The pliocene species of Hipparion andtheir biostratigraphical meanings

THE PLIOCENE SPECIES OF H I P P A R I O N AND THEIR BIOSTRATIGRAPHICAI~ MEANINGS(I)

c. I)e~ret

perpignan

by MARIA-TERESA A L B E R D I *

ABSTRACT

RI~SUMI~

During the Pliocene, the ~>in the Mediterranean area decreased sharply in comparison with the Upper Miocene. Nevertheless, this species can be used to represent the H i p p a r i o n biozones in order to compare them with the m a m m a l units. H. g r o m o v a e characterizes the Ventian or M N 13 m a m mal unit ; H . c r a s s u m the Ruscinian or the Lower Pliocene or M N 14 unit ; H . f i s s u r a e the Upper Ruscinian or the Lower Pliocene, M N 15 ; H . rocinantis, the Middle Pliocene or Lower Villafranchian, M N 16a. This distribution can be correlated in most of the Eurasian region.

Pendant le Plioc6ne la faune ~t H i p p a r i o n sur le pourtour m6diterran6en diminue consid6rablement par rapport au Mioc6ne sup6rieur. Cependant, les esp6ces de ce genre peuvent ~tre utilis6es pour repr6senter les biozones d ' H i p p a r i o n par rapport aux diff6rentes unit6s mammaliennes. H . g r o m o v a e caract6rise le Ventien ou unit6 NM 13 ; H . c r a s s u m le Ruscinien ou Plioc6ne inf6rieur ou unit6 N M 14 ; H . f i s s u r a e le Ruscinien sup6rieur ou Plioc6ne inf6rieur NM 15 ; H. r o c i n a n t i s le Plioc6ne m o y e n ou Villafranchien inf6rieur, NM 16a. En conclusion de cette 6rude, on peut 6tablir un parall6le entre la distribution de la plupart des H i p p a r i o n de la r6gion euroasiatique.

RESUMEN Durante el Plioceno las faunas de H i p p a r i o n disminuyen en relaci6n al Mioceno superior, incluso la representaci6n fosilifera de las especies existentes es bastante escasa. No obstante esta especie puede ser utilizada para representar las distintas biozonas de H i p p a r i o n a lo largo del Pliocene en las distintas unidades de mamiferos. H . g r o m o v a e caracteriza el

Ventiense o unidad de Mamiferos NM 13 ; H . crass u m el Rusciniense inferior o Plioceno inferior, NM 14 ; H . f i s s u r a e el Rusciniense superior o Plioceno inferior, NM 15 ; e H . r o c i n a n t i s el Villafranquiense inferior o el Pliocene medio, NM 16a. Esta distribuci6n es correlacionable con la mayoria de los datos conocidos de la regi6n euroasi~tica.

KEY-WORDS : HIPPARION, PLIOCENE, MIO-PLIOCENE BOUNDARY, VILLAFRANCHIAN, BIOSTRATIGRAPHY, CORRELATION. SOTS-CLIPS : HIPPARION, PLIOCENE, LIMITE MIO-PLIOCI~NE, VILLAFRANCHIEN, BIOSTRATIGRAPHIE, CORRI~,LATION

(1) This work was expounded in the ~ Journ~es Charles Dep~ret ~>that took place in Perpignan (France) on October 24, 25, 26, 1985. And it is supported by CSIC (Spain).

* Museo Nacional de Ciencias Naturales. CSIC, Jos6 Guti6rrez Abascal, 2. 28006 Madrid (Spain). Geobios, n ° 19, fasc. 4

p. 517-522, 1 fig.

Lyon, aoOt 1986

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518

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INTRODUCTION The genus Hipparion, owing to its wide and quick distribution through different intervals in space and time, is a good biostratigraphic indicator to deduce the possible distribution of this species in some specific epoch (Alberdi 1974 ; Sen & alii 1978 ; Forsten 1980 ; Alberdi & Morales 1981). During the Early Pliocene, the presence of Hipparion decreased drastically in the Mediterranean

faunas. During the Mio-Pliocene this decrease becomes important, being mainly represented by H. crassum, rare and not well known, used as Hipparion biozone in the unit MN 14 (Mein 1977) or in MN 13-MN 14 transit. We are going to dedicate most of this work to emphasize the last mentioned point.

