Biostratigraphy of the mammalian faunas at the pliocene-pleistocene boundary in middle and western Europe

Palaeogeography, Palaeoclimatology, Pa[aeoeeology Elsevier Publishing Company, Amsterdam - Printed in The Netherlands

B I O S T R A T I G R A P H Y OF T H E M A M M A L I A N F A U N A S AT T H E P L I O C E N E - P L E I S T O C E N E B O U N D A R Y IN M I D D L E A N D W E S T E R N EUROPE H. TOBIEN

Palaeontological Institute, Johanne~Gutenberg University, Mainz (Germany) (Received January 12, 1970)

SUMMARY

During Late Pliocene and Early Pleistocene immigration of new mammal groups and genera (microtines, microtoid cricetines, Archidiskodon, Leptobos, Equus); disappearance of existing groups and genera (Trilophomys, other microtoid cricetids, Anancus arvernensis, Mammut borsoni, Hipparion); in-place evolution (Mimomys group, Dicerorhinus megarhinus-etruscus, cervids, Archidiskodon meridionalis) and combination of these phenomena permit the recognition of a sequence of faunistic "niveaus" (levels) between the Pontian (Lower Pliocene) and the Middle Pleistocene in middle and western Europe. An attempt is made to distribute the larger local faunas of middle and western Europe within five or six of these niveaus. The position of the Pliocene-Pleistocene boundary within the sequence of these niveaus is discussed. Stratigraphic relations to marine faunas exist in the Mediterranean aTea. The Middle Pliocene Roussillon-Montpellier fauna is associated with marine molluscs of Astian character. The Montopoli local fauna (Val d'Arno lnferiore, Italy) of Lower Villafranchian age similarly occurs together with Astian molluscs, whereas the Middle and Upper Villafranchian niveaus seem to have the marine Calabrian as equivalents. Some relations of the m a m m a l fauna niveaus to floristic stages (Reuverian, Pretiglian, Tiglian) are indicated, and radiometric data are discussed. An attempt is made, to correlate the sequence of paleomammatogical niveaus in middle and western Europe with those of the European part of the U.S.S.R. Some relations to the marine stages of the Caspian Sea can hereby be demonstrated. INTRODUCTION

In the last two decades the knowledge of the mammalian faunas at the Pliocene-Pleistocene boundary in middle and western Europe has been considerably increased (ToB~EN, 1969). Together with the faunistic description an effort

Palaeogeography, Palaeoclimatol., Palaeoeeol., 8 (1970) 77-93

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H. TOBIEN

was also made to give stratigraphic details and subdivisions mainly of the Villafranchian (referred to here and on the following pages as Lower Pleistocene), and to a lesser degree of the preceding Late Pliocene. There is not space enough here to discuss these subdivisions in detail. The intention is simply to present another attempt to subdivide the Late Pliocene and Early Pleistocene mammalian faunas in middle and western Europe, in which, however, former stratigraphic data will be reconsidered. The subdivision here presented is based on the combination of biostratigraphic data of certain species or genera or groups of those of fossil mammals important in this respect. The principles and methods are traditional: (1) Immigration of new mammalian genera and/or groups of them to middle and western Europe, e.g., Elephas (Archidiskodon), Leptobos, Equus (abbreviated as E - L - E group), microtines, and "microtoid" cricetines. (2) Disappearance and/or extinction of taxa, living in middle and western Europe before such as mastodonts, Hipparion, Trilophomys, and other taxa. (3) In-place evolution of morphogenetic lineages, e.g., Mimomys group, Dicerorhinus megarhinus-etruscus, cervids, Archidiskodon meridionalis lineage, etc. By these criteria a larger number of middle and western European local faunas, included in the time interval Late Pliocene-Early Pleistocene, can be distributed over several "biostratigraphic niveaus (levels)". The local faunas ascribed to a certain niveau are characterized by all or at least one of the above mentioned criteria. The niveaus are named after localities typical for them. The geological time span embraced by a niveau may correspond to a stage or substage or also to a zone of the international stratigraphic nomenclature code (HEDBERG, 1961). The use of the name "zone" for these niveaus is avoided here because several niveaus very probably include a larger stratigraphic time interval than a biostratigraphic zone in the normal and formal sense. Furthermore the stratigraphic code recommends (HEDBERG, 1961, p.11, 12, 22) the use of the word "zone" in a formal sense only in connection with the name of a species or genus typifying the zone. The elaboration of zones as used in the marine biostratigraphy is not envisaged in this paper and is also difficult to realize as far as fossil mammals are concerned. According to the experiences in the field of mammalian stratigraphy the above-mentioned criteria applied to a local fauna seem to be sufficiently useful and workable tools for stratigraphic purposes, at least at the present state of knowledge and in view of the relative scarcity of fossil mammals and mammalian faunas. Therefore the provisional, informal designation "niveau" ( = level) will be used throughout this paper. The niveaus correspond in many respects to THALER'S (1966, p. 186) "zones biochronologiques". Every niveau is designated by two letters of one or two localities which are most characteristic for the niveau concerned (for instance: Rou-A1 niveau, abbreviation for: Roussillon and Alcoy faunistic niveau, see Table I). The first

Palaeogeography,PalaeoclimatoL,Palaeoecol., 8 (1970) 77-93

S t a g e s and substages

Niveau 5eque~:c(?

'

i

g

Middle

.

-:

:Am=:

:

Csarnotian)

[L o w e r iPliocene (= P o n t i a n s,l.)

I Middle [Pliocene I(= Ruse inianl

[

Uppe r Plioeene

__

Lower Vilhtfra:~('hia n

!

I'$ .~ I V i l l a f r a n c h i a n

*~ ~

gl.

