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Home , Archaeobelodon, Choerolophodon, Phiomia, Platybelodon, Stegolophodon, Zygolophodon

J. van der Made & A.V.Mazo Museo Nacional de Ciencias Naturales, Madrid

Proboscidean dispersals from towards Western Europe

Van der Made, J. & Mazo, A.V., 2003 - Proboscidean dispersals from Africa towards Western Europe - in: Reumer, J.W.F., De Vos, J. & Mol, D. (eds.) - ADVANCES IN RESEARCH (Proceedings of the Second International Mammoth Conference, Rotterdam, May 16-20 1999) - DEINSEA 9: 437-452 [ISSN 0923-9308] Published 24 May 2003

Proboscideans of African origins dispersed towards Western Europe during two distinct periods: browsing forms did so between some 18.5 and 14 Ma and grazers between some 2.7 and 0.8 Ma ago. The earlier dispersals occurred when Western Europe had a warmer and less seasonal clima- te than today, the later dispersals occurred at moments of fundamental global climatic change when open landscapes increased in Europe. Between 8 and 6 Ma ago, the changing environment caused African to adapt to grazing. It is this adaptation that allowed them to enter again Western Europe.

Correspondence: J. van der Made & A.V. Mazo, Consejo Superior de Investigaciones Científicas, Museo Nacional de Ciencias Naturales, C.José Gutiérrez Abascal 2, 28006 Madrid, Spain; e-mail [email protected] (JvdM) and [email protected] (AVM)

Keywords: Proboscidea, dispersal, Africa, Western Europe, biogeography

INTRODUCTION where involved. Tassy (1990) and Antunes From the onwards, proboscideans (1990) showed that the initial dispersal of the are well known from most continents. Since proboscideans was diachronous and that there the beginning of this century, their origin is was more than one dispersal event. Thomas supposed to be in Africa. Describing material (1985) supposed that there were two major from the Fayum, Andrews (1904, 1906), pro- faunal exchanges and Van der Made (1996) posed for the first time that the origin of the recognised up to six dispersal events between proboscideans is in the African . Africa, northern Eurasia, the Indian Schlesinger (1912) determined Subcontinent and the area formed by SE angustidens in the lower Miocene of Algeria. Europe and Anatolia. These events ranged in Matsumoto (1924) supposed that the zygod- age between 21 and 12.5 Ma. The probosci- onts descended from and the deans that have been assumed to disperse bunolophodont elephantoids from , from Africa include Hemimastodon, Gompho- both from the Palaeogene of Africa. This idea therium, , , was followed by Tobien (1978). , and . Berggren & Van Couvering (1974) and The entry of Mammuthus (or ‘Archidisko- Madden & Van Couvering (1976) popularised don’) in Europe coincides with that of Equus the ‘Proboscidean Datum Event’. This event and is considered to be an important strati- was supposed to be a major faunal exchange graphical marker (Azzaroli et al. 1988). This between Africa and Eurasia during the Early event is believed to have occurred some 2.7 Miocene. This ‘event’ attracted much discus- Ma ago (Sardella et al. 1998). The dispersal sion about when it happened and which taxa of antiquus towards Europe has

437 ADVANCES IN MAMMOTH RESEARCH DEINSEA 9, 2003

aroused much less interest, possibly because lophodon are cited from the Turolian of this has often been confused with China (Tobien et al. 1986, 1988; Guan 1996). Mammuthus. Though the possibility of other A skull and mandible from the Upper dispersals will be discussed below, an interes- Miocene of Hohenwarth in Austria were ting pattern arises: the proboscidean dispersal interpreted as intermediate between events towards western Europe form two and (Zapfe 1957) and clusters, one in the Early-Middle Miocene many isolated teeth have been considered as and one in the Late and Pleistocene. intermediate (Schlesinger 1917; Lehmann It is the aim of this paper to establish which 1950; Mottl 1969; Tobien 1978). Material dispersals occurred and when they occurred, from Concud has been assigned to Anancus and to study their context. or a closely related form (Lehmann 1950; Bergounioux & Crouzel 1958; Mazo 1977) THE TAXA and has an age of 6.8 Ma (Van Dam 1997). Material from the Late Miocene of Perim Hemimastodon (Tassy 1986) might be among the oldest Hemimastodon is known only from two teeth material, but the exact age is not known. from Bugti, described by Pilgrim (1908) and Material from Mpesida, Kenya has an estima- Tassy (1988). The exact age is not well ted age of 6.4-6.0 Ma (Kalb et al. 1996) and known, but the bulk of the fauna might be as material from Langebaanweg QSM and old as 21 Ma (Van der Made 1996). A phylo- Sahabi is slightly younger. The age of these genetic analysis by Tassy (1990) suggests that localities was controversial, but a revision of Hemimastodon is the sister taxon of the the Tetraconodontinae suggests that the age . The primitive character of might be about 5.5 Ma (Van der Made these teeth is in line with an old age, but is 1999a), though the possibility even exists that not sufficient ground to assume that the taxon one of the localities might be in regressive was present in the Indian Subcontinent prior sediments deposited during the TB3.3 cycle to the major Early Miocene faunal exchange of Haq et al. (1997). Even if a simple lineage at about 21 Ma; though primitive, the taxon is not accepted, there is no consensus on the may have come from Africa. time and place of origin of the latter two genera. Tobien (1972, 1973) has interpreted some remains from western Europe that ori- Historically Gomphotherium, being a bunod- ginally were placed in Gomphotherium, as ont elephantoid, has been traced back to primitive platybelodonts. Tassy (1984) essen- Phiomia (Tobien 1976, 1978). Many authors tially followed this idea, but introduced the supposed an evolution from Gomphotherium name for these proboscideans to Tetralophodon to Anancus (Schlesinger and tentatively assigned material from 1917; Lehmann 1950; Mottl 1969; Tobien Legetet to this (Tassy 1984). Astibia & 1978), though Tassy (1985) did not accept Mazo (1988) and Mazo (1996) assign the this lineage. Material from Portugal that has western European material to Gomphotherium. been considered intermediate between Of all the forms treated above, Gompho- Gomphotherium and Tetralophodon (Antunes therium is the only one that experienced & Mazo 1978), was considered to be platybe- dispersal from Africa to Western Europe, that lodont (Tassy 1985). Both Gomphotherium is beyond discussion. There is a temporal gap and Tetralophodon have been reported from between Phiomia and the earliest Gompho- Eppelsheim, Germany (Lehmann 1950; therium. Material from Legetet (Africa; Set I, Tobien 1980, 1996), but in SW Europe, this MN3 equivalent; faunal sets Pickford 1981; association has never been found (Mazo MN units - Mein 1990, De Bruijn et al. 1992) 1977, 1996). Gomphotherium and Tetra- and material from Bugti (Pakistan; MN 3

