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PLEISTOCENE VERTEBRATE FAUNAS OF HUNGARY


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PLEISTOCENE VERTEBRATE FAUNAS OF HUNGARY

Developments in Palaeontology and Stratigraphy, 8

PLEISTOCENE VERTEBRATE
FAUNAS OF HUNGARY
by

D. Jiinossy
Geological-PaleontoIogicaIDepartment of the Natural History Museum,
Budapest, Hungary

ELSEVIER
Amsterdam-Oxford-New York-Tokyo 1986


This book is the revised translation of
A MAGYARORSZAGI PLEISZTOCEN TAGOLASA GERINCES FAUNAK ALAPJAN
AkadCrniai Kiad6, Budapest, 1979
Translated by
A. Demeter
Joint edition published by
t-lsevier Science Publishers B.V., Amsterdam, The Netherlands
and
Akademiai Kiadb, Budapest, Hungary
The distribution of this book is being handled by the following publishers
for the U.S.A. and Canada
Elsevier Science Publishing Co., Inc.
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Elsevier Science Publishers
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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 0-444-41142-9 (Series)

(c)Akadeniiai Kiadb, Budapest, 1986
Printed in Hungary


CONTENTS

PREFACE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

7

CHARACTERISTIC SEDIMENTARY ROCKS OF
THE PLEISTOCENE OF HUNGARY . . . . . . . . . . . . . . . . . . . . . . .

9

The filling of basins . .
Fiuvial sediments . .
Sediments deposited by
Freshwater limestone .
Karstic forms . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . .
wind . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . .

VERTEBRATE FAUNAS O F THE LOWER PLEISTOCENE OF HUNGARY . . . . .

9
9
12
14
16

18

Boundary faunas between the Pliocene and the Pleistocene
(Csarnbtasubstage) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Vertebrate faunas of the Lower Pleistocene sensu stricfo
(Beremend and Upper Villany substages) . . . . . . . . . . . . . . . . . . . . . .
Vertebrate faunas of the last phase of the Lower Pleistocene
(Betfian and Biharian substages) . . . . . . . . . . . . . . . . . . . . . . . . . .

46

VERTEBRATE FAUNAS O F THE MIDDLE PLEISTOCENE O F HUNGARY

70

The initial phases of the Middle Pleistocene
(Tarkoian. Vertesszolosian and Upponyian substages)
The upper phase of the Middle Pleistocene
(Castellumian and Solymarian substages) . . . . . .

19
27

. . . . . .

71

. . . . . .

102

VERTEBRATE FAUNAS OF THE UPPER PLEISTOCENE O F HUNGARY

. . . . . 114

The initial phase of the Upper Pleistocene
(“Riss-Wiirm”. “pre.Wiirrn”. Siittoian and Varboian substages) . . . . . . . . . . . . 115
Lower Wiirrn” faunas of Hungary (Subalyukian and Tokodian substages)
. . . . . . . 133
Faunas of the “Middle Wiirm” of Hungary
(Ista116skoian substage) . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
144
Faunas of the “Upper Wiirm” (Pilisszantoian and Palankian substages) . . . . . . . . . 149


6

CONTENTS

A REVIEW OF THE PLEISTOCENE VERTEBRATE FAUNAS OF HUNGARY

. . . . 168

Fauna and climate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Chronologyof thePleistoceneof Hungary based onvertebrate faunas . . . . . . . . . .

168
171

CONCLUSION . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

184

REFERENCES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

191

INDEX OF LOCALITIES . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

199

. . . . . . . . . . . . . . . . . . . . . . . .

201

INDEX OF VERTEBRATE TAXA


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 microstratigraphic 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 continued 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

8

PREFACE

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 exclusively 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 Pleistocene (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 at 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 geomorphological 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 (Halavits, 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. Naturally, 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.


PLEISTOCENE VERTEBRATE FAUNAS OF HUNGARY

70

“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 Tiszafured 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

-

100

Cegled

Ocsod
vecs
Szeieveny
Csongrad
Mindszent

300

LOO

500

600

700

I

I

I

I

I

1

800 r
n
1

I

Jaszladany
Kengyel

200

I

Depth :

m

o m
m
0

- 0

7

Hodmezovasarhely

I
I

3

Mako
Oballa

Raktaloranthkza
Szeged

=

111

0

I

e
Csarnotan

0Villanyian

0 Wirm

Lower

a

Lower Biharian

Pannonlan

Fig. 1. Stratigraphical data of time-marking vertebrate fossils found so far in deep borings in the
Great Hungarian Plain (after Kretzoi and Krolopp, 1972).