Upper Vilh{frallchian

c

i

Rou-Al Iliveatl

L

We-Csa ni~ eau

? Aj l niveau

-. :=::

El-Me ~ive:{u

Ro-Va niveau

Ar-Se niveati

:

existing

existing

,111111011115

P, fItgnIUttIIS Gc;'*~ecnlone ; 5 5lctclto*u I s

1964)

{

1964) !

group I ( F e j f a r , 1964)

(Fejfar,

g r o u p II

group II / F e j f a r . 1:)64/

(Fejfar,

g r o u p II

g r o u p II and III p.p. (Fejfar 1964)

existing

existing

B i h a r i a n : Middle' P l e i s t o c e n e

Jrilo/)]ll)HlkS

M i d d l e P l e i s t o c e n e (= post T e g e l e n n i x e a u s )

Series

E-L-E

K-L-E

--

existing

existing

existing

b~I'OU 1~

.

cervids as in Aj ( H e i n t z , 1968)

In Rou other

Cevvo('(,rt~ llttsa (W~,ze)

(Rebielice)

C'OFPO('CFI¢% 1~1~'~(I

c e r v i d s of the L o w e r Villafrx::chitu: (He ~ z, 1968)

c e r v i d s el the Middle Villafranchian (He ntz, I 9 6 8 )

c e r v i d s of the U p p e r Vill~'ffranchian ( H e i n t z , 1968)

Ccrvids

group: Eh?p]la,s - l . o / ~ t o l l o ~ A'qtms group

II~¢>,q(l)']lltllI5

n~garhnnt~

i JHC,q'rZI']IiI~I~S

I Ino~at'/lilltts

e/rltsc'ns

t't~'lt'q('lt~

[)l('c'JUl'll/illtS

5

Vialette,

W(Jlfol'sheilll,

Astian (Roussillon, Montpellier)

AstiRn

Astian

Astian ),Iontopo[i J

CMabrian

L 7__

!

'~Calabrian

Marine stages

Reuverian IWiJifersheim

no R e u v o r i a n ~Etou&il'e~, Vialette)

= = = ==

?Pretiglian {LoIIe p.p.)

Tiglian (Tegelen, Val d ' A r n o Montevarchi)

F l c n ' i s t i c slat es

:

i

('a

6.5

A

m.v.:

, F;tt,uz~il't,5

3.4 m . y J

B 3 m.y. 7Z~

[ll,g. (Roccaneyr,1) 2,5-2.6

1.8 m.y. (Chilhac. Coupei )

I~ {diome{l ic daht

!A: P l i o c e n e - P l e i s t o c e n e bokl/ld~tFv b a s e d on E - L - F : ~l'o{ip i B: P l i o c e n e P l e i s t o c e n e b o u n d a r y b a s e d on l';t(litHIlOil'lC d a t a ( S a v a g e and r i l l ' I t s , 1 9 6 8 ;

R o u s s i l l o [ 2, Ah:ov, M o n t p e l l i e r , G~dgdlii

Podhsico, Gundersheim pp.. I~anovce, A r o n d o l l i , S e v n e s S61e, Ngmes, ? V i l h f f r a n c a d ' A s t i

W~Ze. Csal'llolkl-2,

A j n a c s k a , Rqbielico( ? )

Etouaires. Mol~di, I3crc~ti. Malu~tc:li

R o c c a n c y r a , St. V a l l i e r , P a r d i n e s , P u c b l a de V a h ' e r d e , B e r e m e n d ( ? ) , Matee~ti, Berbe~ti. Tuluce~ti V i l l a r o y L Kislang( ? I

[ T e g e l e n , Set ~,ze, N o r w i c h C r a g W e y b o u r n e C r a g of E a s t - R u l l t o i l , Chagny p.p., C h i l h a c , C n u p e l , E r p f i n g e n , L e f f e p.p., VaI d ' A r n o s u p e r i o r e (Mont e v a r c h i - S e r i e s ) , Olivola, Villany-3(?), Irime~ti, Bug ule~ti

]

! Loc'alflies ( s ~ , b < t e d l

POSSIBI K 5It{ XI'IGHAPIIIC SI:Qi I NCK U I 5t)3II{ MAMMALIAN FAt NAS .\[" Fill' PI IO( I XI2-PI KIST()CI N[ BOLNDARY IN MIDDI K .\ND \kESFF }IN I [ROI'I

TABLE I

80

H. TOBIEN

named locality with its local fauna represents the type of the niveau ( = "gisementtype" of THALER, 1966, p.186), as it shows all or most of the criteria typicalfor this niveau. The second locality or local fauna gives further criteria for the niveau. I f a subdivision of a niveau ( = subniveau, sublevel) should be possible, the second abbreviation could be used to typify this subniveau (e.g., A r - T e subniveau means: subniveau of the Ar-Se niveau, characterized by the local fauna of Tegelen, The Netherlands). The niveaus, briefly described on the following pages ~ are located between the Pontian sensu lato ( = Lower Pliocene) and the Biharian (Middle Pleistocene) of KRETZOI (1965, p.607). The earlier niveaus belong to the Late Pliocene, the younger ones to the Early Pleistocene. Altogether six, possibly only five niveaus result (Table I). Besides the typical local fauna each niveau is exemplified by several other local faunas, which are neither complete nor fully typical in every case, and they may be changed and improved. THE MAMMALIAN F A U N A NIVEAUS IN MIDDLE A N D WESTERN EUROPE

Brief characterization of the fauna niveaus (see Table I) The classical fauna from the Roussillon area in southeastern France (HuGuENEY and MEIN, 1966) is typical for the Rou-A1 niveau. This fauna is definitely younger than the Hipparion faunas of the Pontian sensu lato; Dicerorhinus megarhinus and Trilophomys are typical, but Mimomys, other primitive arvicolids, and "microtoid" cricetines are absent, as are also members of the E - L - E group since they are very probably later immigrants to western and middle Europe. In Alcoy (Province of Alicante, Spain) micromammals were discovered some years ago (HARTENBERGERet al., 1967, p.511). Trilophomys, Ruscinomys and Stephanomys occur, yet the last two on a lower evolutionary level than in the Roussillon fauna. This gives the Alcoy local fauna a slightly older aspect. The Weze (Poland, KOWALSKI, 1964) and Csarnota (Hungary, KRETZOI, 1962) local faunas, which are typical for the We-Csa niveau are characterized by eastern immigrants such as: (1) primitive Mimomys species belonging to group I of FEJVAR (1964, p.104); and (2) other archaistic arvicolids and "microtoid" cricetines such as Baranomys, Germanomys, Stachomys, " Ungaromys", Promimomys, Leukaristomys, Laugaritiomys. Trilophomys is another microtoid cricetine genus, but it already appears - - very probably also as an eastern immigrant - - in the earlier Rou-A1 niveau. Besides Trilophomys W61fersheim (north of Frankfurt, western Germany; TOmEN, 1952) has produced Mimomys forms of group I as well as members of the Baranomys-Germanomys group, and also large mammals. Some taxa among them

