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Home , Albanohyus, Aureliachoerus, Enhydriodon, Giraffokeryx, Herpestides, Indian subcontinent

Intercontinental relationship - and the Indian Subcontinent 45 Jan van der Made*

A great number of genera, and even Palaeogeography, global climate some , are cited or described from both Europe and Africa and/or the Indian Subconti- nent. In other cases, an ancestor-descendant re- After MN 3, Europe formed one with lationship has been demonstrated. For most of . This land mass extended from Europe, the Miocene, there seem to have been intensive through to China and SE Asia and is faunal relationships between Europe, Africa and here referred to as . This term does not the Indian Subcontinent. This situation may seem include here SE Europe. At this time, the Brea normal to uso It is, however, noto north of Crete was land and SE Europe and During much of the Tertiary, Africa and formed a continuous . The Para- were isolated . There were some peri- tethys was large and extended from the valley of ods when faunal exchange with the northern the Rhone to the Black , and continents occurred, but these periods seem to further to the east. The Tethys was connected have been widely spaced in time. During a larga with the Indian and large part of the Middle part of the and during the earliest East was a shallow sea. During the earliest Mio- Miocene, Africa and India had been isolated. En- cene, Africa and Arabia formed one continent that demic faunas evolved on these continents. Fam- had been separated from Eurasia and India for a ilies that went extinct in the northern continents considerable time. Similarly the India n Continent during the Paleogene, tlorished in these southern had been isolated for a very long period (RÓGL & continents and other familias originated. Also in STEINIGER1983). Due to this long isolation, the the northern continents evolution went on. The southern continents had endemic faunas. result was that the northern and southern conti- During the Miocene, the African and Indian nents had totally different faunas. plates moved closer to Eurasia causing the fur- Then happened what MADDEN & VAN COUVER- ther elevation of the Alps, and other ING (1976) named the "Proboscidean Datum mountain belts. The Paratethys and Tethys be- Eventn. They apparently believed that it was a carne smaller and extensive Breas in the Middle single synchronous faunal exchange event be- East beca me shallow and finally beca me tween Africa and Eurasia, involving many taxa, land. The area of SE Europe beca me connected including proboscideans. THOMAS (1985) recog- to Eurasia, while the collapse of the ares north of nized two majar phases of faunal exchange. T ASSY Crete separated it from Anatolia. The Sea (1990) showed that the dispersal of the probosci- formed, and Arabia became disconnected from deans was diachronous. The UDatum Eventn turns Africa. During much of the Miocene, India may out to be a complex of events (VAN DERMADE have had a contact with Asia at the place of the 1996, 1997b). These events took place during the present Himalayas, but there is little evidence of Early and Middle Miocene. Faunal exchange be- faunal exchange through this ares. The Parateth- tween Eurasia and Africa during the Late Miocene ys was another barrier. Faunal exchange between is not well documented; the African record is very northern Asia and the southern continents may incomplete. The Indian record is better for the have occurred through the ares between Himala- Late Miocene, but nevertheless, relationships be- yas and Paratethys. tween the faunas are not very clear. The events While plate tectonics gradually changed the around the latest Miocene Messinian Salinity Cri- , sea level fluctuations had more direct sis are again well studied. effects. Sea level changes were, geologically J speaking, instantaneous, and had a magnitude of up to a hundred metres and occasionally succes- sive events had accumulative effects (HAQ et al. 1987). Between 15 and 10.5 Ma ago, sea level dropped in three successive events ayer two * Dr. Jan van der Made, Museo Nacional de Ciencias hundred metres (HAQ et al. 1987). Sea level chang- Naturales, José Gutiérrez Abascal, 2, E-2BOO6Ma- e8 were relaled 10 global climale; Ice accumula- drid, Spain tion on Antartica was responsible for sea level

The Mk)cen8 LAnd Memmals ~ E¡wope. - Pp. 4&7-472. 457 drops (MILLER et al. 1996). While extensive land Stratigraphy masses were slowly being formed out of a number of smaller fragments, sea level changes connect- MN units or Units are widely ed and disconnected these Breas and allowed for, usad for Europe and northern Africa (DE BRUIJNet or inhibitted faunal exchange. The effects of the al. 1992). The MN units do not have a clear defi- sea level changes were strongest during the Early nition of the boundaries. It appears that global Miocene, whereas from the Middle Miocene on- events, causing majar dispersal or ward, the land-sea distribution seems to have events, tend to be within MN units, whereas the been less affected and less important in control- transition of one unit to the other is diffuse. Here ling the geographical distribution of . frequently informal reference will be made to Changes in global climate had a direct impact MN 4e (early MN 4), MN 41 (late MN 4) etc. to on the distribution of species: there are distinct indicate the part of the MN unit before or after an dispersal and extinction events. When, during the important dispersal evento In the same way MN 8 Early Miocene, global climate lowered the sea may be used instead of MN 7+8. In addition level, and allowed for faunal exchange between reference is made to continental stages and their Africa and Europe, global climate siso permitted biozones: Ramblian (zones Z-A), Aragonian (zones Asian to disperse into Europe. Late Mio- B-G), Vallesian (H-I) and Turolian (J-M) (DAAMS& cene global climate, may have had effects on sea FREUDENTHAl1988, VAN DAM 1997). Though these level, but its effect on the distribution of animals zones are defined with local (Spanish) fauna, the was directo fact that the local faunal events used tend to Faunal exchange between Europe and Africa coincide with global events, makes a more or less and the Indian Subcontinent occurred through SE precise correlation with a much larger Brea possi- Europe and Anatolia. Miocene climates do not ble. A number of long profiles, frequently in or necessarily have recent equivalents. Neverthe- correlated to type sections, were sampled for less, certain characters may have been the same: palaeomagnetism in arder to date biozones and higher temperatures towards the equator, the MN units (KRIJGSMANet al. 1996). These dates are climate of northern Eurasia may always have used here, but differ considerably in the MN 4-6 been more seasonal with respect to temperature range as used by for instance STEININGERet al. than the central African climate, etc. As we will (1996). see in the following sections, the distribution of For Africa a series of "faunal sets" were de- animals that may have been dependent on Borne fined (PICKFORD1981). Like the Spanish zones, degree of humidity suggests, that the distribution fuese sets tend to capture global events. Many of more and less humid climates during a large African localities are radiometrically datad (PICK- part of the Miocene may have shown only a FORD 1986b). limited variation. In general, the conditions in SE The best