438 VAN DER MADE & MAZO: Proboscidean dispersals

equivalent) has been assigned to Gompho- Zygolophodon. Meswa Bridge is older than therium (Pickford & Andrews 1981; Pilgrim the first major Miocene faunal exchange 1912). In Europe the genus is cited from between Africa and Eurasia. During the Late Navère (France; late MN 3; Bulot & Miocene Mammut replaced Zygolophodon in Ginsburg 1993) and Artenay (France; early Europe and northern Asia. A skull from (pro- MN 4; Ginsburg & Antunes 1966; Tassy bably) the Odessa region was interpreted as 1977a) and from an MN3 equivalent locality intermediate form (Kubiak 1972). in Japan (Takai 1954 as cited by Tassy 1990). Adispersal of Zygolophodon from Africa to Qiu et al. (1999, p. 445) believe that indeter- Europe and Asia is widely accepted, though minate proboscideans were present in China the opinions differ on when this may have in MN 2. However, the arguments for an age happened. Some authors (Tobien 1976, 1996; comparable to MN2 are not convincing and Tassy 1977b, 1996) think that Zygolophodon MN 3 age seems more likely. The first deter- arrived in Europe together with Gompho- minable Chinese proboscidean is Gompho- therium, while others (Stehlin 1926; Gaziry therium (Qiu et al. 1999) of MN 4 age. The 1976; Mazo 1977) think that the teeth of this most accepted opinion holds that Gompho- age that are attributed to Zygolophodon, are therium evolved in Africa and dispersed morphological variants of Gomphotherium. A during MN3 to the Indian Subcontinent, in mandible fragment of Gomphotherium from MN 3 (early/late?) to China and Japan, and Paracuellos 5 (province of Madrid; MN6) has late in MN3 or at the MN3-4 transition to a p4 and a m1 with zygodont and a m2 with Western Europe. bunodont morphology (Mazo 1985, 1996), suggesting that identifications based on Deinotherium isolated specimens are disputable. For Tassy Deinotheres are found in Africa, the area of (1985) Zygolophodon is present in the SE Europe and Anatolia, and the Indian Orleanean of France and Spain, though not in Subcontinent. The smaller forms are often Portugal. The oldest locality where it is cited placed in and the larger is in Marsolan (Tassy 1977b), which is placed (and later) forms in Deinotherium. Different in MN3 and where Brachyodus onoideus is species of both genera are cited from the present (Bulot & Ginsburg 1993). Within different (sub)continents, which either implies MN3 Brachyodus intermedius evolved into B. multiple dispersals or a polyphyletic origin of onoideus, which implies that Marsolan is in Deino-therium. For the time being, we assign the later part of MN3. From early MN6 all deinotheres to Deinotherium. The origin onwards the presence of Zygolophodon in of the deinotheres is assumed to be in Africa Europe is indisputable (Elgg, Switzerland; (Osborn 1936). The earliest record outside Schinz 1824). Zygolophodon is described or Africa is in Bugti (Pakistan; Pilgrim 1908) cited from the Orleanian and Vallesian of and might be as old as 21 Ma (Van der Made China (Tobien et al. 1988; Guan 1996) and 1996). Deinotheres arrived later in Europe Kazakhstan (Gabunia 1981; Tassy 1985). (Tassy 1990) and are found in localities like Artesilla (Spain; Aragonian zone C or late Choerolophodon, Platybelodon MN4; Azanza et al. 1993). The entry in The oldest record of Choerolophodon is pos- Europe is supposed to have happened 16.5 sibly from Buluk (Kenya; dated 17,2 Ma; Ma ago (Van der Made 1996). Tassy 1990). The genus is known from Bugti (Lydekker 1884a,b). However this ‘locality’ Zygolophodon has many levels, including the Middle The oldest true zygodont known is Miocene and the exact provenance of the Eozygodon from Meswa Bridge (Kenya; 22 material is often not known (Pickford 1988). Ma; Tassy & Pickford 1983), and is related to The oldest extra-African record is in Chios

439 ADVANCES IN MAMMOTH RESEARCH DEINSEA 9, 2003

(Greece; MN5; Tobien 1980), it is also the presence of Mammuthus at about 3 Ma in known possibly from the Kamlial Formation Cernatesti and Tulutesti (Rumania). The and certainly from the Chinji Formation oldest citations from Western Europe are (Lydekker 1884a,b; Tassy 1983). The presen- from Huelago, with an age of some 2.7 Ma ce of Choerolophodon in China (Tobien et al. (Mazo 1989; Van der Made & Mazo 2001), 1984; Guan 1996), has been doubted (Tassy Montopoli, with an age close to 2.58 Ma 1994). Both Chios and the base of the Chinji (Italy; Azzaroli et al. 1988; Sardella et al. Formation are believed to be about 14 Ma old 1998) and the Red Crag (England; Lister and the base of the Kamlial Formation might 1996) at about 2.5-3 Ma. be as old as 16.5 Ma (Van der Made 1996). Though the genus has been cited from Spain Elephas (Bergounioux & Crouzel 1958), this material The oldest record of Elephas is E. ekorensis is currently considered to be milk dentition of from Ekora and Kanapoi, with an age of over Anancus (Mazo 1977, 1996). 4 Ma (Maglio 1970; Beden 1979; Fitch & The oldest record of Platybelodon is from Miller 1976). Elephas first appeared in the Loperot (Kenya, MN4 equivalent; Tassy Indian Subcontinent with Elephas planifrons 1990). The oldest extra-African record of the 3.3 Ma ago (Hussain et al. 1992). One of the genus is from Shanwang and Tongxin (China; oldest citations of Elephas antiquus in Guan 1996) and Belometchetskaya (southern Western Europe is from Huescar, Spain Russia; Gabunia 1973). Shanwang is correla- (Mazo 1989). Huescar is placed in the latest ted to MN 5 (Qui et al., 1999). Suids indicate Early Pleistocene, between some 0.9 and 1 that Belometchetskaya is older than Ding- Ma (Sesé et al. 2001). E. antiquus was also jiaergou (in the lower level at Tongxin) and correctly cited from Soleilhac, France (Bout that both are MN5 equivalent (Guan & Van 1964). This locality is placed in the early der Made 1993; Van der Made 1996). The Middle Pleistocene (Roebroeks & Van possible existence of a primitive platybelodont Kolfschoten 1995). Archaeobelodon has been discussed above. The available data suggest that Choerolo- THE DISPERSAL EVENTS phodon originated in Africa and dispersed not later than 14 Ma ago, possibly even earlier, The Early Miocene setting but probably not earlier than 16.5 Ma ago, to During the late and early Miocene the Indian Subcontinent and the area of SE Africa was separated from Eurasia by the Europe and Anatolia. Also Platybelodon Tethys, and many , including pro- might be of African origin and dispersed boscideans, evolved in isolation. Also the about 14 Ma ago to the Caucasus area, Asia Indian Subcontinent was isolated. The fossil and the Indian Subcontinent. None of these record of this period is poor in both areas, forms reached Western Europe. and it is not known whether, or how much faunal exchange may have occurred between Mammuthus both areas. While tectonics slowly drove The oldest record of Mammuthus is from Africa and India closer to Eurasia, an impor- Langebaanweg QSM in South Africa (Maglio tant and abrupt sea level low stand at around & Hendey 1970; Kalb et al. 1996). The age 21 Ma produced a land connection between of this locality has been controversial, the these areas and a major faunal exchange suids suggesting an aberrant age (Hendey occurred. From this time on sea level low 1981), but a recent revision of the Tetracono- stands allowed from repeated dispersal dontinae resolved the problem (Van der Made events. Accumulative sea level lowering 1999a) and the locality might have an age of during the late Middle Miocene resulted in 5.5 Ma. Radulescu & Samson (1994) claim more permanent land connections; whereas