1 More details about the niveaus and their local faunas are given in TOBIEN(1969).

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MAMMALIAN FAUNAS AT THE PLIOCENE PLEISTOCENE BOUNDARY

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(e.g., ursid, tapir, suid) are somewhat more advanced than the earlier species from the Roussillon. Rou-AI niveau and We-Csa niveau together correspond to THALER'S(1966) "Zone de Perpignan". For the Rou A1 niveau KREtZOfS stage name Ruscinian. (1962, p.375), and for the We-Csa niveau his Csarnotanum (p.376; linguistically perhaps simpler: = Csarnotian = Csarnotium = Csarnotien = Csarnotiano = Csarnotiense) may be used (Table !). The Aj niveau is represented by Ajnacska (FEJFAR, 1964). One main difl'erence between this and the We-Csa niveau seems to be indicated by evolved Mimerays species with cement-filled synclines and a higher degree of hypsodonty ~ts in FEJFAR'S (1964, p. 105) group II. Mhnomvs species of group II are also represented in Rebielice (Poland; KOWALSKI, 1964, p.87). Among the larger mammals of Ajnacska members of the E L E group are missing. This might be a lack of knowledge, but it might also be due to the fact that Ajnacska is earlier than the immigration of members of the E L E group. At the moment l prefer the latter interpretation until taxa of the E L E group are discovered either there or in contemporaneous local faunas. In this case the Aj niveau must be cancelled and its local faunas should be classified in the next younger Et-Mo niveau. Most of the local faunas belonging to the Et Me niveau include at least one taxon of the Elephas Leptobos-Equus group, with the exception of Vb~lette: the cervids, however, are identical with those from Etouaires (HEINTZ, 1967, p.544, 1968). As in the earlier niveaus, the rhinocerid is Dicerorhimts megarhimts. The Mimomys species, as far as they are known, can be referred to Fejfar's group II. Trilophomys and the Baranomys-Germanomys group have obviously disappeared in the Et-Mo niveau. Anancus arvernensi3 and Mammut borsoni occur as holdovers ("superstites") from the earlier niveaus, just as Hipparion does in eastern local faunas. Representative for this niveau are the local faunas of Etouaires (French Central Massif, SCHAUB, 1941) and Montopoli (Val d'Arno Inferiore, Italy, AZZAROLI, 1963). The study of the cervids by HEINTZ (1967, 1968) contributes broadly to the characterization of the E t - M e niveau and the following Re Va and Ar Se niveaus, respectively. Since members of the E - L - E group in the local faunas of the Et Me niveau occur and as these, according to HAUG (1911, p.1767) characterize the Villafranchian (see p.86), it would be convenient, to locate the E t - M o niveau - - as Lower Villafranchian - - at the base of the whole VillalYanchian. The Villafranchian, so understood, would correspond to the Villanyum of KRETZOI (1962, p.377). The local faunas typical for the Ro-Va niveau are Roccaneyra (French Central Massif) and St. Vallier (southern Rh6ne Valley, France). Both have always been considered younger than Etouaires and Vialette (V~RET, 1954, pp. 175 176: Palaeo~eoLTraph.v, PalaeocHmatol., Pa&eoecol., 8 (1970) 77 93

82

H. TOBIEN

BOUT, 1960, p.169). Dicerorhinus etruscus now has taken the place of the former D. megarhinus, and Archidiskodon meridionalis, obviously in a slightly more advanced form than in the E t - M o niveau, is widespread. The cervids are different from those of the earlier niveau. The Mimomys species still belong to group lI. The Ro-Va niveau can be termed Middle Villafranchian. KURTI~N (1963) and particularly AZZAROL~ (1967) delivered more details as far as the larger mammals of the Middle Villafranchian are concerned. The ecological change from the E t - M o niveau to the Ro-Va niveau (i.e., from Lower to Middle Villafranchian) is remarkable in the local faunas of western Europe. Etouaires and Vialette are woodland faunas as most of the local faunas of the Ruscinian and Csarnotian are, whereas in Roccaneyra, Pardines (French Central Massif) and Villaroya (province of Logrofio, Spain) steppe/savannah elements are dominant, among them mainly antelopes (KURT~N, 1963, p.14, fig.l; 1968, fig.6). This also holds true for some local faunas of the following Ar-Se niveau ( = Upper Villafranchian). In a similar way as the steppe/savannah faunas of southern and southwestern Europe received their character from the east during Pontian time, i.e., from the large central Asiatic or southern palearetic steppe/desert area (TOBIEN, 1956, p.209), the steppe/savannah phase of the Middle and Upper Villafranchian can also be explained by the westward extension of this steppe areal into western Europe which, however, was influenced before and afterwards by atlantic-humid conditions (for more details see TOBIEN, 1971). The Ar-Se niveau is typified by the local faunas of the upper part of the Val d'Arno Superiore ( = Montevarchi Series, Italy) and Sen6ze (French Central Massif). For the stratigraphic position and the faunistic characteristics of the larger mammals of these and other local faunas of the Ar-Se niveau I refer to AZZAROLI (1967). Besides Mimomys species of Fejfar's group II members of his group III and other more modern taxa such as Clethrionomys, Pliomys episeopalis have already been encountered in some places, for instance at Schernfeld (Bavaria, Western Germany; HELLER, 1967, p.203). Faunas with these elements may be transitional to KRETZOI'S (1965) Biharian (early Middle Pleistocene). Also Arehidiskodon meridionalis seems to be represented in more advanced populations. A subdivision of the Ar-Se niveau, into two or perhaps three subniveaus is obviously possible (AzzAROLI, 1967; HEINTZ, 1968, p. 2185).

Relations to marine stages (see Table I) Stratigraphic relations of marine stages to the mammalian biostratigraphic niveaus outlined in the preceding section can be found in a relatively favourable manner in the Mediterranean region: the Montpellier mammalian remains were imbedded in marine Astian sands. The Roussillon fauna has been found in brackish and fresh-water marls immediately above marine Astian deposits. The Rou-A1 niveau therefore would belong - referred to the marine stratigraphic scale - - to the Astian. Palaeogeography, Palaeoclimatol.,Palaeoecol., 8 (1970) 77 93