UPPER MIOCENE - - LOWER PLIOCENE Between France and Spain in the Lower Pliocene, we find H. crassum with a large and short appendicular structure and upper cheek teeth very richly plicated. It is abundant in French places like Perpignan and Montpellier (Dep6ret 1890 ; Michaux 1975), and rare in the only quoted Spanish places: Mina de Lignito of Alcoy and Zeneta in Alicante (Alberdi 1974a and b), the former today absolutely lost as a fossil locality. In 1981, Alberdi & Morales included Alcoy in the MN 13 unit, on account of the presence of a H. gromovae tooth, which could indicate either a brief persistence of that Hipparion in the lowest part of Ruscinian or that it is typically Ventian in the sense that these later Miocene faunas are considered representative of the period between the Later Miocene and the Lowest Pliocene, not Ruscinian, probably conditioned by the Mediterranean drainage. This fauna is characterized by having a lot of new macromammal elements compared to the Upper Miocene. In the Upper Miocene or Ventian levels (eg. Venta del Moro) there are three different Hipparion : H. primigenium, rare in number. This species appeared at the same stage in Granada (Arenas del Rey, Spain). H. gromovae, attributed and used as Hipparion biozone of MN 13 unit (Alberdi 1982 ; Alberdi & Morales 1981).

Hipparion sp. morphotype I, in the evolutive line of Turolian species (Alberdi & Morales 1981). In a recent paper, about the passage of mammal faunas across the Gibraltar Straits, De Bonis & alii (1985) state their doubts as to whether Hipparion passage was from Europe to Africa or vice-versa during the Messinian. Owing to the Hipparion fauna distribution and evolution in the Iberian Peninsula, to the relation between the Teruel and Granada levels Hipparion forms at that time (Alberdi 1974b) and to the presence of H. gromovae in Southern Spain and H. sitifense in Northern Africa (we think both are the same species), the passage must have been from Spain to Africa. On the other hand, Eisenmann (1980) outlines a succession of Hipparion in North Africa with size decreased, similar to the Teruel basin. The problem of nomenclature of H. gromovae and H. sit,lense is difficult to solve because the description of the former and the re-description of the latter (badly defined in the last century. Pomel 1897) were done in consecutive years. Besides the new material from North African places describes by Eisenmann (1980) is different from the material of Arambourg studies (1956). But that is another problem. I think H. gromovae is better described and widely distributed than H. sitifense.

RUSCINIAN OR LOWER PLIOCENE Just after the Ventian, H. crassum levels appear in the Mediterranean area, Montpellier and Perpignan (France). With reference to this species, it is difficult to establish the origin as in the old collections most of the

specimens proceed from Perpignan and yet they are generally just labelled Roussillon. We believe that this forma needs a comprehensive study of all those remains. At first sight, the specimens from all the sites come from the Lower Ruscinian of the Mediterranean

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but it is necessary to determine whether these localities are homogeneous or, on the contrary, there exist significant differences. Other Mio-Pliocene places, containing Hipparion, are Casino and Baccinello V3 (Italy), with paleontological and geological problems, whose stratigraphic situation is not so clear (De Giuli & alii 1983). The Casino basin situation (Giannini & alii 1971 ; Lazzarotto & Sandrelli 1978) is distinguished by the existence of two lagoon discordant episodes. The age attributed to those lagoon cycles is uncertain due to lack of stratigraphic details. There exists a Dipoides problematicus tooth from the lower cycle classified by Mein (in Lazzarotto & Sandrelli 1978) as an aged forma dated Turolian. The origin of this fossil, as well as the others from Casino, is unknown but Lazzarotto & Sandrelli (1978) have established that most of the information about vertebrates from Casino belongs t O upper lagoon cycle. The presence of Tapirus arvernensis (De Giuli & alii, 1983) among Casino fossils excludes the Turolian age for it. Gu6rin & Eisenmann (1982) dated this Tapirus species as Pliocene and Pleistocene but include Casino and Baccinello V3 among places in Upper Miocene containing Tapirus priscus (?). It is possible to deduce that the Dipoidesproblematicus belonged to the lower cycle and Tapirus and Hipparion to the upper cycle. The Casino upper lagoon cycle is compared to V3 level of Baccinello dated by Hiirzeler & Engesser (1976) either Upper Turolian or Lower Pliocene (MN 13 or MN 14). This is in agreement with Azzaroli's (verbal comm.) in Lazzarotto & Sandrelli (1978). De Giuli & alii (1983) think Casino is possibly slightly more recent than Baccinello but need a deeper examination. In fact, the presence of Hipparion sp. and Tapirus sp. demonstrates the hypothetical existence of two deposits (upper lagoon cycle of Casino and Baccinello V3). Above Baccinello V3 lies Arcille series (Lower Pliocene) : MN 15 fauna is also quoted here (HOrzeler & Engesser 1976). Studying the lagoon sediments accumulated in Casino basin, we can appreciate that the upper cycle represents the beginning of a new