440 VAN DER MADE & MAZO: Proboscidean dispersals

initially the combination of climatical chan- moment, deinotheres, Hemimastodon and ges and sea level controlled faunal dispersal Gomphotherium reached the Indian Sub- events, after some 11 Ma ago, the importance continent and the latter genus may have of sea level was reduced (Haq et al. 1987; reached China and Japan. There is no eviden- Van der Made 1996, 1999; Miller et al. 1995; ce for a proboscidean dispersal to Europe. Rögl 1998; Rögl & Steininger 1983; Thomas The only large of African origin that 1985). reached Europe in this initial event is the Initially, the majority of dispersals were anthracothere Brachyodus (Van der Made from Eurasia to the southern (sub)continents. 1999b). During the same event it reached Most of the time, this was the predominant India, and possibly Japan where it was found direction of dispersal, though between 17.5 with Gomphotherium (Takai 1954 as cited by and 12.5 Ma and around 5 Ma ago, dispersals Tassy 1990) and Siberia (Vislobokova 1994) to the north were common (Thomas 1985; as well. Pickford & Morales 1994; Van der Made 1996, 1997, 1999b). Changes in composition Gomphotherium-dispersal towards of Spanish rodent faunas around 14 Ma ago, Europe - about 17.5 Ma have been interpreted as resulting from a The first proboscideans that reached Europe major climatical event coeval to a major did so during the later part of MN3 or at the change in marine oxygen isotope values (Van MN3-4 transition and include Gompho- der Meulen & Daams 1992). Taxa of (sub)- therium and perhaps Zygolophodon. The tropical origins or taxa that are limited by rodent Democricetodon (of Anatolian origin), minimum temperatures or seasonality started the carnivore Hyanailouros (of African ori- to go extinct in Europe from about 12.5 Ma gin), and the bovid Eotragus and the suoid onwards, whereas they lived on longer in Taucanamo (both of Asian origins) entered areas more to the south (Van der Made 1992, Europe at the MN 3-4 transition, around 17.5 1993, 1997, 1999b). The dominant direction Ma ago and at about the same moment a of dispersals and the duration of taxa with number of taxa of Eurasian origins entered tropical origins or dependence on temperature Africa. (Van der Made 1996, 1999b). The may be indicative of climate at a particular proboscidean localities of Marsolan and time. It has been hypothesised that Europe Navère seem to be slightly older than the had a tropical climate during part of the Early entry of Democricetodon in western Europe and Middle Miocene (Pickford & Morales (Bulot & Ginsburg 1993), or at least do not 1994), alternatively, the European climate have this rodent. The question is whether the may have been simply warmer or less season- time difference is important. al than in the present (Van der Made 1999b). Deinotherium-dispersal towards Europe Initial dispersal to the Indian - about 16.5 Ma Subcontinent - about 21 Ma Deinotherium entered Europe when also the During the initial faunal exchange event, at rodents Megacricetodon, Eumyarion and least 15 genera of large mammals and many Fahlbuschia (of Anatolian origins) and the small mammals of Eurasian origin entered chalicothere Chalicotherium, the suid Africa and India (Van der Made 1999b), this Bunolistriodon and the tragulid Dorcatherium corresponds more or less to Thomas’ (1985) (all with a previous record in Africa and the NDP1 dispersals towards the Afro-Arabian Indian Subcontinent) entered western Europe. plate. This dispersal event is supposed to About five mammal genera entered Africa. have occurred during the sea level low stand Some mammals, both of African and Asian of the TB2.2 cycle of Haq et al. (1987) (Van origins entered the Indian Subcontinent, der Made 1996). Not later than at this including the primate Dionysopithecus. This

441 ADVANCES IN MAMMOTH RESEARCH DEINSEA 9, 2003

dispersal event corresponds largely to Later Middle Miocene to Early Pliocene Thomas’ (1985) NDP1 dispersals to Eurasia dispersal events and is believed to be allowed for by the sea An event 12.5 Ma ago, involved several dis- level low of the TB2.3 cycle of Haq et al. persals from Africa towards western Europe (1987) (Van der Made 1996, 1999b). A homi- (Van der Made 1996, 1999b), but none of noid, possibly Griphopithecus (Andrews et proboscideans. At the same time, there were a al. 1996), dispersed to Europe either during number of local and total extinctions in this or another event 15.5 Ma ago (Van der Europe (Van der Made 1992, 1993). Made 1999b). That event did however not that are limited by minimum temperatures, involve any Proboscidea. The dispersals to such as crocodiles, disappeared from central the north were more important than those to Europe, but lived on more to the south. the south and this is the time when Europe Dorcatherium, that was represented by four has been interpreted to have had a tropical species in Europe, decreased its species climate (Pickford & Morales 1994). The diversity to one, but remained diverse in the northward migrations might reflect the Indian Subcontinent. Several lineages that tendency towards the climatic optimum. had been present in Europe for millions of years went extinct. Later dispersal events The possible Zygolophodon-dispersal - were predominantly towards Africa (Vrba about 14 Ma 1995; Van der Made 1999b), though during As discussed above, there are different opi- the latest Miocene there were again relatively nions on the first European Zygolophodon; many dispersals from Africa towards Europe, the genus is either supposed to have been but there is no convincing evidence that these present late in MN3/early in MN4, or in the involved proboscideans. The oxygen 18 iso- earlier part of MN 6. In the latter case it may tope curves from the Atlantic suggest high have entered in the major MN5 dispersal temperatures during the late Early and early event, when the primates Pliopithecus and Middle Miocene, a very important drop Plesiopliopithecus, of african origins, the between some 14 and 12.5 Ma and a gradual suoids Sanitherium (with a previous record in further decrease towards the present. These Africa and the Indian Subcontinent) and curves have been related to faunal events Conohyus (of Indian origin) and the rodent (Bonifay 1980; Pickford 1987; Van der Cricetodon, of Anatolian origin, entered Meulen & Daams 1992). Europe. At this time Platybelodon dispersed towards the area extending from the Black The Plio-Pleistocene setting Sea area to northern China and Choero- The late Pliocene and Pleistocene are charac- lophodon entered SE Europe, an important terised by climatic fluctuations of increasing faunal exchange took place. Hyraxes and magnitude and length. These fluctuations aardvarks, of African origins, spread towards caused important biogeographical reorganisa- SE Europe and the hyraxes to the Indian tions. The relation between astronomical Subcontinent as well. Several bovid genera of (Milankovic) cycles and climate has been Asian origins dispersed towards Africa, the known for a long time, but it is in recent Indian Subcontinent and SE Europe. This years that great advances have been made in event has been related to the sea level low the recognition of these cycles in geology and stand at the onset of the TB2.5 cycle of Haq paleontology. The cycles are changes in the et al. (1987) some 14 Ma ago (Van der Made shape of the orbit and the orientation of the 1996, 1999b) and corresponds partially to axis of the earth. The eccentricity of the orbit, Thomas’ (1985) NDP2 dispersals towards the obliquity the earth's axis relative to the Eurasia. plane of the orbit, and the precession of the equinoxes are subject to cycles with durations