MAMMALIAN FAUNAS AT THE PLIOCENE-PLEISTOCENE BOUNDARY

83

Even higher niveaus must be placed, as it seems, into this marine stage: the local fauna of Montopoli (Val d'Arno lnferiore) has been encountered in marine deposits, which are characterized by their molluscs as Pliocene, i.e., Astian (AzzAROL1, 1963, p.15; 1967, p.11). Apparently at the present state of knowledge the marine Astian fauna, as defined by their molluscs, does not change its character from the Middle Pliocene (Rou-A1 niveau) to the Lower Villafranchian ( E t - M o niveau). In the local mammalian faunas of the Lower Villafranchian members of the F - L E group occur as in Etouaires: Leptobos, and in Montopoli: Archidiskochm meri~#o,alis. The International Geological Congress (London, 1948) h~s recommended that the Lower Villafranchian be placed in the Pleistocene. On the other hand, in the frame of marine stratigraphy the Astian is a stage of the Pliocene. Therefore the marine Astian still extends into the Pleistocene, as far as it is defined by continental mammals. So the Pliocene-Pleistocene boundary in the marine facies must be located stratigraphically higher than the same boundary based on the change of continental mammals and on the recommendation of the I.G.C. (London, 1948), which both locate this boundary comparatively lower. Similar discordancies are also known from other boundaries of the Tertiary divisions such as Miocene-Pliocene, Oligocene Miocene and Eocene Oligocene, so Far these boundaries are established in continental facies (by mammals) and in marine facies (by molluscs, microorganisms etc.), respectively. The later Villafranchian niveaus should correspond to the marine Calabrian. These correlations, however, are not fully certified, as local mammalian faunas of the Ro Va and Ar Se niveaus are missing up to now in a /ithostratigraphic connection with Calabrian deposits in the Mediterranean area (see also AZZAROLI, 1967, p. 12). Concerning the correlation of Villafranchian mammal faunas and marine deposits of the North Sea along the British east coast (Crag Series) I refer to AZZAROH (1967, p. 9) and BOUT (1968, p.57). According to these authors elements of the Middle Villafranchian (Ro Va niveau), are probably represented in the Red Crag, whereas elements of the Upper Villafranchian occur in the Norwich Crag and Weybourne Crag of East Runton (Ar-Se niveau). As far as is known the marine molluscs associated with these local faunas are difficult to correlate with those of the Mediterranean area. A correlation, however, might be possible through the mammalian faunas after a more precise knowledge of their biostratigraphic position.

Reh~tions to.l%ristic stages (see Table 1) This section only intends to demonstrate relations to local floras as far as they are in a lithostratigraphic connection with mammalian local faunas or have been found together. According to K1RCHHEIMEr (1940, pp. 151, 159) and LESCHrK (1956) the local flora of Walfersheim (north of Frankfort, Western Germany) must be parallelized Palaeo,~eo,~,raphy, Palaeoclimat~l., Palaeoecol., 8 (1970) 77 93

84

H. TOBIEN

by its carpological and palynological components with the Reuverianin The Netherlands. The We-Csa niveau, therefore, can be compared with the Reuverian. It is the unanimous opinion of the paleobotanists, that the Reuver flora still has a definite Pliocene character. The other end of the niveau sequence proposed here offers a further possibility for a connection: The mammalian local fauna of Tegelen (The Netherlands) is associated with a local flora, in which a certain number of typical Tertiary elements are missing. The Tegelen flora already has a Pleistocene, interglacial charactel. A similar flora has been found in the upper sections of the Val d'Arno Superiore, i.e., in beds above the discordancy ( = Montevarchi Series), which contain the classical rich mammalian fauna, the beds below the discordancy (Castelnuovo dei Sabbioni Series), on the other hand, have produced a Reuverian flora (FLORSCHOTZ, 1954; BOUT, 1968, p.65). From the Le Puy Basin (French Central Massif) pollen floras are known that have been referred to the Tiglian and Pretiglian respectively (BouT, 1968, p.65). Typical Reuverian floras, connected with mammalian faunas, in general seem to be lacking in the French Central Massif so far. According to the paleobotanical data, the earliest Plio-Pleistocene mammalian local fauna in the Le Puy area (Vialette) seems to have lived together with a flora that no longer had a Pliocene, i.e., Reuverian character, but with a younger, Pleistocene vegetation. A comparable situation seems to hold true for the Etouaires local fauna in the adjacent Allier Basin: here, in direct stratigraphic superposition, a flora has been discovered which is identical with the Ceyssac local flora in the Le Puy Basin and which cannot be ascribed to the Reuverian, but to a later floristic level (BouT, 1968, p.66). The mammalian remains from the stratigraphically lower lignites in the Leffe section (province of Bergamo, north Italy; VENZO, 1965, p.378, fig.4) occur in an interstadial which must be characterized as moderate-cool according to the palynological investigations of F. Lona (see VZNZO, 1965). VENZO (1965, p.378) compares this horizon with the Pretiglian. Very probably it could correspond to our Ro-Va niveau ( = Middle Villafranchian). From the upper parts of the Leffe section Archidiskodon meridionalis forma evolutiva and Dicerorhinus etruscus are cited (VENzO, 1965, fig.4). The palynological data from this horizon (F. Lona in VENZO, 1965) indicate a relationship to the Tiglian, and through this to the Tegelen mammalian local fauna ranked in the Ar-Se niveau ( = Upper Villafranchian). According to the preceding data the Reuver flora of W61fersheim and of the We-Csa niveau is already replaced in the Et-Mo niveau ( = Lower Villafranchian) by local floras of Tiglian or Pretiglian habitus that characterize furthermore the plant associations of the two following niveaus.

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MAMMALIAN FAUNAS AT THE PLIOCENE PLEISTOCENE BOUNDARY

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Radiometric data BOUT et al. (1966), BOUT (1968) and SAVAG~ and CURTIS (1968) published K - A r data of mammalian local faunas from the French Central Massif. These data are furthermore important for the knowledge of the duration of the Pleistocene, especially of its lower part. B o u t (1968, p.68) issued the following figures: Etouaires is later than the basaltic lava l¥om Roca Neyra, dated as 3.9 m.y., and earlier than the lowermost tuffilic layer of the Perrier section, dated as 3.5-3.4 m.y. The figure 3.9 was determined by G. H. Curtis. H. J. Lippolt (in B o u t et al., 1966) found only 3.1 for the same rock. The local fauna of Roccaneyra was deposited in a tuffaceous agglomerate with an age of 2.5-2.6 m.y. The local faunas of Coupet, Chilhac, Crozas of the latest Villafranchian delivered absolute ages of 1.8 m.y. Also younger than 2 m.y. must be the latest Villafranchian faunas of Sinzelles and La MaIouteyre in the vicinity of ke Puy (French Central Massif). In SAVAGE and CURTIS (1968, p.15) the following data can be fonnd: The sanidine-bearing tuff immediately above the layer with the mammalian fauna of Etouaires has an age of 3.4 m.y. A basalt at the base of the section near the village of Roca Neyra, corresponding lithostratigraphically to the Etouaires horizon, produced 3.5 m.y. This is an obviously corrected figure, compared to the earlier value of 3.9 in BOUT (1968). The sanidine from a b o m b in the volcanic ash with the mammalian fauna of Roccaneyra is 2.5 m.y. old. The lava that overlies the local fauna of Mont Coupet near Le Puy gave 1.92 m.y. From the preceding values results a duration of the Villafranchian (i.e., from the E t - M o niveau to the Ar Se niveau) from about 3.4-3.5 m.y. to 1.8-.I.9 m.y., which means an absolute time span of 1.5 1.7 m.y. (see Table I). The time succession resulting from the K - A r data is lithostratigraphically controlled, at least for the Etouaires and Roccaneyra local faunas. Furthermore, the absolute figures confirm the relative ages received by the paleomammalogical investigations for at least those local faunas which have produced absolute datations. It wotfld also be interesting to have absolute datations from the earlier niveaus, which are here placed in the Middle and Upper Pliocene. As let as Ajnacska is concerned a radiometric age should be obtainable from the volcanic ashes in the section of the site. In any case, the time interval between the Et Mo niveau (typified by Etouaires with 3.4 3.5 m.y.) and the upper boundary of the Pontian (about 6.5 m.y.; EVERNDEN et al., 1964, p.167; D. g. Savage, personal communication, 1965), which in this paper is understood as Middle and Upper Pliocene (see Table 1), is obviously longer than the time span co~ering the three niveaus of the Villafranchian with 1.5-1.7 m.y.