record for the Indian Subcontinent is Europe and Anatolia, may have acted as a filler to in . A series of "faunas" were described faunal exchange. There is a well studied palien from Pakistan, later the names of these "faunas" record in Anatolia, that is correlated to the were applied to the formations from which they MN scale (BENDA& MEULENKAMP1990); there were were collected and still later, the names have important changes in vegetation. Both reasons been applied as stages: Bugti, Murree, Kamlial, may explain why certain laxa, did not take part in Chinji, Nagri, Dhok Pathan (SHAH1984), most of one faunal exchange event, but were very prom- which have been palaeomagnetically sampled inent in a later evento and which yielded detailed range charts (BARRY& A peculiar event took place at the end of the FLYNN1990). Collections of older from the Miocene. The connection between the Mediterra- Bugti Brea, include fossils from several levels nean and the was lost. Evapora- (PICKFORD1987), but the bulk of the material is tion in the Mediterranean is greater than the input probably from more or less the same age anterior through precipitation and rivers. The sea level fell Murree. by up to several kilometres, and Europe and Asia A framework of faunal correlations and the beca me connected, allowing for an exchange of study of intercontinental dispersal events were terrestrial animals. Since this event caused the matched to continental-marine tie-points, palaeo- formation of extensive salt deposits, it is known magnetic data and radiometric dates, and finally as the Messinian Salinity Crisis (HSü et al. 1977). to the eustatic sealevel curves of HAOet al. (1987) (VAN DER MADE 1996, 1997b).Table 45.1 repre- sents stratigraphic data and intercontinental dis- persals of mammals.

458 In this paper, reference will be made to bio- the latest Oligoceneand earliestMiocene fauna is stratigraphic units and ages, even if an original not very well known. Similarly,the record of description of a taxon does not give these data. mammals prior to MN 6 is very restricted in SE The references in this section will usually provide Europe and Anatolia, and prior to MN 5 virtually the link. If the presence of a taxon in Europe in a absent, though there is a limitad number of mi- particular MN unit is mentioned, but no citation is cromammal localities. given, the source is usually DE BRUUN et al. (1992). Insectivora: Insectivores were present in the Paleogene of Africa, but some new ar- rived during the Early Miocene (BUTlER 19788). Taxa taking part in faunal exchange Several genera of Erinaceidae, presumably of Asian origins, are first cited from Africa (BUTlER As we have seen above, there have been many 19788) from localities that are placed in Set I and dispersals from Eurasia to the southern conti- Set 11.Soricidae have been cited from Set 11and nents as well as in the other direction. When Set IV localities from Africa, but there are impor- genera or species dispersed from Africa or the tant hiatuses in the record; the origin of these Indian Subcontinent to Europe, nearly always the dispersal routes were through the , soricids seemsto have been Asia (BUTLER1978a). Talpidae were present in Europe during the Mio- Anatolia and SE Europe. If a or species was cene, but did not disperse to the south of present in Africa and in the Indian Subcontinent Anatolia; they did not enter Africa, nor the Indian and dispersed to Europe, it is frequently not clear Subcontinent. They were presumably limited by where the colonizing populations carne from. In the extension of "desert soils" (dry soils with a any case, the latest non European populations are likely to have lived in Anatolia, whether the dura- high pH) (VAN DER MADE 1992). tion was geologically significant or noto Similarly, Chiroptera: Chiroptera are already known from it frequently cannot be established if a dispersal the European Paleogene. The record from Anato- joto the southern continents originated from Eu- lis is very incomplete (ENGESSER& ZIEGLER 1996). rape or Asia. They have entered Africa prior to the Miocene, There are many examples of laxa that origi- but during the Early Miocene and again during nated in some part of Asia and that migrated at the Late Miocene various families of Asian origins the same moment, or at different moments, to entered Africa (BUTLER19788). Europe and to the southern continents. The pres- ence of such a taxon in these areas does not Primates: The oldest European hominid (or hom- necessarily mean that there has been a faunal inoid in the older classifications) remain is from exchange between Europe and Africa or the Indi- Engelswies and is tentatively placed in Gríphopí- an Subcontinent. thecus by ANDREW5et al. (1996) who also assume At any moment in palaeontological history, a clase relationship between Gríphopíthecus and most of the present species are separated from Kenyapíthecus wíckerí. Gríphopíthecus is well other species by clear morphological and metri- known from MN 6e and MN 61 in Anatolia and cal differences. Taking time into account, there is from MN 61 and MN 7 +8 in . a continuum in morphology and size. Plíopíthecus and Plesíoplíopíthecus are likely subdivides this continuum in species, genera and to have African ancestors, but may have lived other laxa. The limits between these laxa are some time in SE Europe and Anatolia, before frequently taken at convenient places: at a hiatus entering Europe in MN 51. in the record, at a moment when evolution went Dryopíthecus entered Europe before MN 9, more rapidly, or at a long distance dispersal evento though the older record is rather poor. There is no Though the arrival in a new environment may evidence that it lived for a longer period outside accelerate evolution, the first individuals in the Africa before entering Europe. It may have en- new continent are identical to the populations in tered Europe at the same time as the little su id the continent of origino The decision to place the Albanohyus, which has an African record imme- populations that dispersed to the other continent diately previous to the middle MN 6 dispersal in a different species, genus or tribe, is practical, evento but disguises the relationships. Graecopíthecus (= Ouranopíthecus) is well In the Indian subcontinent there is virtually no known from MN 10-11 in Greece and is placed in Olígocene and earliest Miocene record. It is here the Homininae (ANDREW5et al. 1996). assumed that most of the uBugti faunau arrived by Consensus seems to be growing that Oreo- migration, this however is not sure. Also in Africa, píthecus is a hominid (ANDREW5et al. 1996). The

459 Tab.45.1. Faunaldispersals in the different (sub)continents.The left collumn gives eustatic sea level cycles of HAOet al. (1987).Four pairs of collumnsgive the stratigraphy(unit or biozoneand the age of its lower boundaryin Ma; selectedlocalities with dates in Ma or correlationto the Blow zones;see sectionon stratigraphy)and the taxa that enter in a particularunit. Thin horizontallinesindicate (predominantly local) stratigraphy, thick horizontallinesmark the dispersalevents discussed in this paper.The presumedor possibleorigin of the taxa is indicatedwith the following abbreviations: Af = Africa, Am = America, As = Asia, EA= Eurasia,Eu = Europe,ET = the SE Europeanand Anatolian Subcontinent,In = Indian Subcontinent,ME = Middle East.Question marks may indicate doubts on the origin, or on whether this taxon enterad in this unit.

