi)-77

Sorex runtonensis Hinton-l

Chiroptera indet.-2

Prospalax priscus (Nehring)-2

Beremendimys noszkyi Kretzoi-1

Rhinocricetus (?) sp.-2

Dolomys milleri Nehring-26

Mimomys mPheIyi Kretzoi-35

Mimomys sp indet (small species)-6

Lagotherium beremendense (PetCnyi)-93

Alopex ( ?) praeglacialis Kormos-9

Ursus (s 1.) sp indet.-1

Gale praenivalis (Kormos)-1

Mustela palerminea (Petinyi)-3

Putorius stromeri Kormos-I4

Xenic tis pilgrimi (Kormos)- 124

Stratigraphical analysis of this fauna shows that it is greatly different from locality

no 2 of Csarnbta and locality no 7 of Osztramos Not only had the archaic cricetids

(Cricetinus, Baranomys), relicts of the Pliocene, disappeared from the area but also the

forest elements (flying squirrels, asiatic Petauristines, dormice Glirulus, etc.) Steppe elements predominated in the animal communities, with the percentile predominance

* Described later by Janossy (1976b and 1978b)

of Dolomys and Mimomys Obsolete shrews (Episoriculus, Petenyia) and mole-rats (Prospalax) are characteristically present The fauna is intermediate between the one

at Csarn6ta and locality no 3 of VillBny Especially significant stratigraphically is the

presence of primitive evolutionary stages (mkhelyi-stehlini) of Mimomys, evolving towards large species (“pliocaenicus”, ostramosi, savini)

Finally, truly remarkable is the fauna in this series which Krolopp and myself

salvaged in the autumn of 1973, a t a site we called “locality no 11 of Beremend”

During the course of very intensive quarrying, mining engineer Mrs Krhlik collected antilope remains (horn cores, parts of toothrows, fragments of limb bones) from a cleft filled with red clay, brought to the surface by blasting, which she presented to the collection of the Hungarian Natural History Museum We paid a visit to the site upon receipt of the material and succeeded in obtaining for sedimentation a portion some kilograms in weight of an enormous block of red clay from the northwestern corner

of the quarry The large- and small-mammal fauna assemblage thus collected comprised the following taxa (with numbers also given):

Francolinus capeki wezensis Jhossy-5

Falco sp (small spncies)-l

Passeriformes i ndet - 1

Talpa sp.-5

Desmana aff nehringi Kormos-71

Beremendia fissidens (Petknyi)-7

Petenyia hungarica Kormos-2

Sorex aff minutus Linnt-1

Sorex aff runtonensis Hinton-3

Soricidae indet.-I

Sicista cf praeloriger Kormos-I

Prospalax priscus (Nehring)-7

Allocricetus (aff bhiki S c h a u b ) 4

Micromys cf praeminutus Kretzoi-3

Estramomys simplex JBnossy-1 I

Chiroptera indet.-2 (molars)

Mimomys mkhelyi-stehlini g r o u p 2 2 M

Mimomys sp (large)-2 molars

Hypolagus beremendensis (Petknyi) l

Ochotona sp.-I

Felidae s 1 (large, Leo or Machairodontidae)-2

ct Tragospira pannonica Kretzoi-18

The stratigraphical nature of the fauna is similar to that of locality no 5 of Bere-

mend, and is thus transitionary between Csarn6ta no 2 and Villhny no 3 Here

Dolomys is completely absent, from which one may infer that the age of the fauna is somewhat younger than that of the other two faunas Besides the above-mentioned

antilope finds (Tragospira), worthy of note is the occurrence of the eomyid Estramomys simplex JBnossy in the VillBny mountains, an extinct Tertiary form described from

FAUNAS O F THE LOWER PLEISTOCENE 3I

Osztramos The discovery of these finds is partly a matter of collecting techniques (the separate teeth may be obtained by washing through sifts with 0.5 mm mesh size), though in the material in question complete maxillae and mandibles have also been found

