Late Miocene turnover in the Spanish mammal record in relation to palaeoclimate and the Messinian Salinity Crisis

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Abstract

The latest Miocene and earliest Pliocene is a period of marked changes in geography and climate in the circum-Mediterranean area. Its geographical situation gives Spain a key position, and its fossil mammal record reflects these important geographical and climatic changes particularly well.

The Spanish mammal record shows a gradual change of composition towards the typical Pleistocene and recent fauna, marked by several extinction and dispersal events. These events have the effect that northern Eurasian elements increase, while taxa that locally go extinct may live on in tropical areas. Superposed on this large scale trend, there are some events of short lived incursions of animals of African or south Asian origins, which seem to reflect a short reversal of this tendency and temporary geographical changes related to the Messinian Crisis. These latter incursions are detected very well in Spain but also in other parts of southern Europe.

The Messinian Salinity Crisis is caused by interplay of tectonics, and global sea level variations which in turn depend on climate. While some of these changes were permanent, causing the replacement of central Italian insular faunas by mainland faunas, others were temporary probably allowing the short lived incursions of African mammals into SW Europe and certainly allowing for mammals to reach the Balearics, Corsica and Sardinia.

We document an important event around 6.1 Ma, of mammals adapted to dry or open environments dispersing from Africa and SE Europe to SW Europe. During the Messinian desiccation phase, animals adapted to more humid environments reached some of the west Mediterranean islands, suggesting that, at least during part of the desiccation phase, a part of the former sea bottom was not desertic. Between 6.3 and 5.3 Ma tectonics united the island of Toscany to the European mainland, thereby closing a seaway of some 10 km wide and 100 m deep.

Introduction

The study period, from some 7 to 4 Ma ago, is centred around the Mio-Pliocene transition, and in terms of the Neogene Mammal Units (Mein, 1990, De Bruijn et al., 1992, Mein, 1999) covers the later part of MN12 and the whole of MN13 and 14. It also covers the transition between the Turolian and Alfambrian ages, which correspond to the stages of the same name, that have their type sections close to Teruel and a little further north in the valley of the Alfambra. The local zonation of the Turolian contains zones J to M, in which L corresponds to MN 12 and M, with the sub-zones M1–3 to MN13 (Dam, 1997). The Alfambrian covers largely the same time as the Ruscinian, but was introduced because it has a good type section with superposed mammal localities (Mein et al., 1990). Palaeomagnetics in the Turolian type area suggest that the lower boundary of this stage is 8.7 Ma old, while the upper boundary is clearly younger than 6.1 Ma ago (Dam et al., 2001).

The “Mid-Vallesian Crisis”(Agustí and Moyà-Solà, 1990) is a major event that occurred at the transition of MN9–10 and early in MN10, around 9.7 Ma (Agustí et al., 2001). After this event, there was a period with a relatively stable fauna, in which bovids, giraffids and abundant equids suggest open and more or less dry environments. Data provided by Bruijn et al. (1992) suggest that MN12 does not differ so much from MN11 and is noted in Western Europe in the appearance of few new genera (only “Valerimys”— = Huerzelerimys — and Ruscinomys) and the disappearance of few others (Prospalax, Dorcatherium, Micromeryx). In the Turolian type area, the MN11–12 transition, between 7.5 and 7.9 Ma ago, is noted in the appearance of more evolved species in the genera Huerzelerimys, Parapodemus and Occitanomys (Dam, 1997). More data provided by Bruijn et al. (1992) indicate that the Alfambrian age (or Ruscinian, or MN14 and 15) was also a period with a relatively stable fauna, while the transition to the Villafranchian (starting MN16) was again a period of change. The faunal events late in the Turolian and around the Turolian–Alfambrian transition thus mark the transition between two relatively stable periods. These events are studied in more detail here.

The mammalian faunas of the type areas of the Turolian and Alfambrian have been subject of many publications. Weerd (1976) and Dam (1997) treated the rodents and Alcalá Martínez (1994) the large mammals. The Desmaninae, Carnivora, Suidae, Cervidae, Bovidae are included by Rümke (1985), Fraile et al. (1997), Made et al. (1992), Van der Made, 1997a, Van der Made, 1997b, Azanza (2000), Azanza et al. (1997), and Nieto et al. (1997). Palaeomagnetism of the Turolian was published by Krijgsman et al. (1996), while Dam (1997) and Dam et al. (2001) published charts of the Turolian localities and mammals in combination with the palaeomagnetic “dates”. Mein et al. (1990) published the Alfambrian.

Section snippets

The events

Fig. 1 indicates the ranges of the taxa that appeared by dispersal or disappeared by (local) extinction in south western Europe (Spain, France and Italy) and Fig. 2 compares these ranges with those of the same taxa in other areas, particularly central, eastern and south-eastern Europe, as well as with south-western Asia. In the following, the most important events are discussed in chronological order.

Here we discuss dispersals and (local) extinctions, not so much grades of evolution. However,

Biogeography and its relation to palaeogeography and palaeoclimate

The study period is a time of marked changes in the European mammalian fauna and these occurred in four major events. The timing, mode and geographical provenance and extent of these dispersals provide information on palaeogeography and the effect of climatic and environmental change on the fauna.

Conclusion

The time slice between 7 and 4 Ma is one of important changes in Mediterranean geography and in the circum-Mediterranean mammalian faunas; in both cases Spain has a key position. An interplay of tectonics and global climate controlled these changes, some of them being transitory, while others had a more permanent character.

Within the Mid-Miocene there was a spectacular drop in global temperature, followed by a more gradual decrease in temperatures towards the state in the Pleistocene, with only

Acknowledgements

This research is partially based on the study of fossils in numerous institutions in Europe and Turkey. We thank our colleagues in those institutes for help and access to material and J. Agustí for comments on the manuscript. This paper is a contribution to the ESF network EEDEN, projects PB98-0691-CO3-01, PB98-0513, BXX2000-1258-CO3-01, BTE2002-00410 and BTE2003-0301 and the “Unidades Asociadas” program of the CSIC.

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