460 Tab. 45.1. (continued).

genus is endemic at the Miocene island of Tusca- from SE Europe or directly from Africa (during the ny. In view of the fact that its affinities are not Messinian Crisis). perfectly clear, it is equally possible that it carne The cercopithecid Macaca is first citad in Eu- on the island from Africa, SE Europe and Anatolia rape from MN 13 in Spain (ANDREWSet al. 1996). or from Europe. The study of the possible origins The genus is siso known from Sardinia. It seems of other animals on the Tuscany island might likely, that Macaca reached Sardinia during the shed light on its afea of origino Messinian crisis (MN 13), though there is no early The cercopithecid Mesopithecusdecreased in record of the genus. size from MN 11 (M. de/soni) to MN 12 (M. pen- te/icus), to MN 13 (M. cf. monspessu/anus) (DE Creodonta: Creodonts are a primitive arder of BONISet al. 1990). The earliest citations are from ,that declined in Eurasia,but that re- MN 11 in SE Europe and in Maragheh, Iran and mained abundantin Africa. In MN 3 the creodont the earliest citations in Europe are from MN 13 Hyainailouros dispersed to the Indian Subconti- (ANDREWSet al. 1996). This suggests, that the nent and it is first known from early MN 4 in entry in Europe is a later dispersal evento either Europe. 461 : Carnívora are not known from the seems to be closely related to Pseudaelurus, Paleogene of Africa, starting in the Early Miocene which is cited from Europe from MN 3 (FRAilEet they entered in several waves. Carnivores were al. 1997). It seems likely that both arrived by probably absent in the earliest Miocene of the dispersal from some Asian centre of evolution. Indian Subcontinent. There is no a priori reason The same may be the case with Felis which is to suppose that no European carnivores may known from MN 13 in Spain and from equally cid have entered Africa and the Indian Subcontinent, deposits in Africa (FRAilE et al. 1997, HENDEY though SCHMIDT-KITTlER(1987) assumed that the 1981). carnivores entering Africa were of Asian origins. The Amphicyonidae Cynelos, Amphicyon and Tubulidentata: The origin of the Tubulidentata is Ysengrinia were already present in Europe during in Africa. The earliest aardvarks outside Africa are the earliest Miocene and are cited from Africa, from MN 6e in Anatolia (FORTELlUS1990) and from respectively from Set 1, Set lila and again Set lila the Chinji Formation in Pakistan (PICKFORD1978) (SCHMIDT-KITTlER 1987; MORALES et al. in press). and are sIl assigned to the genus Orycteropus. Amphicyon is cited from Bugti (PILGRIM1912). The more or less similar age suggest a single The first appearance of the ursid Hemicyon in dispersal evento Orycteropus may have been MN 3 in Europe and in Set II in Africa (SCHMIDT- present in SE Europe and Anatolia till the Upper KITTLER 1987)may well be a single dispersalevent Miocene (it is very well known from Pikermi and and the same is the case for the ursid Agrio- Sarnas, but it is found in many other localities) therium, which appeared apparently simultane- and may have dispersed in MN 13 to western ously in Europe in MN 13 (MORALES1984) and Europe (Brisighella). Africa (HENDEY 1981). ALFÉREZet al. (1988) mentioned a tubuliden- The oldest record of the in Africa is tefe from C6rcoles (MN 4, Spain), but PICKFORD Luogale from Set 11(SCHMIDT-KITTlER 1987). This (1993) claimed it is a suoid. genus seems to be closely related to Parsgsle from MN 2 in Europe. Lutrinae (Kenyalutra) seem : Proboscidea are known from the to have appeared even eartier in Africa (Set 1)than Eocene of the Indian Subcontinent and Africa and in Europe (MN 4) (SCHMIDT-KITTLER1987; WILLEM- may have originated from a Chinese SEN1990) and both dispersals seem to be inde- genus (WELLS & GINGERICH1983). However, by the pendent events. In the latest Miocene, Plesiogulo Early Miocene there was nothing similar to a and Enhydriodon appeared in Africa (HENDEY1981). proboscidean in Eurasia, whereas proboscideans Plesiogulo had a previous record in Spain (FRAILE had radiated in Africa. When, during the Early et al. 1997) and the lutrine Enhydriodon had a Miocene, Africa beca me connected after a long previous record in the Indian Subcontinent and time with Eurasia and the Indian Subcontinent, a Europe (WILLEMSEN 1990). diverse proboscidean fauna dispersed. Hemimas- and Herpestidae had a record in todon, Gomphotherium and Choerolophodon are Europe previous to the arrival of Kichechia, Lep- present in the Bugti fauna (T ASSY 1990). Possibly toplesictis and Mioprionodon in Africa in Set I and a little later Gomphotherium and Zygolophodon Herpestides in Set 11(FRAILE et al. 1997, SCHMIDT- appeared in Europe, both are certainly present in KlTTLER 1987). Viverra with a previous record in MN 4e and are even claimed to be present in Europe appeared in the latest Miocene of Africa MN 3 (BULOT& GINSBURG1993). Anancus appeared (HENDEY 1981). in the Late Miocene of Europe and Africa, it is Stenoplesictidae (= Percrocutidae) probably however not sufficiently clear where this genus had a pre-Miocene origin in Asia. Stenoplesictis, originated (A. V. MAZO pers. comm.). with felid affinities, first appeared in Africa in Set The lophodont cheek teeth of Deinotherium I and