Interesting from the ecological aspect is that here we have to deal with aquatic sediments This is shown by the relatively large number of fish finds (the Crucian carp,

Curussius, which can be identified from its pharyngeal teeth, lives in small lakes),

but there are also many aquatic desmans (even if the number of finds of Desmunu does not reflect the number of individuals), and many of the bones show signs of having been carried stony substrates across

The fauna next in age in view of the above is the old material collected by Kormos

a t Villhny (Fig 7)

Khroly Hoffmann discovered in 1874 clefts filled with red clay in a quarry on the northern slope of MCszkiihegy (Templomhegy), the most easternly peak of the Villiny mountains (“the quarry opposite to the railway station”) Later, Lbczy, Jr collected from the large quarry on the ridge of the hill (then called the “quarry of Prince Frederick”) Kormos had worked for 30 years on the material of various clefts and finally Kretzoi (1953-1955) and myself (from 1975) have carried out excavations Of

the 11 Villany localities listed by Kretzoi (1956), nos 3 , 5 and 11 will be discussed here

because of their relevant age

The faunas of localities nos 1, 2 and 4 are insignificant and unsuitable for further

analysis (Kretzoi, 1956); the localities with larger numbers are of younger age

Undoubtedly, the site named as locality no 3 is the richest of all the Villhny faunas Kormos called this locality “Villany-Kalkberg-Nord” (Picture 6) The site is situated

on the northern wall of the eastern end of the ridge quarry A cleft, on the average 1 m

wide and lying in a direction east to west, cuts through the Lower Malm Oxfordian limestone, and is filled with red sediment cemented by secondary calcite formation

Fig 7 Lower Pleistocene vertebrate fossil localities in the village of Villany (Villany nos 1-1 1, and Somssich Hill nos 1-2)

Kormos collected at this locality from the 1910s until 1939, but his layer-by-layer fauna lists do not contain quantitative data, a practice of this field of science which was customary at the time The fauna list, updated and reviewed by Kretzoi (1956) and myself (Jhnossy, 1977, 1978a) is as follows:

Bufo viridis (Laurenti)

Lacerta viridis (Laurenti)

Natrix natrix (L.)

Zamenis jugularis caspicus (Gmelin)

Testudo lambrechti Szalai

Lyrurus cf partium Kretzoi

Francolinus subfrancolinus Jhnossy

Talpa minor Freundenberg

Desmana nehringi Kormos

Sorex runtonensis Hinton

Sorex minutus (L.)

Beremendia fissidens (Pettnyi)

Petenyia hungarica Kormos

Episoriculus gibberodon (Pettnyi)

Crocidura kornfeldi Kormos

Picture 6 The inoperative quarry of Templomhegy (MeszkBhegy) of Villany (photo by the author)

FAUNAS OF THE LOWER PLEISTOCENE 33

Erinaceus sp indet

Myotis baranensis Kormos

Myotis steiningeri Kormos

Myotis schaubi Kormos

Myotis wiisti Kormos

Vespertilio majori Kormos

Eptesicus praeglacialis Kormos

Rhinolophus aff ferrumequinum (Schreber)

Citellus primigenius Kormos

Glis sackdillingensis Heller

Dryomimus eliomyoides Kretzoi (not Eliomys sp.)

Prospalax priscus (Nehring)

Apodemus sylvaticus (L.)

Apodemus alsomyoides Schaub

Rhinocricetus Phiki (Schaub)

Mimomys aff pliocaenicus Major

Mimomys fejPrv6ryi Kormos

Mimomys hungaricus Kor m 0s

Mimomys petenyii MChely

Mimomys pusillus MChely

Kislangia rex (Kormos)

Myodes sebaldi (Heller)*

Lagurodon sp indet

Hystrix sp indet

Estramomys simplex Jhnossy * *

Hypolagus beremendensis (PetCnyi)

Pliolagus beremendensis (Kormos)

Canis mosbachensis Soergel

Canis lupus ssp indet

Vulpes ( ? vulpes L.)