442 VAN DER MADE & MAZO: Proboscidean dispersals

of some 100, 40 and 20 ka respectively The arid belt that runs at present from the (Foucault 1993). The shorter cycles have a north of Africa to the Middle East and conti- greater impact on higher latitudes. Before nues into central Asia may have been present some 2.7 Ma ago, the 20 ka cycle had a in some form for a long time, but it may have greater influence on global climate, after that been more permeable to mammal dispersals date the 40 ka cycle became dominant and 2.7 Ma ago. These changes are probably rela- after some 1 Ma ago the 100 ka cycle ted to the growing importance of the 40 ka (deMenocal 1995). Important biogeographical cycle, causing periods of a greater seasonality reorganisations occurred around 2.7 and 1 Ma and lower temperatures. Lower temperatures ago, whereas the period inbetween was relati- directly affect the degree of aridity, though vely stable. other mechanisms may have acted on the environment, causing biogeographical adjust- Mammuthus dispersal - about 2.7 Ma ments. Mammuthus entered Western Europe slightly before 2.6 Ma, possibly about 2.7 Ma ago. Elephas dispersal - about 0.8 Ma Around this date important changes occurred. Elephas () entered Europe Large mammals that dispersed into western some 0.8 Ma ago. As we have seen, between Europe at this time include the equid Equus, about 1.2 and 0.8 Ma ago the 100 ka eccen- the deer Eucladoceros, the bovid Leptobos tricity cycle posed itself as the dominant etruscus, and of course the proboscidean cycle forcing the climate and the impact of Mammuthus. Several of these animals, inclu- the glaciations on the European environment ding Mammuthus, are typical of open habi- became gradually stronger. Species from dif- tats. More or less simultaneously, forest ani- ferent environments sharing a tolerance for mals, like Tapirus and Mammut went extinct. cold periodically lived together as the ‘glacial The dispersal of Equus, of American origin, fauna’. With the successive glaciations, they is traditionally related to a first cold period in adapted to the unique glacial environment. a series that lead to the severe glaciations of Species with a lesser tolerance for cold for- the Middle and Late Pleistocene. The accu- med the ‘interglacial fauna’ but lived during mulation of continental ice caused a sea level the glaciations in refugia in southern Europe. low stand, allowing for the crossing of the Whereas the separation into ‘glacial’ and Bering Strait by Equus. ‘interglacial’ faunas seems to have occurred Another change that occurred at this time, after 0.8 Ma, when the 100 ka cyclicity was is the southward shift of the limits of the well established, the transition period Palearctic bioprovince. This is well reflected between some 1.2 and 0.8 was characterised in the distribution of the deer. Deer originated by the appearance of a number of new taxa. in Asia, entered Europe some 21 Ma ago, These taxa do not seem to have evolved in maintained the southern limit of their distri- Western Europe, consequently they must have bution within Anatolia till some 2.5/2.7 Ma dispersed from an area outside (western) ago, when they entered the Indian Subcon- Europe. Using paleomagnetism in combina- tinent and North Africa (Maglio 1978; tion with biostratigraphy, the dispersals can Hussain et al. 1992; Barry & Flynn 1990). tentatively be arranged in three clusters. Lagomorphs first entered the Indian Subcon- tinent (Steensma & Hussain 1992). It has Just before 1.07 Ma. Localities that are been hypothesised that Cervidae, Talpidae, supposed to be only slightly older than the the equid Anchitherium and Tapiridae have Jaramillo event include Venta Micena and been limited for millions of years in their Pirro Nord. Traditionally these localities have southward expansion by an area that was too been located around 1.2 Ma, though there is arid for them to cross (Van der Made 1992). no objective way to estimate their exact age.

443 ADVANCES IN MAMMOTH RESEARCH DEINSEA 9, 2003

Taxa that first appear in these localities inclu- or eastern Europe. Coelodonta, Rangifer and de the cervid Eucladoceros giulii and the some of the bovids seem to have appeared bovids Bison degiulii, Soergelia minor, earlier in Germany and Austria than in France Praeovibos sp. (?) and Capra alba. All taxa and have been interpreted as belonging to the were probably typical of dry and/or open glacial fauna, though their glacial adaptations landscapes, though some of the bovids may may not yet have been fully developed at this have been inhabitants of more mountainous early stage. S. kirchbergensis is considered to environments while other bovids and the belong to the interglacial fauna. This suggests large cervid are likely to have preferred the possibility of more than one event. plains. Probably all these taxa have their ori- The vast majority of taxa that dispersed gins in Asia or Eastern Europe. into Western Europe were typical of open landscapes. This does not seem to be reflec- 1.07 - 0.78 Ma. Localities that are as old as ted in an equivalent number of extinctions of the Jaramillo event, or slightly younger, such animals that preferred closed habitats, but in as Untermassfeld, Vallonet, Atapuerca TD4-6, an increase in biodiversity. It seems therefore etc. have the first records of a number of spe- that the European landscape became progres- cies that still live, as well as others extinct sively more open, though closed habitats con- now. These taxa include: the hyaenid tinued to exist. The entry of Elephas antiquus Crocuta, the felids Panthera leo and Panthera forms part of these changes. Between about pardus, the suid Sus scrofa, the deer 2.7 and 0.8 Ma Mammuthus was the only Capreolus (?) and Cervus elaphus, and bovids elephantine in Europe, but after the entry of of the Bison menneri/voigtstedtensis group. Elephas it became part of the developing gla- The carnivores have their ultimate origin in cial faunas, whereas the latter became part of Africa, but were prior to 1.07 Ma present in the interglacial faunas. It is possible that the the Middle East. The ungulates have their ori- opening of the landscape allowed for a gin in northern Asia or Eastern Europe. The second elephantine species in Europe, follow- suid and the deer are more typical of a tem- ed by an adaptation of each lineage to a diffe- perate closed environment. Whereas the hya- rent environment, the newcomer ending up enid is more typical of an open environment. with the less severe ‘interglacial niche’.

Shortly after 0.78 Ma. Localities that are not DISPERSALS, EVOLUTION AND very much younger than 0.8 Ma include EUROPEAN ENVIRONMENT Voigtstedt, Süssenborn, Bad Frankenhausen, Figure 1 indicates the distribution in time of Hundsheim, Huescar etc. Taxa that first proboscideans in Western Europe. Thick lines appear in these localities include the probos- represent the range of groups or lineages of cidean Elephas antiquus, the rhinos proboscideans, dots indicate the approximate Coelodonta, Stephanorhinus hundsheimensis age of the type locality of the type species of and Stephanorhinus kirchbergensis, the cer- a genus. Dispersals of proboscideans from vids Megaloceros soleilhacus (?), Megalo- Africa towards Western Europe occurred in ceros savini and Rangifer and the bovids two clusters about 17.5, 16.5 and possibly 14 Hemitragus bonali, Praeovibos priscus, Ma ago and about 2.7 and 0.8 Ma ago. The Ovibos suessenbornensis and Soergelia elisa- figure also indicates eustatic sea level after bethae. Most of these animals may have pre- Haq et al. (1987). The earliest three dispersal ferred an open landscape, though this is pro- events are separated by periods of isolation of bably not the case for S. kirchbergensis. Africa. After two successive and cumulative Hemitragus was probably more typical of the drops at 14 and 12.5 Ma sea level remained mountainous environments. The deer, rhinos low, save for a short period after 5 Ma, and and bovids probably had their origin in Asia the land connection between Africa and

444 VAN DER MADE & MAZO: Proboscidean dispersals

Eurasia became probably permanent. It is dramatic drop in temperature occurred. The noteworthy that in the long period between later dispersals occurred when temperatures 12.5 and 2.7 Ma, when land connections did were lower. Temperature itself was probably exist, no dispersals could be demonstrated, not a limiting factor. As proboscideans are neither around 21 Ma ago. The availability of very large animals, it is to be expected that land connections was a prerequisite, but it they are less sensitive to temperature, but was apparently not the real factor controlling temperature may have acted on the environ- migrations most of the time. ment and in this way it could have been a It is evident that the earlier dispersals limiting factor. The proboscideans that dis- occurred during high temperatures as indica- persed towards western Europe are not only ted by the isotope record (Fig. 1). The period two groups according to the time of dispersal, between 17.5 and 14 Ma ago had a climatic but also according to masticatory adaptations. optimum, between some 14 and 12.5 Ma a The proboscideans that reached Western

Figure 1 Oxygen isotope records (after Flower & Kennett 1993) and eustatic sea level (after Haq et al. 1987) and range chart of the Proboscidean genera in Western Europe. Dots refer to the position of the type localities of the type species.The range of the Gomphotherium-Tetralophodon-Anancus and Zygolophodon-Mammut groups are indicated as if two lineages, though speciation events may have occurred within each group.