Africanictis, with hyaenid affinities, in Set suggest that it had a specialized diet. Deino- lila. In the afea of SE Europe and Anatolia Steno- therium (or Prodeinotherium) is known from Bug- plesictidae appeared in MN 6 and in western Eu- ti. It reached Europe during the later part of MN 4 rape in MN 7 (MORALESet al. 1998, WERDELlN and never really entered in China. The later and 1996). more limited dispersal of this proboscidean seems The hyaenid Protictitherium is found in Eu- to be due to its ecological requirements. rape in Bézian (MN 4) (WERDELlN& SOLOUNIAS 1996). The earliest African hyaenids appeared in Hyracoidea: The oldest known are from Set IV (SAVAGE 1978). the African Oligocene (MEYER1978). The oldest The machairodontine felid Afrosmilus first ap- known hyraxesoutside Africa are a form clase to peared in Africa Set I (SCHMIDT-KITTLER1987) and Geníohyus from Bugti (PICKFORD1986c) and P/ío-

462 from Pasalar in Anatolia (FORTEUUS1990), to Eurasia (VAN DER MADE 1996). Melambes on Crete and Can Uobateres and other Cainochoerinae (Kenyasus), of unkown origin, Vallesian European localities (PICKFORDet al. 1997, entered in Set I Africa. Albanohyus is first known FISCHER& HEIZMANN1992). For a long time Mela- from Set IV in Africa and later from MN 7-9 in mbes was believed to be Vallesian, but it seems Europe (VAN DER MADE 1996b). much more likely the locality is MN 6 (VAN DER "Conohyus" sindiense from MN 41 equivalent MADE 1996). This gives the impression of a first strata in the Indian Subcontinent is the oldest dispersal towards the Indian Subcontinent in MN 3 tetraconodontine known and gave rise to Cono- and a subsequent dispersal towards SE Europe hyus that dispersed into Europe in MN 51. During and Anatolia early in MN 6 and subsequent dis- the Vallesian this genus dispersed to Africa, where persal towards in MN 91. it gave rise to Nyanzachoerus. "Conohyus" sindi- ense evolved into Sivachoerus in the Indian Sub- Perissodactyla: Tapirs were present in Eurasia continent and dispersed possibly twice to Africa during the Oligocene and Miocene and the equid during the late Miocene, giving rise to severa I Anchitherium during the Aragonian and Early species of "Nyanzachoerus" (VAN DER MADE in Vallesian, but they did not disperse into Africa and press). the Indian Subcontinent. In the Vallesian, Hippari- In MN 8 Propotamochoerus (Suinae) dispersed on, also ot American origins, dispersed into Eur- either from Asia or the Indian Subcontinent to asía and immediately siso into the southern con- Europe. Hippopotamodon (= Microstonyx), prob- tinents. ably of Asian origin, appeared in MN 9 in Europe Chalicotheriidae have their origins in the north- and in Anatolia and in the Nagri Formation in ern hemisphere and were represented in the ear- Pakistan. Suinae did not enter Africa during the liest Miocene ot Europe with Schizotherium. When Miocene. the first connection occurred between Africa the northern and southern continents, Cha/icotherium Hippopotamoidea: Brachyodusof African origin moved into Africa and the Indian Subcontinent. dispersedin MN 3 to Eurasiaand the Indian Sub- At this stage, Cha/icotherium was not yet present continent. Hippos evolved in Africa from anthra- in Europe; it moved into Europe, when the tirst cotheres.Hexaprotodon dispersed during the late deinotheres carne into Europe. Phy//oti//on is cit- Miocene to the Indian Subcontinentand a tetra- ed from Bugti and may have given rise to Ancy- protodont hippo (placed either in Hexaprotodon /otherium. This group or lineage entered Europe or Hippopotamus)migrated in MN 13 to Europe not later than the Middle Miocene and Africa (VAN DER MADE this volume). during the Late Miocene (BUTlER 1978b). The group is not well known so that the timing of the Tragulidae: Tragulidae are primitive ruminants dispersals remains dubious. and have their ultimate origin in Eurasia. The Rhinocerotidae were present in northern Eur- oldest Dorcatherium is known from Africa from asía during the Oligocene. During the early Mio- Set O (PICKFORD1986a), possibly from the Bugti cene they first moved into Africa and the Indian fauna (UTragulus sivalensis"; RAZA & MEYER 1984) Subcontinent. The initial dispersal into Africa about and from MN 41 in Europe (FAHLBUSCH1985). 21 Ma ago involved tour genera (HOOIJER1978). Dorcabune,with a previous record in the Indian Subcontinent, is probably present in MN 10 in Suoidea: Sanítheríum(= Díamantohyus;Palaeo- Crete (VAN DER MADE 1997a). choeridae)first appearedin Set I in Africa and in the Bugti fauna of the Indian Subcontinent(PICK- Primitive pecorans: Material from African Set 1 FORD 1984, VAN DER MADE & HUSSAtN 1992) and and 11localities was assigned to cf. Strogulo- may have evolved from Palaeochoerusor a sim- gnathus (= ), Palaeomeryx, Propalaeo- ilar form (VAN DERMADE 1998), which is last ryx, Hindeterminate BovidaeH, Gelocus and Nam;- known from MN 2 in Europe. In MN 51, a later b;omeryx by STROMER(1926), WI1WORTH(1958) stage of evolution of Sanitherium enteredCentral and HAMILTON(1973) and MORALESet al. (1995) Europe,and was present in Chios. and HProdremother;um(?)" and "Gelocus(?)" by The origin of the is in Asia. Nguruwe PILGRIM(1912) from Bugti probably represent (Listriodontinae)first appeared in Africa in Set l. primitive pecorans. The bovoid H;spanomeryx is Bunolistriodon appeared in Africa in Set 1, the first known from Anatolia from MN 6e (MovA- Bugti fauna and later in MN 41 in Europe and SoLA 1987) and appeared in western Europe in China.Listríodon evolved in the Indian Subconti- MN61. nent from Bunolistríodonand dispersed in MN 6