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 discrepancies 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-widesediment stratumis dated


12

PLEISTOCENE VERTEBRATE FAUNAS O F HUNGARY

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 vertebrates 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 “loessdurci” in the French literature, containing 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 intermediate 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 environment to a warmer (though in the present sense by no means “Mediterranean”) environment. 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.


14

PLEISTOCENE VERTEBRATE FAUNAS OF HUNGARY

Freshwater limestone
Freshwater limestones deposited in a series of local basins around previous springs

deriving their source along fault lines in the mountains of western Hungary (the edges
of the Buda and the Gerecse mountains) have long been known to contain vertebrate
fossil localities (Figs. 3a, 3b). I mention here only as a matter of interest to the history
of science that the first, unfortunately rather trivial, vertebrate fossil remains were
collected by Pettnyi from freshwater limestone deposits of the Gelltrt Hill in Budapest.
Although the complex synthesis of data relevant to these rocks has been reviewed by
Schrtter (1953) and Scheuer and Schweitzer (1973, 1974), many of the limestones
could be correctly allocated stratigraphically only recently. This achievement is
primarily due to examination of the loose sediments which accompany the freshwater
limestone formation by means of the washing method.

0

1

2km

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.


CHARACTERISTICSEDIMENTARY 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 geomorphology, 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


16

PLEISTOCENE VERTEBRATE FAUNAS OF HUNGARY

Szabadsig Hill (Csillebtrc) in Budapest. Based on their mollusc fauna, they are undoubtedly 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 unquestionably 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 Templomhegy biotic zone of the Biharian phase of faunal genesis (according to older terminology, “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 largemammal 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 TokodHegyeskG and the other at Tata, both in western Hungary. In a recent revision,
I have shown that though both are Upper Pleistocene (“Lower Wiirm”, SubalyukTokod 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 combination 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 corresponding 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 containing
vertebrate palaeontological remains are known.

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.


18

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 isolation, 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 postPannonian 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 increasingly 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 elements to the area. Amongst others, characteristic elements are the Upper Pliocene
mastodons (Anuncus urvernensisand 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
(Propotamochoerusand 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 Pleistocene (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 newcomers, 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


20

PLEISTOCENE VERTEBRATE FAUNAS OF HUNGARY

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 1X 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).

Dipfopoda indet.%
Gastropoda indet .%
Pisces indet.%
Megalobatrachidae-2
Triturus sp.-sporadic
Salamandra sp.-rare
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
* The two Pliogallus species described by Gaillard are to be deleted from the list (see Janossy,
1977).


FAUNAS OF THE LOWER PLEISTOCENE

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)%
Sorex s. 1. sp.%
Hemiechinus cf. samsonowiczi Sulimski-few
Chiroptera indet.-common
cf. Dolichopithecus sp. indet. %
Macaca sp. indet.-sporadic
Sciurus sp. indet.-1
Eutamias (?) sp. indet.-few
Pliopetes hungaricus Kretzoi-few
Pliopetaurista pliocaenica (DepCret)-2
Glirurus gemmula Kretzoi-1
Dryomimus eliomyoides Kretzoi-ca. 1000
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
Apodemus dominans Kretzoi-ca. 8000

Rhagapodemusfrequens 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
Lynx (?) sp. indet.%
Cervus (?Rusa) sp. indet.X
(Megaceros cf. dupuisi Stehlin-unlikely, according to Kretzoi)
“Alces” sp. indet.+
Small Cervidae (Cervus pyrenaicus DepCret?)X
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

21


22

PLEISTOCENE VERTEBRATE FAUNAS OF HUNGARY

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 grasslandsteppe 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 yellowish clay, occasionally cemented with calcite spots (Picture 5).

Picture 3. Location of the south-southwestern localities (indicated by numbers)
o the quarry system of Osztramos (photo by L. Kordos, 1972).
f


24

PLEISTOCENE VERTEBRATE FAUNAS OF HUNGARY

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).