445 ADVANCES IN MAMMOTH RESEARCH DEINSEA 9, 2003

Europe between some 17.5 and 14 Ma ago 14 Ma from Europe, went extinct in Anatolia had low crowned cheek teeth, wide cusps or (Van der Made 1992, 1993, 1996). Still some crests, thick enamel, relatively few cemen- animals of southern origins migrated north- tum, relatively short M3 and at some time in ward and some of those that had done so life had premolars and molars in use. These before, still lingered on. proboscideans were probably browsers that The next important climatical change is the may have eaten hard, but not very abrasive ‘Mid-Vallesian Crisis’, when seasonality food. They may have eaten fruit. Deino- became more accentuated (Van Dam 1997; therium and the Zygolophodon-Mammut Fortelius et al. 1996; Van der Made 1990). group were probably folivorous. Anancus This is probably the reason that suoid species may have evolved in the Palaearctic from diversity in Europe dropped markedly; in earlier migrants and acquired longer molars, lower Vallesian localities five to six suoid particularly the M3, and lost its premolars. species may be found together, but in youn- This suggests that it adapted to a diet poorer ger localities rarely more than one. This mar- in nutrients, so that greater amount of food ked the end of the Cainochoerinae, Listrio- had to be ingested. The proboscideans that dontinae and Tetraconodontinae in Europe, reached Europe between 2.7 and 0.8 Ma ago, three suid subfamilies that have their oldest had high-crowned cheek teeth, with antero- record in the tropics and it also was the end posteriorly compressed cusps, valleys filled for the European Schizochoerinae (ranges with cementum, thin enamel and elongate after Van der Made 1990; after Van molars, in particular elongate M3. These ani- der Made 1997; see also Fortelius et al. mals were grazers. This type of dentition 1996). All but the Listriodontinae lived on in evolved in Africa some 6 Ma ago. the tropics and the Tetraconodontinae became The browsing proboscideans entered very diverse there. Other groups were also Western Europe, when a large number of ani- affected, for instance European mainland pri- mals of (sub)tropical origins did so too. Up to mates went extinct. Around 7-8 Ma ago, the four species of Dorcatherium (Tragulidae) ente- last European tragulids and deinotheres went red Europe at the same time as Deino- extinct. Thus, many of the lineages of tropi- therium.Living tragulids tend to be frugivo- cal origins that entered Europe before 12.5 rous and to a lesser extend folivorous Ma ago went extinct after that date. It is like- (Kingdon 1989). The suid Bunolistriodon was ly that this is related to the effect of decre- folivorous (Hunter & Fortelius 1994; Van der asing temperatures and increasing seasonality Made 1996). The primates Griphopithecus, on the vegetation. It became probably Pliopithecus, Plesiopliopithecus and Dryo- progressively more difficult for frugivores pithecus were probably frugivorous and and folivores to come through the winter. folivorous. Tropical environments have the Vegetation with a C4 photosynthetic path- year round fruit and leaves, while environments way is favoured by low atmospheric CO2 with cold winters don’t. This may have been levels compared to that with a C3 pathway, a limiting factor to dispersal. Around 12.5 Ma and includes grasses. Carbon 13 isotope a number of animals went extinct in western records suggest that such vegetation started Europe, that either had migrated earlier becoming important at low latitudes between towards the north, or that were sensitive to 5 and 8 Ma ago, while this happened much temperature. Crocodiles went extinct in central later at high latitudes (Quade et al. 1989; Europe, tragulid species diversity dropped Cerling 1992; Cerling et al. 1993, 1997). from four to one, whereas diversity remained Carbon isotopes in tooth enamel show that high in the Indian Subcontinent, and the last deinotheres maintained a pure C3 diet, while members of the Bunolistriodon meidamon other shifted to a C4 dominated lineage, that had disappeared already around diet (Cerling et al. 1997). When the vegeta-

446 VAN DER MADE & MAZO: Proboscidean dispersals

tion changed in the tropics, proboscideans de l’Aragonien) d’Artesilla a Villafeliche (Saragos- there adapted to grazing and evolved hypsod- se), Bassin de Calatayud-Teruel (Espagne) - Bulletin ont cheek teeth, with thin enamel, a high du Muséum national d'Histoire naturelle, Paris, 4e number of plates, much cementum, etc. It is sér. 15, section C (1-4): 105-153 this adaptation that allowed proboscideans to Azzaroli, A., De Giuli, C., Ficarelli, G. & Torre, D., enter and survive in Europe and northern 1988 - Late Pliocene to Early Mid-Pleistocene Asia. The exact timing of their dispersals Mammals in Eurasia: faunal succession and dispersal coincides with important climatical changes, events - Paleogeography, Palaeoclimatology, suggesting that it were these changes that Palaeoecology 66: 77-100 triggered the dispersals. Barry, J.C. & Flynn, L.J. 1990 - Key biostratigraphic events in the Siwalik sequence - in: Lindsay, E.H., ACKNOWLEDGEMENTS Fahlbusch, V. & Mein, P. (eds.) - European Neogene Research by J. van der Made benefitted from Mammal Chronology - Plenum Press, New York & financial support from the DGICYT (project London: 557-571 numbers PB96-1026-C03-02 and PB98-0513) Beden, M., 1979 - Les eléphants (Loxodonta et Elephas) and the ‘Unidades Asociadas’ program. d’Afrique orientale: systématique, phylogénie, intérêt biochronologique - Thèse Doctoral des Sciences, REFERENCES Poitiers: 1-567 Andrews, C.W., 1904 - Further notes on the mammals of Bergounioux, F.M. & Crouzel, F., 1958 - Les mastodonts the of Egypt. Part I. - The Geological d’Espagne. - Estudios geológicos 14 (40): 224-343 Magazine, decade V, new series, I: 109-115 Berggren W.A. & Van Couvering, J.A., 1974 - The Late Andrews, C.W., 1906 - A descriptive catalogue of the Neogene. Biostratigraphy, geochronology and paleo Tertiary Vertebrata of the Fayûm, Egypt - British climatology of the last 15 million years in marine and Museum (Natural History). London continental sequences - Paleogeography, Andrews, P., Harrison, T., Delson, E., Bernor, R.L. & Palaeoclimatolgy, Palaeoecology 16 (1-2): 216 pp. Martin, L., 1996 - Distribution and biochronology of Bonifay, M.F., 1980 - Relations entre les données isoto- European and soutwhest Asian Miocen catarrhines - piques océaniques et l’histoire des grandes faunes in: Bernor, R.L., Fahlbusch, V. & Mittman, H.W. européennes plio-pleistocenes - Quaternary Research (eds.) - The evolution of western Eurasian Neogene 14: 251-262 mammals faunas - New York, Columbia University Bout, P., 1964 - Étude stratigraphique et paléogeographi- Press: 168-208 que du gisement de mammifères fossiles du Antunes, M.T., 1990 - The proboscideans data, age and Pleistocène moyen de Solilhac près de Puy-en-Velay paleogeography: evidence from the Miocene of (Haute Loire), France - Geologie en Mijnbouw 43: Lisbon - in: Lindsay, E.H., Fahlbusch, V. & Mein, P. 83-93 (eds.) - European Neogene Mammal Chronology - Bulot, Ch. & Ginsburg, L., 1993 - Gisements à mam- Plenum Press, New York & London: 253-262 mifères miocènes du Haut-Armagnac et âge des plus Antunes, M.T. & Mazo, A.V., 1978 - Les plus anciens anciens Proboscidiens d’Europe occidentale - mastodontes tétralophodontes (Langien inférieur Vb), Comptes Rendues de l'Académie des Sciences, Paris, évolution et remarques sur la tétralophodontie - 316, série II: 1011-1016 Ciencias da Tierra (Universidade Nova de Lisboa), Cerling, T.E., 1992 - Development of grasslands and Portugal, 4: 9-30 savannas in East Africa during the Neogene - Astibia, H. & Mazo, A.V., 1988 - Los Mastodontes Paleogeography, Palaeclimatology, Palaeoecology (Proboscidea, Mammalia) del Mioceno Medio de (Global and Planetary Change Section) 97: 241-247 Tarazona de Aragón (Zaragoza, España) - Estudios Cerling, T.E., Harris, J.M., MacFadden, B.J., Leakey, geológicos 44: 329-338 M.G., Quade, J., Eisenmann, V. & Ehleringer, J.R., Azanza, B., Cerdeño, E., Ginsburg, L., Van der Made, J., 1997 - Global vegetation change through the Morales, J. & Tassy, P., 1993 - La faune des grands Miocene/Pliocene boundary - Nature 389: 153-158 mammifères du Miocène Inférieur (MN 4, zone C Cerling, T.E., Wang, J. & Quade, J., 1993 - Expansion of