463 : Citations of very early Bovidae and the after the dispersal of the first into Anatolia and the great diversity in the primitive forms in Central second into Africa. Asia, Mongolia and China (VISLOBOKOVA 1997) and There are early citations of the antilopine Ga- the sudden appearance of bovids belonging to zella (= ? Homoiodorcas) from Africa (Set lila, IV), different subfamilies in Europe, the Indian Sub- the Middle Chínji Formation in Pakistan and pos- continent and Africa, suggest that bovids have sibly from MN 6, but certainly MN 9 in Anatolia their origin in north or . (HAMILTON1973, 1978, THOMAS 1984, KÓH- The oldest bovids outside north or central LER1987). Hispanodorcas and the hippotragines Asia are the 18 Ma old boselaphine Eotragus from Palaeoryx and Protoryx had previous records in the Indian Subcontinent (SOLOUNIASet al. 1995) Greece and Anatolia (GENTRv& HEIZMANN1996). and the Eotragus from MN 4e in Europe, both These tour genera appeared in MN 12 in Spain probably coming from north or central Asia. Forms (NIETOet al. 1997). that have been attributed to Protragocerus first From the Vallesian onwards, till nearly the end appeared in Europe in MN 51, in the Indian Sub- of the Miocene, the African fossil record is poor, continent in the Lower Chinji Formation and in while there is an extremely rich bovid fauna in the Africa in Set IV:Miotragocerus (= ? Tragoceridus) Turolian of Greece and Turkey (SOLOUNIAS1981, of Eurasian origin entered some 6 Ma ago in KÓHLER1987, collections in Ankara). But few rela- Africa (GENTRY1978). Sivoreas entered the Indian tionships have been established between these Subcontinent 13.8 Ma ago (BARRY & FLYNN 1990) bovids, so that biogeographical interpretations and is found in the Vallesian of Africa (THOMAS are hazardous. 1981). A number of early citations of Eotragus are : The oldest giraffids known are Te- based on horn cores, that more probably belong rue/ia from MN 2 in Spain (MoYA-SoLA 1987), to primitive hypsodontinae. This mar be the case Progiraffa from Bugti in the Indian Subcontinent with a specimen from Gebel Zelten (HAMILTON (PILGRIM1912) and Zaraffa and Pa/aeotragus from 1973). Some very hypsodont teeth from Midra- Set 111in Africa (CHURCHER1978), whereas the ash-Shamali (WHYBROWet al. 1982), of about the giraffoid C/imacoceras is known from Set II (PICK- same age, lend support to this idea. The horn FORD1981, 1986a), which probably arrived inde- core from the Kamlial formation PILGRIM(1939) pendently from Asia. assigned to Eotragus, might siso represent a Giraffokeryx is cited from the Indian Subcon- hypsodontine. At present it is not clear, whether tinent from 16.5 Ma onwards and may have dis- the appearance of Kubanotragus in MN 5 in the persed to SE Europe and Anatolia in MN 5 and and 13.8 Ma ago in Pakistan and Tur- Africa in Set IV (BARRY & FLYNN 1990, GENTRY& coceros in MN 61 in Anatolia (KÓHLER 1987, BARRY HEIZMANN 1996, CHURCHER1978). This gives the & FLYNN 1990) present different dispersal events impression of a dispersal from the India n Subcon- or local evolution from such primitive forms. Hyp- tinent towards Africa and SE Europe late in MN 5. sodontus is known from early MN 5 or MN 6e However, doubts have been expressed on deter- equivalent localities in Anatolia, Pakistan, south- minations based on dental material (MONTOYA & ern Russia and Africa (KOHLER1987, GENTRY1978, MoRALES1991). Bramatherium is cited from the THOMAS 1984). Less hypsodont specimens from Indian Subcontinent from Aragonian times and China suggest a dispersal of Hypsodontus from appeared in the Turolian in Anatolia (MONTOYA& China or central Asia towards the other areas MORALES 1991). There are a number of other some 14 Ma ago. Miocene giraffids, but their relationships and are- The short premolars suggest, that a small as of origin are poorely known. brachyodont bovid from Ad Dabtiya that GENTRY (1987) assigned to Eotragus, might well belong to Cervidae: Cervidae first appeared in Europe in the hippotragine Tethytragus (= Caprotragoides). MN 3, but did not reach Africa and the Indian In MN 6e the genus was present in Anatolia and Subcontinent during the Miocene. nearly immediately two lineages can be recog- nized that dispersed to Europe in MN 61 (VAN DER Lagomorpha: Lagomorpha haya their origin in MADE 1994). Protoryx evolved in Anatolia from Asia. This is also the case for the Ochotonidae, Tethytragus and appeared in MN 8 in Africa (KÓH- which later spread to Europe and stilllater, during LER 1987). Gentrytragus is known from Set IV in the Early Miocene, enterad Africa. Austro/ago- Africa and Arabia (GENTRY1978, THOMAS1983). mys is cited from Set I (STROMER 1926) and Ken- Tethytragus and Gentrytragus (both were former- ya/agomys from Set 11(PICKFORD 1986a). The Lepor- Iy placed in Caprotragoides) possibly diverged idae did not enter Africa during the Miocene.