519

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Pliocene transgression that is completed with the upper marine sediments ; in fact the lower lagoon cycle in the basin could be an internal basin lying under a higher marine level dated Tortonian. After the regression that took place during the Messinian and that came formed the evaporitic basins of the Mediterranean. The beginning of the Pliocene transgression which originated a lagoon stage that at the end of the transgression could again be inundated by marine water. The only and not goot evidence of this fact, is the conglomerate covering the lagoon cycle containing rare marine fossils and there is not an appreciable discordance between the upper lagoon cycle and Pliocene marine sediments (Lazzarotto & Sandrelli 1978). The age of the sediments deposited in the upper lagoon cycle in the Casino basin could be dated Lower Pliocene, probably earlier Ruscinian, but the new information obtained in the field cannot refute the Lazzarotto & Sandrelli (1978) affirmation that it is Upper Messinian in age (without excluding a Pliocene age). The taxonomy of these Hipparion, difficult due to lack of remains, could be H. aff. crassum, to judge by the features of the cheek teeth for Casino, and Hipparion sp. morphotype I for Baccinello V3, the latter also in Venta del Moro (Spain). Other deposits situated at this stratigraphical level, are the Sahabi where the Hipparion described by Tobien (1982) as H. sp. cf. sitifense and H. sp. cf. primigenium fit perfectly in the Venta del Moro species. Other places containing H. crassum are included as ~alta (Turkey) attributed to Upper Ruscinian (Sen & Heintz 1977) with a dolichopodial Hipparion named H. longipes (Heintz & alii 1975). And GOdOllO (Hungarian) dated also Upper Ruscinian by Kretzoi (1981). Anyway there is not a complete study yet. In view of all this we can use H. crassum as biozone of MN 14 mammal unit, although it can be previously associated to H. gromovae in MN 13-MN 14 transit, Hipparion sp. morphotype I can also be associated. Orrios (Spain), near to Venta del Moro, is situated in the MN 14 unit with scarce presence of this Hipparion sp.

UPPER RUSCINIAN OR LOWER PLIOCENE

Hipparion is better represented in MN 15 mammal unit (Ruscinian s.s.) although it is only present in some places. In Spain it is represented in La Calera

and Villalba Alta (Teruel) and Layna (Soria). The

Hipparion sp. morphotype II from La Calera and Villalba Alta (Alberdi & Morales 1981), with medium to

- - 520 - small size, simple cheek teeth morphology and slender legs, is correlated with H. garedzicum from Bazaleti (Georgia, URSS) (Gabunia & Alberdi in litt.) In Layna (Soria, Spain) H. fissurae is larger and longer and has much longer legs than the one mentioned above, and is in the evolutive trend of Hipparion

Turolian forms (Alberdi 1974b) used as Hipparion biozone of MN 15 mammal unit. In 1981, Alberdi & Morales distinguished this forrna as Ruscinian s.s., MN 14 and MN 15 units, but perhaps it would be better to say that H. crassum characterizes the MN 14 unit and H. fissurae the MN 15 unit. HIPPARION S P E C I E S

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•

SPAIN

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FRANCE

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ITALY

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U.R.S.S.

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CHINA

Vl LLARRUYA

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ILl uJ z .~ I.U a (-.) £30

LAS HIGUERUELAS MN 16o

OLL

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R DE ALMORADIEL KVABEBI

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LAYNA MN15 z

LA CALERA

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BAZALETI PERPIGNAN

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MONTPELLIER MN 14

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VENTIAN MIN VENTA

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MORO

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CASI NO

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MIOCENE Fig. 1 - -

Chronological succession of some typical Hipparion species from localities of Eurasia. Distribution chronologique de quelques esp~ces d'Hipparion typiques provenant de diff6rents gisements d'Eurasie.

M I D D L E P L I O C E N E OR L O W E R V I L L A F R A N C H I A N Finally, in Middle Pliocene or Lower Villafranchian, the mammal unit MN 16a is characterized by H. rocinantis typically caballine, very common in Spain till the arrival of Equus. We find this species in

Villarroya, La Puebla de Almoradiel and Las Higueruelas. It can also be found in Kvabebi (Georgia) at same stratigraphic stage and as H. houfenense in China where it is mentioned as Ruscinian (Zhan-

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X i a n g & M i n - Z h e 1985). Hipparion f r o m t h o s e localities has all the s a m e characteristics in the species level. M a y b e , it is p o s s i b l e to p u t here also Hipparion f r o m R o c c a n e y r a (France). A l b e r d i & B o n a d o n n a (1986)

521

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p l a c e d it as Hipparion ( p r o b a b l y rocinantis ?) in this level, b u t the g e o l o g i c a l setting o f R o c c a n e y r a d e p o sits is n o t so clear.

CONCLUSION

A m o r e exhaustive s t u d y o f d i f f e r e n t species o f Hipparion is necessary in o r d e r to establish the right evol u t i o n a n d p a l e o b i o g e o g r a p h y o f the E u r a s i a n r e g i o n

that, a l t h o u g h b r o a d l y o u t l i n e d in the p r e s e n t p a p e r (fig. 1), m u s t be a n a l y s e d in m o r e d e p t h in f u t u r e studies.

Acknowledgements

thank J. Arroyo for the Illustration, J.L. Casaseca for the typescript and Sheila Stewart for the correct this English text.

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M a n u s c r i t d6finitif re9u le 10.04.1986