Vulpes praecorsac Kormos

Vulpes (?) praeglacialis (Kormos)

Ursus cf gombaszogensis Kretzoi

Ursulus stehlini Kretzoi

Baranogale beremendensis (PetCnyi)

Vormela petenyii Kretzoi

Mustela palerminea (PetCnyi)

Pannonictis pliocaenica Kormos

Xenictis pilgrimi (Kormos)

Epimachairodus hungaricus Kretzoi

“Mammuthus wiisti (Pavlova)”

Stephanorhinus etruscus (Falconer)

* For revision, see Jinossy, 1973a; Janossy and Meulen, 1975

** Collected by Topal in 1973

Equus sp indet

Cervus sp indet

Capreolus sp indet

Bovidae indet

Gazellospira cf torticornis (Aymard)

Tragospira cf pannonica Kretzoi

Procamptocerus cf brivatense Schaub

Hemitragus cf bonali Schaub

This represents the type locality of the “Villhyian”, or rather, the “Upper Villhnyian” fauna Though there have never been any quantitative collections, because of its diversity the material containing both macro- and microfauna is very suitable for that purpose The most important stratigraphical property of this fauna

is the complete absence of the genus Dolomys in the vole fauna which, just as in the fauna of locality no 11 of Beremend, may not be attributed to facies differences because of the proximity of its geographical position It is striking that not only are

Mimomys stehlini-mkhelyi replaced by M pliocaenicus but in general the genus

Mimomys diverges into various species and evolutionary lines simultaneously The reason put forward in the discussion of the fauna of locality no 3 of Osztramos indicates that this “mutational explosion” may be attributed to climatic factors (Fig 8)

I

1 mm

Fig 8 Dentition of the most important index fossil of the Lower Pleistocene, Mimomys

plicocaenicus Forsyth-Ma- jor

a) lower molar of adult specimen; b)occlusal surface

of the first molar of a young specimen; c) and d) lateral view of the first molar of the same specimens Charac- teristic is the strong forrna- tion of roots (after Cha- line, locality: Saint-Vallier,

“Loess durci”)

FAUNAS OF THE LOWER PLEISTOCENE

Picture 7 Locality no 3 of Osztramos during excavation (photo by L Kordos, 1971)

As mentioned several times in the introduction, locality no 3 of Villhy and locality

no 3 of Osztramos represent, in the geological sense, such close periods of time (a sub-Mediterranean and a Carpathian aspect, respectively), that parallel presentation

of the two materials seems to be justified and instructive

Locality no 3 of Osztramos is situated in the southern corner of section XI of the

quarry, at 340 m a.s.1 The red clay sediment deposited in a cave formed by a cleft running in a northwest to southeast direction was blasted in 1971 (Picture 7) At the time of palaeontological collections, in 1969 and 1970, the upper part of the profile,

10 m high and 5-6 m wide, was made up of yellowish sediment closed downwards by a calcite shelf, below which reddish brown sediment was found The material was collected from 9, arbitrarily delimited (20-30 cm wide) strata Analysis by layers has not been carried out so far (the aim ofsuch study would be to detect differences between the layers) According to JQnossy (1969b, 1970), the following faunal list is known, mainly from layers 2 and 3

Talpa cf fossilis Pettnyi

Desmana sp (large)

Sorex cf praeminutus Sulimski

Sorex sp (araneus group)

Beremendia jissidens (Pettnyi)

Petenyia hungarica Kormos

Crocidura cf kornfeldi Kormos

Erinaceus sp (size of europaeus)

Rhinolophus cf ferrumequinum (Schreber)

Rh euryale group

Myotis cf dasycneme (Boie)