447 ADVANCES IN MAMMOTH RESEARCH DEINSEA 9, 2003

C4 ecosystems as an indicator of global ecological phants and their relatives - Oxford University Press, change in the late Miocene - Nature 361: 344-345 Oxford, New York & Tokyo:124-135 De Bruijn, H., Daams, R., Daxner-Höck, G., Fahlbusch, Guan Jian & Van der Made, J., 1993 - Fossil Suidae from V., Ginsburg, L., Mein, P., Morales, J., Heizmann, E., Dingjiaergou near Tongxin, China - Memoires of the Mayhew, D.F., Van der Meulen, A.J., Schmidt-Kittler, Beijing Natural History Museum 53: 151-199 N. & Telles Antunes, M., 1992 - Report of the Haq, B.U., Hardenbol, J. & Vail, P.R., 1987 - RCMNS working group on fossil mammals, Chronology of Fluctuating Sea Levels Since the Reisensburg, 1990 - Newsletters on Stratigraphy 26 - Science 235: 1156-1166 (2/3): 65-118 Hendey, Q.B., 1981 - Palaeoceology of the late Tertiary deMenocal, P.B., 1995 - Plio-Pleistocene African fossil occurrences in "E" Quarry, Langebaanweg, Climate - Science 270: 53-59 South Africa, and a reinterprtation of their geological Fitch, F.J. & Miller, J.A., 1976 - Conventional context - Annals of the South African Museum 84 Potassium-Argon and Argon-40 / Argon-39 (1): 1-104 dating of volcanic rocks from East Rudolf - in: Hunter, J.P. & Fortelius, M., 1994 - Comparative dental Coppens, Y., Howell, F.C., Isaac, G.L. & Leakey, occlusal mophology, facet development, and micro R.E.F. (eds.) - Earliest man and environments in the wear in two sympatric species of Listriodon Lake Rudolf Basin - University of Chicago Press, (Mammalia, Suidae) from the Middle Miocene of Chicago: 123-147 western Anatolia (Turkey) - Journal of Vertebrate Flower, B.P. & Kennett, J.P., 1993 - The middle Miocene Palaeontology 14 (1): 105-126 climatic transition: east antarctic ice sheet develop- Hussain, S.T., Van den Bergh, G.D., Steensma, K.D., De ment, deep ocean circulation and global carbon Visser, J.A., De Vos, J., Arif, M., Van Dam, J., cycling - Palaeogeography, Palaeoclimatology, Sondaar, P.Y. & Malik, S.B., 1992 - Biostratigraphy Palaeoecology 108: 537-555 of the Plio-Pleistocene continental sediments (Upper Fortelius, M., Van der Made, J. & Bernor, R.L., 1996 - Siwaliks) of the Mangla-Samwal Anticline, Azad Middle and Late Miocene Suoidea of central Europe Kashmir, Pakistan - Proceedings of the Koninklijke and the Eastern Mediterranean: Evolution, Nederlandse Akademie van Wetenschappen 95 (1): Biogeography and Palaeoeoclogy - In: Bernor, R.L., 65-80 Fahlbusch, V. & Rietschel, S. (eds.) - Later Neogene Kalb, J.E., Froehlich, J. & Bell, G.L., 1996 - Paleo- European biotic evolution and stratigraphic correla biogeography of late Neogene African and Eurasian tion - Columbia University Press: 348-377 Elephantoidea - in: Shoshani, J. & Tassy, P. (eds.) - Foucault, A., 1993 - Climat. Histoire et avenir du milieu The Proboscidea. Evolution and palaeoecology of terrestre - Fayard: 328 pp. and their relatives - Oxford University Gabunia, L.K., 1973 - The fossil vertebrate fauna from Press, Oxford, New York & Tokyo: 117-123 Belometchetskaia - ‘Metsniereba’, Tbilisi: 1-136 Kingdon, J., 1989 - East African Mammals, IIIB - Gabunia, L., 1981 - Traits essentiels de l’évolution des University of Chicago Press: 436 pp. Kingston, J.D., faunes de mammifères de la région Mer Noire- Marino, B.D. & Hill, A., 1994 - Isotopic evidence for Caspienne - Bulletin du Museum national d’Histoire Neogene Hominid paleoenvironments in the Kenya naturelle, Paris, 4e série, 3, section C, 2: 195-204 Rift Valley - Science 264: 955-959 Gaziry, A.W., 1976 - Jungtertiäre Mastodonten aus Kubiak, H., 1972 - The skull of Mammut praetypicum Anatolien (Türkei) - Geologisches Jahrbuch B22: (Proboscidea, Mammalia) from the collection of the 3-143 Jagiellonian University in Craców, Poland - Acta Ginsburg, L. & Antunes, M.T., 1966 - Considération sur Zoologica Cracoviensia 17: 305-324 les mastodontes du Burdigalien de Lisbonne et des Lehmann, U., 1950 - Über Mastodontenreste in der Sables de l'Orléanais (France) - Revista da Faculdade Bayerischen Staatssammlung in München - de Lisboa, 2nd series C 14: 135-150 Palaeontographica 99 (a): 121-228, plates 10-22 Guan Jian, 1996 - On the shovel-tusked elephantoids Lister, A., 1996 - Evolution and taxonomy of Eurasian from China - in: Shoshani, J. & Tassy, P. (eds.) - The - in: Shoshani, J. & Tassy, P., (eds.) - The Proboscidea. Evolution and palaeoecology of ele- Proboscidea. Evolution and palaeoecology of ele-