464 Rodentia: Rodentia are very well studied in Eu- The dispersal events rape. The collections from the Indian Subconti- nent and Africa are much smaller. Especially for During the Oligocene and earliest Miocene Africa, the available information might be rather (MN 1-2; Set O; "pre-Bugti") Africa and the Indian incomplete, and therefore it seems better not to Subcontinent had endemic faunas. Chapattimyid treat individual genera (apart from the fact that rodents were present in the Eocene of the Indian this would require much more space). Subcontinent and survived there ti" the earliest THOMAS(1985) listed as families that dispersed Miocene (JACOBSet al. 1981, KUMARet al. 1996). into Africa and Arabia during his NDP1 event: The primitive ruminant Leptomeryx and an an- Zapodidae, Sciuridae, Dipodidae, Cricetodontidae thracothere have been cited from the Oligocene and Ctenodactylidae (Sayimys) and during the of India (KUMARet al. 1996). The African faunas of NDP2 event the Gliridae. Cricetodontidae and that time include primates, creodonts, tubuliden- Sciuridae seem to have their origin in Eurasia tates, proboscideans and hyracoids. Tragulids (LAvoCAT1978, THENIUS1979) and are first re- were present in the earliest Miocene of Africa and corded from Africa from Set I (PICKFORD1986a). may have been present in the Indian Subconti- Dipodidae may have their origin in Europe and nent. Eurasian faunas had Carnivora, rhinos, pal- entered only in the north of Africa (LAVOCAT1978). aeochoerids, suids, primitive pecorans, giraffoids, Ctenodactylidae, Sciuridae and Cricetodontidae, etc. The subcontinent of SE Europe and Anatolia all supposedly of Asian origins, dispersed into the had been connected with Europe during the Oli- Indian Subcontinent during the Early Miocene (DE gocene; the Early Miocene faunas are not well BRUIJN & HUSSAIN 1984). Data in the time range 21- known, but probably showed many resemblanc- 16.5 Ma are poor, but the available data suggest es to European faunas. a majar dispersal event around 21 Ma from Asia In the next sections a series of faunal ex- to Africa and the Indian Subcontinent. DE BRUUN change events are treated. Table 45.1 compiles & HUSSAIN (1984) supposed the Gliridae to have the primary data. migrated from Europe to the Indian Subcontinent in the Middle Miocene, where they are known S,.chyodus event, 21 Ma: The anthracothere from Chinji equivalent levels. The glirid dispersals Brachyodus, of African origin, enterad Europe, to the Indian Subcontinent and Africa may form Asia and the Indian Subcontinent. part of a single evento Dispersals into Africa involved a larga number THOMAS (1985) listed the Pedetidae and Thryo- of chiropterans, erinaceids, the rodent familias nomyidae as dispersing from Africa to Eurasia Cricetodontidae and Sciuridae, the lagomorph during the NDP2 evento Pedetidae are first known Austrolagomys, the amphicyonid Cynelos, the from Set I localities in Africa (STROMER 1926, viverrids Kichechia, Legetetia, Leptoplesictis and PICKFORD1986). DE BRUIJN& HUSSAlN (1984) listed Mioprionodon, the stenoplesictid Stenoplesictis, Pedetidae? from a locality from a Kamlial equiva- the felid Afrosmilus, the rhinos Brachypotherium, Aceratherium, HDicerorhinus" and Chilotheridium, lent level. The seems to have dispersed towards Eurasia during MN 5/6 (THOMAS 1985) or the chalicothere Chalicotherium, the palaeoch- during the late Miocene (THENIUS1979). DE BRUIJN oerid Sanitherium, the suids Bunolistriodon and & HUSSAJN (1984) indicated that Rhizomyidae dis- Nguruwe and primitiva pecorans, all of Eurasian persed from the Indian Subcontinent to Africa origins. Possibly at this time the rodent families and Myocricetodontidae, of the same geograph- Dipodidae, Zapodidae and Ctenodactylidae en- ical origin, to Africa and Anatolia. The localities tered Africa. they cite indicate that the arrival in Africa is not Dispersals into the Indian Subcontinent in- later than MN 41 or Set lila, whereas the arrival in ciudad probably the creodont Hyainailouros, the Anatolia is not later than MN 6. proboscideans Hemimastodon, Gomphotherium, The oldest Muridae known are from Pakistan Choerolophodon and Deinotherium, the hyrax from a locality (DE BRUUN & HUSSAIN 1984) that is aft. Geniohyus and the anthracothere Brachyo- slightly younger than 16.5 Ma. This genus gave dus, sIl of African origino Dorcatherium may have rise to Progonomys and was replaced by that been present already or may have come from genus about 11.8 Ma ago (JACOBS et al. 1990). Africa. The Amphicyon, the rhinos Pa- This genus appeared in MN 9 in Anatolia (SEN raceratherium (= ?Baluchitherium), Brachypo- 1990) and in latest MN 9 in Europe (VAN DAM therium, Aceratherium, Chilotherium, the chali- 1997). Progonomys entered Africa in MN 10 (LA- cotheres Chalicotherium and Phyllotillon, primi- VOCAT 1978). tive pecorans, the giraffid Progiraffa and the rodents Sciuridae, Cricetodontidae and Cteno-

465 dactylidae,sIl of Eurasianorigin entered the Indi- This eventis within MN4, and at the begin- an Subcontinent. ning of zone C of the Aragonian and Set 111.It This event took place at the beginning of seemsto have been allowedfor by the regression MN 3, zona Z of tha Ramblian,Set 1,HBugtiH. In of the T82.3 cycle. numerous places, a regrassion is documented (Aquitaine,Untere Süsswasser Molasse in central Hominid event, 15.5 Ma: A hominid (Gríphopí- Europaetc.), which is the regressionat the onset thecus?), of African origin, entered Europe and of the TB2.1 cycle of HAQ et al. (1987). This possibly the first hypsodontine bovids, of Asian regressionvery likaly allowed the massivefaunal origin, enteradthe Indian subcontinent.However, exchange described above. these bovids may have been presentearlier, since they enterad during the previous event in Africa. Creodont event, 17.51 Me: The creodont Hysin- This event is possibly at the beginning of MN 5, si/auras, of African origin and the rodent Demo- zona D, Set IIIb and the TB2.4 cycle. cricetodon, of Anatolian origin dispersed into Eu- rape. The entry of the proboscideans Gompho- Conohyus/Pliopithecus event, 14 Me: The ca- therium and Zyg%phodon, of African origin, in tharrines Pliopithecus and Plesiopliopithecus,of