Myotis cf schaubi Kormos

Myotis sp 1-11

Plecotus sp

Estramomys simplex Jhnossy

Citellus primigenius Kormos

Glis cf sackdillingensis Heller

Prospalax priscus (Nehring)

Apodemus sp 1-11

Micromys cf praeminutus Kretzoi

Allocricetus khiki Schaub

Trilophomys cf schaubi Fejfar

Germanomys sp

Mimomys exilis (Kretzoi)

Mimomys ostramosensis Jtinossy et Meulen

Mimomys hungaricus Kormos

Mimomys pitymyoides JBnossy et Meulen

Mimomys tornensis Jhnossy et Meulen

Mimomys pusillus-reidi group

“Myodes sebaldi Heller”

Lemmus aff, lemmus (L.)

“Hypolagus beremendensis (Pettnyi)”

Ochotona sp

Canidae indet (size of Vulpes)

Cervus philisi group

Sus strozzii group

This fauna is the most important of all the animal communities of Osztramos, as far

as the taxonomy of the rodents ofthe Lower Pleistocene is concerned Most remarkable

is the occurrence of a surviving representative of the family Eomyidae, which had been

thought to have become extinct in the Miocene This species has been described from this rich locality (Estramomys simplex; Jhnossy, 1969b) In view of recent findings in the Villhny mountains and in the Ukraine, this form is likely to be attributed strati- graphical significance in the future

Also, remains of members of the genus Mimomys in such a complete state of preservation have been found (crania fossilized together with the mandibles) that for the first time the taxonomical status of the relevant species can be clarified, partly by describing a number of new forms (Jhnossy and Meulen, 1975)

Localities no 3 of Villhny and Osztramos contained such similar animal communi- ties that the geological age of those faunas is unquestionably the middle part of the Lower Pleistocene (Upper Villhnyian, middle Villafranchian “event”) The facies differences between the two faunas are not so great as between, e.g., locality no 2 of Csarn6ta and locality no 7 of Osztramos

Most of the species are common to both localities Differences are caused by the absence of southern species a t Osztramos no 3 (Mimomys fejkrvdryi, Kislangia rex and

antilopes among the large-mammal fauna ; also Gazellospira, Tragospira, Procampto- ceras) and the characteristic presence of northwestern f o r m (?Micromys, Trilophomys,

mainly Lemrnus; in the large-mammal fauna only deer : Cervusphilisi group) Villanyia

FAUNAS OF THE LOWER PLEISTOCENE 37

exilis is a common element of Osztramos no 3 but it is absent from Villhny no 3,

even though locality no 5 in the roof, the overlying rock of Villhny, is the type locality

of Villanyia This proves that Osztramos no 3 is somewhat younger than Villhny

no 3 Also, the two Lemmus M a t Osztramos no 3 were found in the upper, yellowish layer of the locality This suggests a climatic basis for the “mutational explosion” of

Mimomys

In view of the above, I have suggested that the fauna of locality no 3 of Osztramos

be designated the Tornan stratigraphical stage

Because of its similar age in the narrow sense, locality no 5 of Villiiny is described

next Kretzoi (1956) characterizes the site as follows: “ right above locality no 3,

a 10-20 em wide cleft with a corroded surface sinks into the karstic surface, which, at a depth of about 1.5-2.0 m is cut by a horizontal shift the rock of the cleft is Oxfordian limestone and its direction coincides with the strike direction.” The following fossils, with numbers of specimens, have been found by Kretzoi (1956)