448 VAN DER MADE & MAZO: Proboscidean dispersals

phants and their relatives - Oxford University Press, Miller, K.G., Mountain, G.S., the Leg 150 Shipboard Oxford, New York & Tokyo: 203-213 Party & Members of the New Jersey Coastal Plain Lydekker, R., 1884a - Mastodon teeth from Perim Island Drilling Project, 1996 - Drilling and dating New - Memoirs of the Geological Survey of India, Jersey Oligocene-Miocene sequences: ice volume, Palaeontologia Indica (10), 3, part 5: 149-154 global sea level, and Exxon Records - Science 271: Lydekker, R., 1884b - Additional Siwalik Perissodactyla 1092-1095 and Proboscidea - Memoirs of the Geological Survey Mottl, M., 1969 - Bedeutende Proboscidier-Neufunde aus of India, Paleontologia Indica (10), 3, part 1: 1-34, 5 dem Altpliozän (Pannonien) Südost-Österreichs - plates Österreichische Akademie der Wissenschaften Madden, C.T. & Van Couvering, J.A., 1976 - The Mathematisch-Naturwissenschaftliche Klasse, Proboscidean Datum Event: Early Miocene migration Denkschriften 115: 1-22 from Africa - Geological Soci-ety of America, Osborn, H.F., 1936 - Proboscidea: a monograph of the Abstracts with Programs: 992 discovery, evolution, migration and description of the Maglio, V.J., 1970 - Four new species of mastodonts and elephants of the world. Vol. I: from the Plio-Pleistocene of northwestern Kenya - Moeriterioidea, Deinotherioidea, Mastodontoidea - Breviora, 341: 1-43 The American Museum Press, New York Maglio, V.J., 1978 - Patterns of faunal evolution - in: Pickford, M., 1981 - Preliminary Miocene mammalian Maglio, V.J. & Cooke, H.B.S. (eds.) - Evolution of biostratigraphy for western Kenya - Journal of human African Mammals - Harvard University Press, evolution 10: 73-97 Cambridge & London: 603-619 Pickford, M., 1987 - Concordance entre la paléontologie Maglio, V.J. & Hendey, Q.B., 1970 - New evidence continentale de l’est africain et les événements paléo- relating to the supposed stegolophodont ancestry of oceanographiques au Néogene - Comptes Rendus de the Elephantidae - South African Archaeolocial l’Académie des Sciences, Paris 304: 675-678 Bulletin 25 (3-4): 85-87 Pickford, M., 1988 - The age(s) of the Bugti fauna(s), Matsumoto, H., 1924 - A revision of Palaeomastodon Pakistan - in: Whyte, P., Aigner, J.S., Jablonski, N.G., dividing it into two genera and with desciptions of Taylor, B., Walker, D., Wang Pinxian & So Chaklam two new species - Bulletin of the American Museum (eds.) - The palaeoenvironment of east Asia from the of Natural History 50: 1-58 Mid-Tertiary - Centre of Asian Studies, University of Mazo, A.V., 1977 - Revision de los mastodontes de Hong-Kong, Hong-Kong: 937-955 España - Universidad Complutense de Madrid Pickford, M. & Andrews, P., 1981 - The Tinderet Mazo, A.V., 1985 - Proboscideos terciarios de Madrid y Miocene sequence in Kenya - Journal of Human su provincia - in: Geología y paleontología del Evolution 10: 11-33 Terciario continental de la provincia de Madrid - Pickford, M. & Morales, J., 1994 - Biostratigraphy and Consejo Superior de Investigaciones Científicas, palaeobiogeography of East Africa and the Iberian Madrid: 41-46 Peninsula - Palaeogeography, Paleoclimatology, Mazo, A.V., 1989 - Nuevos restos de Proboscidea Palaeoecology 112: 297-322 (Mammalia) en la cuenca de Guadix-Baza - Trabajos Pilgrim, G.E., 1908 - Tertiary and post-Tertiary fresh- sobre el Neogeno-Cuaternario 11: 225-237 water deposits of Baluchistan and Sind, with notices Mazo, A.V., 1996 - and mammuthids of new vertebrates - Records of the Geological from the Iberian Peninsula - in: Shoshani, J. & Tassy, Survey of India 37: 139-166 P. (eds.) -The Proboscidea. Evolution and palaeoeco- Quade, J., Cerling, T.E. & Bowman, J.R., 1989 - logy of elephants and their relatives - Oxford Development of Asian monsoon revealed by marked University Press, Oxford, New York & Tokyo: ecological shift during the latest Miocene in northern 136-142 Pakistan - Nature 342: 163-166 Mein, P., 1990 - Updating of MN zones - in: Lindsay, Qiu Zhangxiang, Wu Wenyu & Qiu Zhuding, 1999 - E.H., Fahlbusch, V. & Mein, P. (eds.) - European Miocene mammal faunal sequence of China: palaeo- Neogene Mammal Chronology - Plenum Press, New zoogeography and Eurasian relationships - in: York & London: 73-90 Rössner, G. & Heissig, K. (eds.) - The Miocene land

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mammals of Europe Verlag Dr. Friedrich Pfeil, Stehlin, H.G., 1926 - Über Milchincisiven miocäner München: 443-455 Proboscidier - Eclogae Geologae Helvetiae 19 (3): Radulescu, C. & Samson, P.M., 1994 - Review of the 694-700 "Villafranchian" (s.s.) faunas of Romania - AIQUA- Takai, F., 1954 - An addition to the mammalian fauna of CNR Il significato del Villafranchiano nelle succes the Japanese Miocene - Journal of the Faculty of sioni Plio-Pleistoceniche - Peveragno (CN)- Sciences, University of Tokyo, section II, 9: 331-335 Villafranca (AT), 20-24 june 1994 Tassy, P., 1977a - Les mastodontes miocènes du Bassin Rögl, F., 1998 - Palaeogeographic considerations for Aquitain: une mise au point taxonomique - Comptes Mediterranean and Paratethys seaways (Oligocene to Rendus de l’Académie des Sciences Paris, 284, série Miocene) - Annalen des Naturhistorischen Museums D: 1389-1392 in Wien 99A: 279-310 Tassy, P., 1977b - Découverte de Zygolophodon turicen- Rögl, F. & Steinginger, F.F., 1983 - Vom Zerfall der sis (Schinz) (Proboscidea, Mammalia) au lieu dit Tethys zu Mediterran und Parathethys - Annalen des Malartic à Simorre, Gers (Vindobonien Moyen); Naturhistorischen Museums Wien 85A: 135-163 implications paléoecologiques et biostratigraphiques - Roebroeks, W. & Van Kolfschoten, T., 1995 - The Geobios 10 (5): 655-696 earliest occupation of Europe: a reappraisal of arte- Tassy, P., 1983 - Les Elephantoidea miocènes du Plateau factual and chronological evidence - in: Roebroeks, du Potwar, Groupe de Siwalik, Pakistan - Annales de W. & Van Kolfschoten, T. (eds.) - The earliest occu- Paléontologie 69: 99-136, 235-297, 317-354 pation of Europe - University of Leiden: 297-315 Tassy, P., 1984 - Le mastodonte a dents étroites, le grade Sardella, R., Caloi, L., Di Stefano, G., Palombo, M.R., trilophodonte et la radiation initiale des Petronio, C., Abazzi, L., Azzaroli, A., Ficcarelli, G., Amedelodontidae - in: Buffetaut, E., Mazin, J.M. & Mazza, P., Mezzabotta, C., Rook, L., Torre, D., Salmon, D. (eds.) - Actes du symposium paléontolo- Argenti, P., Capasso Barbato, L., Kotsakis, T., gique Georges Cuvier - Impressions le Serpentaire, Gliozzi, E., Masini, F. & Sala, B., 1998 - Mammal Montbéliard: 459-473 faunal turnover in Italy from the Middle Pliocene to Tassy, P., 1985 - La place des mastodontes miocènes de the Holocene - Mededelingen Nederlands Instituut l’ancien monde dans la phylogénie des Proboscidea voor Toegepaste Geowetenschappen TNO 60: 499- (Mammalia): hypothèses et conjectures - Unpubished 511 PhD thesis, Université Pierre et Marie Curie, Paris, Schinz, H.R., 1824 - Naturgeschichte und Abbildingen volumes 1-3 der Säugethiere - Brodtmanns Lithographischer Tassy, P., 1986 - Nouveaux Elephantoidea (Proboscidea, Kunstanstalt, Zürich: 417 pp. & Atlas: 177 pls Mammalia) dans le Miocène du Kenya: essai de ré- Schlesinger, G., 1912 - Studien über die Stammes- évaluation systématique - Cahiers de Paléontologie. geschichte der Proboscidier - Jahrbuch der Éditions du Centre National de la Recherche Kaiserlich-Königlichen Reichsanstalt 62: 87-182 Scientifique (CNRS), Paris Schlesinger, G., 1917 - Die Mastodonten des K.K. Tassy, P., 1988 - Le statut systématique de l’espèce Naturhistorischen Hof-Museums - Denkschriften der Hemimastodon crepusculi (Pilgrim): l’éternel problè- K.K. Naturhistorischen Hofmuseums, 1, Geologisch - me de l’homologie et de la convergence - Annales de Paläontologische Reihe 1: 1-230 Paléontologie 74 (3): 115-126 Sesé,C., Alberd, M.T., Mazo, A. & Morales, J., 2001 - Tassy, P., 1990 - The "Proboscidean Datum Event": How Mamíferos del Mioceno, Pliòceno y Pleistoceno de la many proboscideans and how many events?- in: Cuenza de Guadix-Baza (Granada, España): revisión Lindsay, E.H., Fahlbusch, V. & Mein, P. (eds.) - de las asocianones faunistica más características - European Neogene Mammal Chronology - Plenum Paleontologia i Evolucio 32-33: 31-26 Press, New York & London: 237-252 Steensma, K.J. & Hussain, S.T., 1992 - Caprolagus Tassy, P., 1994 - Les gisements de mammifères du sivalensis (Lagomorpha, Mammalia) from the Upper Miocène supérieur de Kemiklitepe, Turquie: 7. Siwalik Subgroup of Azad Kashmir, Pakistan - Neues Proboscidea (Mammalia) - Bulletin du Muséum Jahrbuch für Geologie und Paläontologie, National d’Histoire Naturelle, Paris, series 4, 16, Monatshefte 7: 429-434 section C: 143-157