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The Pliocene-Pleistocene boundary As far as the Pliocene-Pleistocene boundary, based on the succession of continental mammalian faunas, is concerned, HAUG'S old definition (1911, p. 1767) must be considered first of all. It emphasizes the importance of Elephas (now Archidiskodon meridionalis and related forms), Bos (now Leptobos) and Equus (now diverse Equus taxa) for the Pleistocene and its lower limit. Furthermore Haug placed the Villafranchian, defined by early members of these eastern immigrants, at the beginning of the Pleistocene. This definition gave, at least for Europe, a rather practicable Pliocene-Pleistocene boundary established on paleomammalogical criteria. It is an open question, however, whether the three Villafranchian genera contemporaneously appeared as immigrants in Europe. KURT~N (1963, p.4) stressed, that Leptobos would have arrived in Europe earlier than Archidiskodon and Equus. On the other hand, in the stratigraphically old Montopoli local fauna (Val d'Arno Inferiore) Leptobos is absent, whereas an archaistic Archidiskodon meridionalis is present (AzzAROLI, 1963, p.18). Also in the Rumanian Lower Villafranchian faunas (M~lu~teni, Bere~ti and others) Leptobos is missing and instead a Bison-like form occurs. Arehidiskodon is also absent and appears only in the Middle Villafranchian. In the Lower Villafranchian, however, two Equus forms are listed by SAMSON and RADULESCO (1965, pp.69-70). Until the question is definitely settled it seems to be reasonable, in this somewhat obscure situation to follow HOPWOOD (1935, p.47): "that these three genera form convenient guide fossils for the separation of the Pleistocene from the Pliocene, and that the presence of any one of them is sufficient reason for assigning a Pleistocene age to the bed in which it is found". Since members of the E - L - E group are present in the local faunas of the E t - M o niveau and since the PliocenePleistocene limit must be drawn below this niveau, the earlier niveaus must thus belong to the Pliocene (Table I: A-boundary). The E t - M o niveau has been ascribed to the Lower Villafranchian. This procedure basically follows Haug's definition, according to which the Villafranchian is characterized by the presence of the E - L - E genera, supplied by the above mentioned statement of Hopwood. Concerning the type locality of the ViUafranchian near the town of Villafranca d'Asti (north Italy) it seems to me to be not yet proven, that beds containing one or two or all three elements of the Villafranchian actually do not occur in the environs of this city, and that the stratotype of Pareto's Villafranchian must be cogently referred to strata with a fauna of Pliocene character (for more details see TOBIEN, 1969). Whether the lower boundary of the Villafranchian and the Pleistocene, as defined above, agrees with the recommendations of the I.G.C. (London, 1948) is another problem. In its pertinent part the recommendation is as follows: "The Commission recommends that, in order to eliminate existing ambiguities, the Lower Pleistocene should include as its basal member in the type-area the Calabrian

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formation (marine) together with its terrestrial (continental) equiwflent the Villafranchian. The Commission notes that according to evidence given this usage would place the boundary at the horizon of the first indication of climatic deterioration in the ltalian Neogene succession". On p.83 it has been demonstrated that the local fauna of Montopoli (Val d'Arno I nferiore, Italy) with Archidiskodon meridionalis is contemporaneous with marine sediments not of Calabrian but obviously of Astian age. As the marine Astian molluscan fauna does not show any sign of climatic deterioration, the E t - M o niveau ( = Lower Villafranchian), to which the Montopoli fauna must be ascribed, should have its position below the Pliocene-Pleistocene boundary. Only the R o - V a niveau - - as it is accepted here with the Calabrian as its rnarine equivalent - - would agree with the recommendations of the I.G.C. (London, 1948), because the Calabrian indicates a first climatic deterioration. Accordingly the Pliocene Pleistocene boundary must be drawn between the E t - M o niveau and the R o - V a niveau, i,e., between the Lower and Middle Villafranchian (Table I: B-boundary). A similar view is advocated for other reasons by AZZARO1A ( 1 9 6 7 ) , The position of the B-boundary is not very far from that proposed by GRICHUK et al., (1965, p.314). Herein the Lower ("warm") Villafranchian without Dicerorhinus etruscus and Hippopotamus major, however, could agree with our Lower Villafranchian. SAVAGE and CURXlS (1968) propose a PIiocene Pleistocene boundary through radiometric data at 3 m.y., since the earliest glacigene deposits on the Northern Hemisphere (Sierra Nevada) have been dated at 2.7 m.y. and similar occurrences on the Southern Hemisphere are dated somewhat younger at 2.5 m.y. This radiometric boundary should fall approximately between the E t - M o and Ro-Va niveaus, i.e., between Middle and Lower Villafranchiam corresponding to the B-boundary on Table I. It very probably coincides with the peculiar ecological change which is found in mammalian local faunas of western Europe: in the Lower Villafranchian woodland faunas (Etouaires is typical) prewtil, whereas in the Middle Villafranchian savannah/steppe elements are predominant. The A-boundary is a time-marker which can stratigraphically rather well be characterized by mammals. This is also the case to a lesser degree with the B-boundary. The B-boundary, however, is defined by absolute data and is further characterized by climatic and ecological phenomena. Thus it would agree better with the recommendations of the I.G.C. (London, 1948). In addition the B-boundary very probably seems to be the correlate of the boundary between Astian and Calabrian. The B-boundary separates two series of the stratigraphic scale: Pliocene and Pleistocene, but cuts through a stratigraphically lower unit, the Villafranchian Stage, since the Lower Villafranchian belongs to the Pliocene, and the Middle and Upper Villafranchian to the Pleistocene. This should be avoided. Therefore the name Lower Villafranchian ( = E t - M o niveau) could be changed into Upper Palaeogeograph.v, Palaeoclimatol., Palaeoecol., 8 (1970) 77 93

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H . TOBIEN

Csarnotian ( = late Upper Pliocene). In this case the Middle Villafranchian would become Lower Villafranchian, whereas the name Upper Villafranchian would stay as it is. Many detailed studies seem to be necessary before a decision in favor of one of the two boundaries can be made. Under the present state of knowledge the B-boundary seems to me - - together with AZZAROLI (1967), SAVAGE and CURTIS (1968) and other paleomammalogists - - to be the more suitable. R E L A T I O N S T O T H E M A M M A L I A N F A U N A S OF T H E E U R O P E A N P A R T OF T H E U . S . S . R .