Celtis sp indet.-2

Bufo sp indet 1%

Bujo sp indet 11%

Rana sp indet.%

Lacerta viridis (Laurenti)X

Ophisaurus intermedius Bolkay %

Ophidia indet. ca 40,000

Aves indet %

Talpa fossilis Pettnyi %

Desmana nehringi Kormos-16

Sorex runtonensis HintonX

Sorex minutus L +c

Drepanosorex margaritodon (Kormos)-1

Beremendia fissidens (Pettnyi)-l84

Petenyia hungarica Kormos x

Soriculus gibberodon (Pettnyi) %

Crocidura kornfeldi Kormos-606

Erinaceus sp indet - 3

Rhinolophus cf ferrumequinum (Schreber)-1

Chiroptera div indet.-57

Citellus primigenius Kormos-28

Prospalax priscus (Nehring)-1 0

Parapodemus sp indet.-1

Apodemus leptodus Kretzoi-1

Apodemus cf sylvaticus (L.)-12

Rhinocricetus Phiki (Schaub)-284

Villanyia exilis Kretzoi-2

Mimomys mkhelyi Kretzoi (= M pliocaenicus group)%

Mimomys petenyii Mthely +

Mimomys hungaricus Kormos %

Mimomys obtusus Kretzoi

Mimomys fejPrvaryi Kormos %

Mimomys arvalinus Kretzoi =pusi1lUs groupt

Mimomys intermedius (Newton) (= savini Hinton)*

Kislangia rex (Kormos)-53

Clethrionomys solus Kretzoi-1

Allophaiomys deucalion Kretzoi-1

Lagotherium beremendense ( Pettnyi) #

Canis sp indet.-2

Vulpes s.1 sp indet.%

Paratanuki martelinm (Pettnyi) #

Ursus cf gombaszogensis Kretzoi-1

Mustela palerminea (Pettnyi)-2

Vormela petenyii Kretzoi +

Baranogale beremendensis (Pettnyi)#

Pannonictis pliocaenicus Kormos-13

Before dwelling further upon the rich assemblage of the cleft fauna so suitable for stratigraphical analysis, we look briefly at the relevant data of the drill material from the Great Hungarian Plain Kretzoi and Krolopp (1 972) reported Mimomys pliocaeni-

cus from layer 504.70-505.10 m at Csongrhd, layer 289.60-292.80 m at Kengyel and

layer 800-802 m at Makb This species (group of species) is overwhelmingly charac-

teristic of Villiinytype faunas [ M osztramosensis, M mkhelyi (partim), etc are

identical with this species] Here again the connection between local and regional sediments is evident although regarding the Great Hungarian Plain our picture is still very incomplete Kretzoi and Krolopp (1972) coined the name Csongriid complex for these sediments containing remains

Of the regional sediments, the freshwater limestone of Siitt6-DunaalmBs will be discussed Until recently, we had had very little palaeontological data from the enor- mous freshwater limestone conglomerate lying above Siitto-Dunaalmb (Picture 8)

Using vertebrate palaeontological data published by Kormos (1925) and Schrtter (1953), Kretzoi (1954b) published a combined list of the Siittii-DunaalmBs fauna with the following vertebrate species:

Testudo siittoensis Szalai

Clemmys mkhelyi Kormos (= Emys orbicularis L.)

FAUNAS OF THE LOWER PLEISTOCENE 39

Cervidue 1-11

Leptobos cf etruscus (Falconer)

Kretzoi says “ the elephant, horse and Meguceros species doubtlessly indicate

that this locality belongs to the ‘meridionalis’ faunas” At present, however, the

“early meridionah faunas” are subdivided into at least four or five fauna waves (substages), so this statement is vague by current scientific criteria Thereafter, with

my colleagues I investigated the area for nearly a decade in quest of more accurate time-marking fossils In the course of this search, we uncovered a molar of an ancient

beaver (Trogontherium sp.) from the gravel mine of the Leshegy-CsGcsoshegy,

fragments of teeth of a canid (Cunis sp.), an undetermined vole from a cleft in quarry

no I of Dunaalmris, a “cast” of a mastodon (Anuncus urvernensis) tooth in the

freshwater limestone of Kiipite, and finally a large number of tooth finds of giant

deer (Meguloceros sp.), though probably of only a single individual, from the sandy

Picture 8 The southwestern working face of the Di6svolgy quarry of Siittii, with locality no 12 (photo by L Kordos)