450 VAN DER MADE & MAZO: Proboscidean dispersals

Tassy, P., 1996 - The earliest gomphotheres - in: Tobien, H., Chen Guanfang & Li Y.Q., 1988 - Shoshani, J. & Tassy, P. (eds.) - The Proboscidea. Mastodonts (Proboscidea, Mammalia) from the late Evolution and palaeoecology of elephants and their Neogene and early Pleistocene of the People's relatives - Oxford University Press, Oxford, New Republic of China. Part 2. The genera Tetralophodon, York & Tokyo: 89-91 Anancus, , Zygolophodon, Tassy, P. & Pickford, M., 1983 - Un nouveau mastodonte Mammut, . Some generalities on the zygolophodonte (Proboscidea, Mammalia) dans le Chinese mastodonts - Mainzer Geowissenschaftliche Miocène inférieur d’Afrique orientale: systemátique Mitteilungen 17: 95-220 et paléoenvironnement - Geobios 16 (1): 53-77 Van Dam, J.A., 1997 - The small mammals from the Thomas, H., 1985 - The Early and Middle Miocene Land Upper Miocene of the Teruel-Alfambra region Connection of the Afro-Arabian Plate and Asia: A (Spain): paleobiology and paleoclimatic reconstruc- Major Event for Hominoid Dispersal? - in: tions - Geologica Ultraiectina 156: 204 pp. Ancestors: The Hard Evidence: 42-50 Van der Made, J., 1992 - Migrations and climate - Tobien, H., 1972 - Status of the genus Serridentinus Courier Forschungsinstitut Senckenberg 153: 27-39 Osborn 1923 (Proboscidea, Mammalia) and related Van der Made, J., 1993 - Artiodactyla and the timing of a forms - Mainzer Geowissenschaftliche Middle Miocene climatical change - Premier Mitteilungen 1: 143-191 Congrès Européen de Paléontologie, Lyon, 7- Tobien, H., 1973 - On the evolution of mastodonts 9/7/1994: 128 (Proboscidea, Mammalia). Part 1: The bunodont tri- Van der Made, J., 1996 - Listriodontinae (Suidae, lophodont groups - Notizblatt des Hessischen Mammalia), their evolution, systematics and distribu- Landesmtes für Bodenforschung zu Wiesbaden 101: tion in time and space - Contributions to Tertiary and 202-276, plates 23-26 Quaternary Geology 33 (1-4): 3-254, microficha 54 pp. Tobien, H., 1976 - Zur paläontologischen Geschichte der Van der Made, J., 1997 - Intercontinental dispersal Mastodonten (Proboscidea, Mammalia) - Mainzer events, eustatic sea level and Early and Middle Geowissenschaftliche Mitteilungen 5: 143-225 Miocene Stratigraphy - Mémoires et Travaux EPHE, Tobien, H., 1978 - On the evolution of mastodonts Montpellier 21: 75-81 (Proboscidea, Mammalia). Part 2: The bunodont Van der Made, J., 1999a - Biometric trends in the tetralophodont groups - Geologisches Jahrbuch Tetraconodontinae, a subfamily of pigs - Transactions Hessen 106: 159-208, plates 10-21 of the Royal Society of Edinburg, Earth Sciences 89: Tobien, H., 1980 - A note on the skull and mandible of a 199-225 new choerolophodont mastodont (Proboscidea, Van der Made, J., 1999b - Intercontinental relationship Mammalia) from the middle Miocene of Chios Europe-Africa and the Indian Subcontintent - in: (Aegean Sea, Greece) - in: Jacobs, L.L. (ed.) - Rössner, G. & Heissig, K. (eds.) - The Miocene land Aspects of vertebrate history. Essays in honor of mammals of Europe - Verlag Dr. Friedrich Pfeil, Edwin Harris Colbert - Museum of Northern Arizona München: 457-472 Press, Flagstaff: 299-307 Van der Made, J. 2001 - Les ongulés d’Atapuerca. Tobien, H., 1996 - Evolution of zygodons with emphasis Stratigraphie et biogéographie - L’Anthropologie 105 on dentition - in: Shoshani, J. & Tassy, P. (eds.) - The (1): 95-113 Proboscidea. Evolution and palaeoecology of ele- Van der Made, J. & Mazo, A.V., 2001 - Spanish phants and their relatives - Oxford University Press, Pleistocene proboscidean diversity as a function of Oxford, New York & Tokyo: 76-85 climate - in: Cavarretta, G., Gioia, P., Mussi, M. & Tobien, H., Chen Guanfang & Li Y.Q., 1984 - The Palombo, M.R. (eds.) - The world of elephants. mastodonts (Proboscidea, Mammalia) of China: Proceedings of the 1st international congress. evolution, palaeobiogeography, palaeoecology - in: Consiglio Nazionale delle Ricerche, Roma: 214- 218 Whyte, R.O. (ed.) - The evolution of the east Asian Van der Meulen, A.J. & Daams, R., 1992 - Evolution of envrionment II: paleoboany, plaeozoology, and palae- Early-Middle Miocene rodent faunas in relation to oanhroplogy - Centre or Asian Studies, University of long-term palaeoenvironmental changes - Hong Kong: 689-696 Palaeogeography, Palaeoclimatology, Palaeoecology

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93: 227-252 Zachos, J.C., Flower, B.P. & Paul, H., 1997 - Orbitally Vislobokova, I., 1994 - The Lower Miocene artiodactyls paced climate oscillations across the Oligocene/ of Tagai Bay, Olhon Island, Lake Baikal (Russia) - Miocene boundary - Nature 388: 567-570 Palaeovertebrata, Montpellier 23 (1-4): 177-197 Zapfe, H., 1957 - Ein bedeutender Mastodon-Fund aus Vrba, E.S., 1995 - The fossil record of african antelopes dem Unterpliozän von Niederösterreich - Neues (Mammala, Bovidae) in relation to human evolution Jahrbuch für Geologie und Paläontologie 104: 382- and paleoclimate - in: Vrba, E.S., Denton, G.H., 406 Partridge, T.C. & Burckle, L.H. (eds.) - Paleoclimate and evolution, with emphasis on human origins - Yale received 24 August 1999 University Press, New Haven and London: 385-424

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