Just as in middle and western Europe the knowledge of the mammalian faunas at the Pliocene-Pleistocene boundary has also been considerably increased in the last two decades in the European and Asiatic parts of the U.S.S.R. Reviews with the pertinent literature are given by NIKIEOROVA (1965, 1968). Furthermore GABOUN~A (1967) has contributed to this subject. During the "International Colloquium on Lower and Middle Pleistocene Geology and Fauna of Europe, M a y - J u n e 1969" under the direction of Prof. Dr. K. V. Nikiforova in Moscow, I received many suggestions and much informations through the lectures and the discussions with the Russian and foreign colleagues to all of whom I wish to confer my best thanks. In the subsequent section an attempt will be made to correlate the above proposed succession of middle and western European mammalian fauna niveaus with those in the European part of the U.S.S.R. (Table II). As far as the sequence of local faunas in the U.S.S.R. is concerned I refer to NIKIFOROVA (1965, 1968). The fauna of the Moldavian Roussillon is very similar to the Roussillon Montpellier fauna of southern France. Anancus arvernensis, Dicerorhinus mega-

rhinus, Hipparion crassum, Propotamochoerus provincialis, Capreolus australis, Parabos boodon, Dolichopithecus ruscinensis and others are common. Together with the larger mammals archaic, rooted microtines without cement which are lacking in the typical Roussillon fauna occur, such as Dolomys milleri, Pliomys kowalskii, Mimomys (Cseria) gracilis, Mimomys ex gr. stehlini and others (NIKIFOROVA, 1965, p.240). Mimomys (Cseria) gracilis and M. ex gr. stehlini are species of Fejfar's group I, as they also occur in the We-Csa niveau. In the reviews of the Moldavian Roussillon fauna NIKIFOROVA (1965, p.240) mentions Equus and an archaistic Archidiskodon meridionalis ( = A. gromovi). Seen from the situation in middle and western Europe it might be possible, that these representatives of the E - L - E group indicate a younger part within the Moldavian Roussillon faunistic complex, which corresponds more or less to the E t - M o niveau. To ascertain this comparison it would be necessary, however, to demonstrate at those localities the occurrence of microtines of Fejfar's group II, together with elements of the E - L - E group. Sites of the Moldavian Roussillon faunistic complex are: the Kutchurgan

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TABLE II M A M M A L I A N F A U N A S E Q U E N C E A T T H E P L I O C E N E - - P L E I S T O C E N E B O U N D A R Y IN M I D D L E A N D "vVESTERN E U R O P E A N [ ) IN T H E E U R O P E A N P A R T OF T H E U . S . S . R .

Stages

Mammalian niveaus in middle and western Europe

Europeanpart of the U.S.S.R. 1

CaspianSea correlations 1

Villafranchian

Ar-Se niveau

Taman complex

Apsheronian

Ro-Va niveau

Chapry complex

Middle and Upper Aktshagylian

.

.

.

.

EI-Mo niveau

B-boundary later part of the MolLower Aktshagylian davian Roussillon fauna (with Equus, Archidiskodon)

....................................................... Csarnotian

A-boundary

'? Aj niveau

?

?

We-Csa niveau

Odessa catacombs; earlier part of the Moldavian Roussillon fauna (with primitive

Lower Aktshagylian

Mimomys)

Ruscinian

Rou AI niveau

'? earliest parts of '? the Moldavian Roussillon fauna

1 Afler NIKIFOROVA(1965, 1968). Gravel o f M o l d a v i a (eventually representing the earlier phase of" the complex); the basins o f the lower Pruth, Saltscha and K a g u l (in parts also belonging to the earlier phase o f the complex) and the K o t l o w i n a village site (at Lake Kagul), with Archidiskodon meridionalis ( = A. gromovi). This last site can p r o b a b l y be ascribed to the y o u n g e r part o f the M o l d a v i a n RoussiUon fauna. To the We Csa niveau m a y be referred f u r t h e r m o r e the f a u n a f r o m the fissure fillings in the f a m o u s c a t a c o m b s o f O d e s s a (ORLoV, 1967; NIKIFOROVA, 1969, p.52). The larger m a m m a l s show a Late Pliocene character, and m e m b e r s o f the E - L - E g r o u p are missing. A m o n g the r o d e n t s such as l a g o m o r p h s , cricetines, Steneol~'ber and others, microtines also occur; e.g., Dolomys is mentioned in the faunal lists. D u r i n g the excursion o f the " I n t e r n a t i o n a l C o l l o q u i u m on Lower and M i d d l e Pleistocene G e o l o g y and F a u n a o f E u r o p e " in Odessa, June 1969, I had the o p p o r t u n i t y - - t h a n k s to the courtesy o f Prof. Dr. I. Y. Yatsko, D i r e c t o r o f the Paleontological M u s e u m o f the University o f Odessa, and one of Palaeogeography, Palaeoclimatol., Palaeoecol., 8 (1970) 77 ~93