Europe might antedate the entry of Democriceto- African origins, the rodent Cricetodon,of Anato- don, and thus be in MN 3. However, there are but lian origin, the palaeochoeridSanitherium, with a very few MN 3 localities with proboscideans. Pos. previous record in Africa and the Indian Subcon- sibly, the localities where these proboscideans tinent, and the suid Conohyus of Indian origin are found are very clase to or identical in age to entered Europe. the earliest localities with Democricetodon. Not later than at this moment, the tubuliden- Soricids, the lutrine Luoga/e, the herpestid tate Orycteropusand the hyrax Pliohyrax,of Afri- Herpestides, the ursid Hemicyon, the suid Kenya- can origin, the suid Kubanochoerusof Afro-lndian sus and the giraffoid C/imacoceras are first re- origin and the carnivore Percrocuta and cervids corded from Set 11.The Set 11localities are much dispersed into SE Europe and Anatolia. There is richer than the Set 1localities, it is possible that a however, virtually no previous large mammal number of these taxa were present already be- record in the SE Europeanand Anatolian afeas. toreo The hyaenid Protictitherium and the bovids Pro- At present, it is not sufficiently clear whether tragocerus,Gentrytragus and Hypsodontus,sIl of there was a single event in Europe, that occurred Asian origins, entered Africa. at the beginning of MN 4 and zone B, nor is it clear The aardvark Orycteropus, of African origin, whether it is synchronous to the African event at and the bovids Protragocerus, Sivoreas, Sivaceros the beginning of Set 11.tf there was an event, it is and Kubanotragus, of Asian origin dispersed into related to either the regression of the TB2.2 cycle the Indian Subcontinent. of HAQ et al. (1987), or to the changes in climate This event is within MN 5 and is at the begin- that caused this minar regression. ning of zone E, Set IV and the TB2.5cycle. At this time the Chinji faunal complex was formed. Bunolistriodon event, 16.5 Ma: The proboscide- an Deinotherium, the chalicothere, Chalicothe- Ustriodon event: The dispersal of Listriodon, of rium, the suid Bunolistriodon and the tragulid Indianorigin, to Anatolia and Eurasiaseems to be Dorc8theriumwith previous records in Africa and an isolated event at the beginning of MN 6 and the Indian Subcontinentand the cricetids Mega- zoneF. cricetodon,Eumyarion and Fahlbuschiaof Anato- lian origins, dispersedto Europe. Tethytragus event, 12.5 Me: The primitive bovoid The amphicyonids Amphicyon and Ysengri- Hispanomeryx and the bovid Tethytragus sIl with nia, felid Africanictis, the giraffids Zaraffa and previous records in Anatolia, the suid Albanohyus Giraffokeryxor Palaeotragusand the bovid sub- and probably the hominid , of African family Hypsodontinae, the first Gazel/a, sIl of origin entered Europe. The bovid Turcoceros ap- Eurasianorigins, and rhizomyid and myocriceto- pea red in Europe and in SE Europe and Anatolia; dontid rodents, of Indian origins, dispersed to its origin is either local in the latter afea or in Asia. Africa. The carnivore Percrocut8 entered Africa. The The catarrhine Dionysopithecusand pedetid bovid Helicoport8x, of Asian origin, and the hamo rodents,of African origin, and the giraffid Giraffo- inid Sivapithecus, of African origin, entered the keryx, of Asian origin, dispersed into the Indian Indian Subcontinent. Subcontinent. This event is within MN 6, probably at the

466 beginningof zonaG, at the beginningof Set V and beginning of MN 10, and late in zone l. It seems the middle Chinji faunal complex and onset of the likely that climatical changes triggered the events. TB2.6 cycle. Gazella event, 6.9 Me: The bovids Gaze//a, Prot- Propotsmochoerus event: Propotamochoerus, of oryx, Palaeoryx and Hispanodorcas first appeared Asian or Indian origin, entered Europe. KOHLER in Western Europe in the later part of MN 12 (1987) assumed a dispersal of Protoryx to Africa (NIETOet al. 1997) in localities with an estimated in MN8. age of 6.8-6.9 Ma (VAN DAM 1997). The MN 12-13 These dispersals did not coincide with a re- boundary is estimated at 6.8 Ma. when certain gression. Two cycles (T82.5-TB2.6) had cumula- murids first appear. These murids are still absent tive effect and global sea level remained very in the Gaze//a localities. The bovids have a previ- low; probably the sea did not impede any faunal ous record in SE Europe and Anatolia. This gives exchange between the majar land masses in the the impression, that the appearance of these Cid . There seem, however, to have been bovids in Spain is related to the events that changes in climate in the circum Mediterranean marked the MN 12-13 transition. Brea. In Turkey, the vegetation seems to have In Africa, a number of new bovids, including changed. The Eskihisar Pollenbild was replaced Cephalophus and Ugandax have their oldest by the Yeni Eskihisar Pollenbild in MN 8 (BENDA& records at about 7 Ma (GENTRY1978). There are MEULENKAMP1990). There were also changes in however incertainties about the origin or first the vegetation of southern Germany, MN 8 was occurrence of these bovids. interpreted as a wetter period (JUNG& MAYR1980). CERLlNGet al. (1997) reported on a global After a long period of absence, castorids entered vegetation change between 6 and 8 Ma ago; again in the Aragonian type afea (DAAMS& FREU- vegetation with a C3 photosynthetic pathway was DENTHAL1988), suggesting a change towards great- replaced by C. vegetation, which includes grass- er humidity. Such environmental changes may es. This was assumed to have occurred due to a have permitted the dispersal of these taxa. decrease of the CO2 concentration in the atmos- phere. The change in vegetation seems to have HíppBrion evem, 11 Me: The Equid Hipparion, of occurred first at the lower latitudes, whereas American origin, ente red the and with- Western Europe was not much affected. The in a short period reached the most distant parts. changes in vegetation caused important changes Hippopotamodon (= Microstonyx) probably of in the fauna in Africa. Pakistan and North and Asian origin entered in Europe and the Indian . Though Europe was much less subcontinent. Mellivora, of Asian/lndian origin, affected, the appearance of tour hypsodont bovids and Conohyus of European, Anatolian or Indian in Spain is worth to be noted. origin, dispersed into Africa. The TB3.1 cycle starts with a drop of ayer 100m sea level. Within Hippo event, 6.3 Me: A hippo (Hexaprotodon/ the Old World were connected al- Hippopotamus),the bovid Redunca?and the cer- ready, but this event seems to have allowed for a copithecids Macaca and Mesopithecus, of Afri- faunal exchange with America. can origins, the bovid Parabos,possibly of Indian This event is at the beginning of MN 9, zone H, origin, and the ursid Agriotherium, possibly of Set VII and the TB3. 