90

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the well known experts of this fauna - - to study briefly the microtine remains in the museum's collections. Among the lower M1 there were three specimens which can be referred to one of the Mimomys species of group I (crowns with well developed roots, without cement in the synclines, with Mimomys ridge, one specimen with an enamel islet in the anterior lobe). The size indicates one of the larger species of group I. Three other M1 inf. are different. They have strong roots, are without cement, and the wear facets resemble Germanomys or a closely related genus. The archaistic microtines, the Late Pliocene character of the larger mammals, and the absence of members of the E - L - E group would favour a correlation with the W e - C s a niveau (Table I1). Yet the ecological conditions are markedly different from those in the local faunas of the same age in western Europe. ORLOV (1967, p.556) underlined the high percentage of steppe inhabitants in the catacomb fauna, such as Paracamelus (commonest animal), Cricetulus, Proochotona, Ochotona, Hystrix, Struthio, Ammoperdix and others, besides woodland and marshy woodland elements. Steppe elements to such an extent are missing in the western European sites. A representative of the Aj niveau cannot be recognized. In view of its uncertain character, which makes it possibly identical with the E t - M o niveau (see p.81), the absence of the Aj niveau in the Russian sequence does not seem to be surprising. The next younger faunistic level in the European U.S.S.R. is the Chapry complex. Besides Anancus arvernensis, Archidiskodon meridionalis and Equus stenonis, Dicerorhinus etruscus already occurs, whereas in the Moldavian complex Dicerorhinus megarhinus is still present, as also in the We-Csa and E t - M o niveaus. A m o n g the microtines Dolomys hungaricus and Mimomys praehungaricus are mentioned besides superstites from the preceding Moldavian complex such as Dolomys milleri, and Mimomys cf. stehlini. The Chapry complex can therefore be correlated with the R o - V a niveau. Localities are the Chapry Beds near Rostov on the Don, the Jergeni Beds of the Sal River, the lower Kujalnik Beds near Odessa, and the high river terraces of southern Moldavia and southwest Ukraine (e.g., Ferladany terrace IX of the Dnjestr). The Chapry complex is followed by the T a m a n complex. In this, an evolved Archidiskodon meridionalis occurs, occasionally accompanied by Anancus arvernensis, furthermore Equus siissenbornensis and Paracamelus. Typical representatives of the rodents are the advanced microtines Mimomys reidi, Lagurodon arankae, Allophaiomys plioeaenicus (NIK~FOROVA, 1968, p.370). NIKIFOROVA (1965, p.241) compares the T a m a n complex with Sen6ze, Tegelen and Upper Val d'Arno. It might be therefore parallelized with a rather late part of our Ar-Se niveau. The main localities of the T a m a n complex are situated along the coast of

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the Asow Sea, near Kairy on the Dnjepr, in the Pruth and Danube valleys and in the environments of Odessa. NIKIFOROVA (1965, p.241; 1968, p.369) emphasizes the more continental character of the Moldavian Roussillon local faunas (camels, ostriches, tortoises, and in the earlier parts a numerical dominance of the lagomorphs among the micromammals) compared with western Europe. Similar observations were made by ()RLOV (1967) for the Odessa catacombs fauna (see p.90). The local faunas of the Chapry and T a m a n complexes also indicate a more continental climate compared to western Europe. As was pointed out on p.82 it is the influence of the large central Asiatic or southern palaearctic steppe region, existant since Neogene time and may be earlier, which is probably responsible for these ecological differences. The farther to the east, the more distinct these differences become evident, particularly in context with the expansion of steppe/savannah conditions to western Europe during Middle Villafranchian times (for more details see TOBIEN, 1971). As far as the marine transgressions of the Caspian Sea are concerned, N1KIFOROVA (1965, p.242, 1968, p.370) parallelizes the faunal sequence 4(1) earlier part of the Moldavian Roussillon complex; (2) later part of the Moldavian complex; (3) Chapry complex; (4) Taman complex - - as follows (Table II): (l) and (2) with the Lower Aktshagylian; (3) with the Middle and Upper Aktshagylian; and (4) with the Apsheronian, and she continues: "'From this results the probability, to compare the marine Aktshagylian deposits not only with the Calabrian (Middle and Upper Aktshagylian) but also partly with theAstian beds (LowerAktshagylian) of the Mediterranean area and the Atlantic. The upper horizons of the Calabrian corresponds already to the Apsheronian of the Caspian Basin" (N1KIFOROVA, 1965, p.242). These statements would not contrast to the parallelizations with western Europe: the Montopoli local fauna of the E t - M o niveau is associated with a marine Astian mollusc fauna (see p.83 and Table I). As is suggested above, the E t - M o niveau can be correlated with the Later Moldavian Roussillon complex, which is referred to the Lower Aktshagylian by Nikiforova. The earlier Moldavian Roussillon complex corresponds to the middle and western European We-Csa niveau, that per se must have Astian as a marine equivalent, since it is overlain and underlain by niveaus which have been referred to the Astian (Table I). The R o - V a niveau, which is compared with the Chapry complex, might have the Calabrian of western Europe as a marine equivalent. The Taman complex, which Nikiforova connects with the Apsheronian presents resemblances with the Ar-Se niveau, which very probably has a later Calabrian as a marine equivalent (Table II) 1. t See also the similar, but in some points differing views of GABOUNIA (1967).

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ACKNOWLEDGEMENTS

The author extends his best thanks to Mrs. Hartenstein-Day, Seeheim, who scrutinized and improved the English of this note. REFERENCES AZZAROLI, A., 1963. Rinoceronti pliocenici del Valdarno inferiore. Palaeontographia Italica, 57: 11-20.

AZZAROL:, A., 1967. Villafranchian correlations based on large mammals. Congr. on Mediterranean Neogene, 4th, Bologna, 1967:15 pp. BOUT, P., 1960. Le Villafranchien du Velay et du Bassin Hydrographique Moyen et Sup~rieur de l'Allier. Jeanne d'Arc, Le Puy, 344 pp. BouT, P., 1968. La limite Pliocene-Quaternaire en Europe occidentale. Bull. Assoc. Franc. Quaternaire, 1: 55-78. BOUT, P., FRECHEN,J. et LIPPOLT, H. J., 1966. Datations stratigraphiques et radiochronologiques de quelques coul6es basaltiques de Limagne. Rev. Auvergne, 80(4): 207-231. EVERNDEN, J. F., SAVAGE, D. E., CURTIS, G. H. and JAMES, G. Z., 1964. Potassium-Argon dates and the Cenozoic mammalian chronology of North America. Am. J. Sci., 262: 145-198. FEJEAR, O., 1964. The Lower Villafranchian vertebrates from Hajmicka near Fil~ikovo in southern Slovakia. Rozpravy Ustred. Ust. Geol., 30:115 pp. FLORSCHOTZ, F., 1954. Le changement dans l'aspect des for6ts ~t la transition du Plioc~ne au Pldistoc~ne en France mdridionale, compard avec celui constatd en Toscane et en Hollande. Actes Congr. Intern. Botan., 8me, Paris, 1954: p.249. GABOUNIA, L., 1967. Sur la signification des faunes de mammif~res du Miocene Supdrieur et du Plioc~ne de la region Ponto-Caspienne. Congr. on Mediterranean Neogene, 4th Bologna, 1967:13 pp. GRICHUK, V. P., HEY, R. W. and VENZO, S., 1965. Report of the subcommission on the PlioPleistocene boundary. INQUA Rept. Intern. Congr. on Quaternary, 6th, Warsaw, 1961, 1: 311-329. HARTENBERGER, J. L., MICHAUX, J. et THALER, L., 1967. Remarques sur l'histoire des rongeurs de la faune ~ Hipparion en Europe sud-occidentale. Coll. Intern. Centre Natl. Rech. Sci., 163: 503-513. HAUG, E., 1911. Trait~ de Gdologie, 2. Les PHiodes G~ologiques. Colin, Paris, 868 pp. HEDBERG, H. O., 1961. Stratigraphic classification and terminology. Intern. Geol. Congr., 21st, Copenhagen, 1960, Rept. Session, Norden, 25:38 pp. HEINTZ, E., 1967. Donndes prdliminaires sur les cervidds villafranchiens de France et d'Espagne. Coll. Intern. Centre Natl. Rech. Sci., 163: 539-551. HEINTZ, E., 1968. Principaux rdsultats systdmatiques et biostratigraphiques de l'dtude des cervid6s villafranchiens de France et d'Espagne Compt. Rend., 266: 2184-2186. HEELER, F., 1967. Die Wiihlm~iuse (Arvicolidae Gray 1821) der altpleistoz:inen S~iugetierfauna von Schernfeld bei Eichst~tt in Bayern. Mitt. Bayer. Staatssamml. Pali~ontol. Hist. Geok, 7: 201-203. HoPwooD, A. T., 1935. Fossil elephants and man. Proc. Geologists' Assoc. Engl., 46: 46-60. HUGUENEY, M. et ME/N, P., 1966. Les rongeurs Plioc~nes du Roussillon dans les collections Lyonnaises. Trav. Lab. G~ol. Fac. Sci. Lyon, 13: 243-266. KIRCHHEIMER, F., 1940. Flora und Gliederung des Plioc/ins in Mitteleuropa. Zentr. Mineral. Geol. Paliiontol., 1940B: 141-167. KOWALSKI, K., 1964. Palaeoecology of mammals from the Pliocene and Early Pleistocene of Poland. Acta Theriologica, 8: 73-88. KRETZOI, M., 1962. Fauna und Faunenhorizont yon Csarn6ta. Ann. Rept. Hung. Geol. Inst., 1959: 344-395. KRETZOI, M., 1965. Die Nager und Lagomorpben von Voigtstedt in ThiJringen und ihre chronologische Aussage. Paliiontol. AbhandL Ser. A., 2: 586-660.

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KURTI~N, B., 1963. Villafranchian faunal evolution. Soc. Sci. Fennica, Commentationes Bhd., 26: 1-18. KURTI~N, B., 1968. Pleistocene Mammals o f Europe. Weidenfeld and Nicolson, London, 317 pp. LESCttIK, G., 1956. Die Entstehung der Braunkohle der Wetterau und ihre Mikro- und Makroflora. Palaeontographica, B, 100(1/3): 26-64. NIKIFOROVA, K. V., 1965. Stratigraphische Equivalente des Villafranchiens in der Sowjetunion. Koninkl. Ned. Akad. Wetensehap., Proc., Ser. B., 68: 237-248. NIKIFOROVA, K. V., 1968. Die Korrelation der unter- und mittelpleistozfinen Ablagerungen im niSrdlichen Eurasien. Bet. Deut. Ges. Geol. Wiss,, Ser. A, Geol. Paliiontol., 13: 367-374. NIKIVOROVA, K. V. (Editor), 1969. International Colloquium on Lower and Middle Pleistocene Geology and Fauna o f Europe; Guide-Book. Nauka, Moscow, 57 pp. ORL<:,V, J. A., 1967. Quelques donn6es sur les vertdbr6s du Plioc,~ne supdrieur d'Odessa. Coll. Intern. Centre Natl. Rech. Sci., 163: 553-556. SAMSON, P. M. und RADULESCO,C., 1965. Die S~iugetierfaunen und die Grenzen Pliozfin/Pleistoz~in und Unterpleistozfin/Mittelpleistozfin in Rum~inien. Ber. Geol. Ges. Deut. Demokrat. Rep. Gesamtgebiet Geol. Wiss., 10: 67-76. SAVAC;E, D. E. and CURTIS, G. H., 1968. The Villafranchian Stage-Age and its radiometric dating. Proc. Soe. Econ. Paleontologists Mineralogists Res. Comm.--Syrup. on Radiometric Dating, and Paleontological Correlation, Los Angeles, April, 1967: [ 9 pp. SCHAUB, S., 1941. Demonstration der Fauna des Ravin des Etouaires und der Montagne de Perrier. Eclogae Geol. Heir., 34: 320. THAt.ER, L., 1966. Les rongeurs fossiles du Bas-Languedoc dans leurs rapports avec l'histoire des faunes et la stratigraphie du Tertiaire d'Europe. Mdm. Museum Natl. Hi.st. Nat. (Paris), S~r. C, 17:295 pp. TOBIFN, H., 1952. Die oberplioz~ne Sfiugerfauna yon W61fersheim Wetterau. Z. Deut. Geol. Ges., 104: 191. TOBIriN, H., 1956. Zur ()kologie der jungtertifiren S~ugetiere vom H6wenegg/Hegau und zur Biostratigraphie der europfiischen Hipparion Fauna. Schriften Ver. Geschichte Naturgeschichte Baar Angrenzenden Landesteile Donaueschingen, 24:208 223 TOBIE:N, H., 1969. Zur Biostratigraphie der S~iugerfaunen Mittel- und Westeuropas an der Wende Plioz/in/Pleistozfin. Tr. Geol. Inst., Akad. Nauk S.S.S.R., in press. (In Russian, with English summary). TOB~EN, H., 1971. Zur ()kologie der Sfiugerfaunen Mittel- und Westeuropas an tier Wende Plioz~n/Pleistoz~in. Bet. Geol. Ges. Deut. Dernokrat. Rep. Gesamtgebiet Geol. Wiss., in press. VENZO, S., 1965. The Plio-Pleistocene boundary in Italy. INQUA, Rept. Intern. Congr. on Quaternary, 6th, Warsaw, 1961, 1:367 392. VIRFT, J., 1954. Le loess /t bancs durcis de Saint-Vallier (Dr6me) et sa faune de mammiferes villafranchiens. Nouvelles Arch. Museum Hist. Nat. Lyon, 4:200 pp.

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