1 cycle. It marks the beginning Asian origin, entered Europe. of the Nagri Faunal complex. The ursid Agriotherium, the mustelidsPlesio- gula and Enhydriodon, the felid Felis, and the Murid event, 9.6 Me: The murid Progonomys, of viverrid Vive"a, with origins in Eurasiaand/or the ultimate Indian origin, and the hyrax Pliohyrsx Indian Subcontinent, the chalicothere Ancylo- dispersed into Europe. Possibly the dispersal of therium, with an origin in the Middle East, the the palaeochoerid Schizochoerus was at the same bovids Leptobos?,of Asian or Indian origin, and time. AlI had previous records in Anatolia. There Miotragoceros,with a previous record in Europe are slightly older citations of Progonomys in Eu- and SE Europeand Anatolia,and (for the second rope, but the massive presence of murids that time) Conohyus,of Indian origin, entered Africa. characterises this event is very late in MN 9 or The timing of these dispersalsis not very sature, MN 10 (VAN DAM 1997). Progonomys and possi- sincethe previousAfrican fossil record is not very bly Sivschoerus spread at this same moment to abundant. Africa. Dorcabune may have extended its range Hexaprotodon.of African origin and different from the Indian Subcontinent into Crete. from the Europeanhippo, enteradthe IndianSub- This event is very late in MN 9 or at the continent.

467 This event occurred within MN 13, at the be- other factors, than the arid belt, limiting faunal ginning of zone M2 and within the Dhok Pathan exchange between the southern and northern unit. There were two regressions during the TB3.3 continents. This indeed seems to be the case. and 3.4 cycles, causing the events that are known Though are not subject of this paper, as the Messinian Salinity Crisis. The first regres- it should be mentioned that folivorous animals sion probably allowed for most of the dispersals. like tapirs and possiblyAnchitherium and animals The dispersals towards the Indian Subcontinent that may have been (partially)frugorivorous, like are more likely to be related to climatical changes. tragulids and certain primates, became rare or went extinct in Europeduring the later part of the Vallesian,when the Europeanclimate may have Conclusions become more seasonal. The Miocene faunal exchange between Eu- The Miocene faunal exchange between Eurasia, rape (or Eurasia),the block formed by SE Europe Africa and the Indian Subcontinent was intense. and Anatolia, Africa and the Indian Subcontinent Today, these areas are known as the Palaearctic, occurred in a seriesof distinct events (table45.1). Ethiopian and Indian bioprovinces. The latter two The magnitudevaries greatly from none (?)to over together are siso known as Palaeotropical bio- 20 taxa dispersinginto a particular(sub)continent province. The recognition of these bioprovinces (note that "taxa" as presentedhere, includesgen- goes back to SCLATERand WALLACEin the nine- era, but siso whole families).During the Earlyand teenth century. The Palaearctic is the Brea north of Middle Miocene,these dispersal events seem to the arid -Gobi belt and tends to be season- have been allowed for by low sea level stands, al in that it has winters and hot summers, the which in turn are causedby a cold global climate, degree depending partially on latitude, and it having probably an influence on faunal distribu- includes Breas with very low temperatures. The tion in its own right. When the Miocene proceed- Palaeotropic is the ares south of the arid belt; it ed, the steplike lowering of the sea level (MILLER tends to have more of a wet-dry seasonality and et al. 1995)caused land massesto become con- includes permanently warm areas. The Ethiopian nected,later changesin sea levellost importante and Indian bioprovinces are physically separated in biogeography and climate changes became by the , but have similar climates. the driving force controlling patterns of faunal These climates are reflected in the vegetation. distribution. This is seen in the timing of the Seasonal variations in the supply of fruits and dispersal events (see murid and Gazellaevents, leaves tend to be greater in the Palaearctic, limit- table 45.1).The differencewith the early Miocene ing the distribution of folivorous and frugivorous is that changes towards a warm global climate animals. (with high sea level) siso caused intercontinental During much of the Miocene a similar biogeo- dispersals (as may have been the case in the graphical pattern seems to have prevailed. As we murids), resulting in more frequent and more have seen above, mammals that may have been complex changes in biogeography. dependent on so me degree of humidity, like ta- The UBrschyodusevent" (about 21 Ma ago, pirs, the equid Anchitherium, cervids, moles etc. MN 3, Set 1, Bugti) is characterizedby massive were present in northern Eurasia, during a long dispersals into the Palaeotropicand only very part of the Miocene, but did not enter Africa and limited dispersals into the Palaearctic.It is only India; there may have been an ares south of during the "Bunolistriodon event" (MN 41, Set Turkey, that was toa arid for them to cross. While lila), UConohyus/Pliopithecusevent" (MN 51, Set there seems to have been an important filter IV, Chinji) and "Tethytragus event" when there controlling faunal exchange between Eurasia and were important dispersals into the Palaeoarctic. the southern continents, the data presented above The "Tethytragusevent" involved siso important suggest, that faunal relationships between Africa dispersalsinto the Palaeotropic.The Upper Mio- and the India n Subcontinent were more intense. cene record in Africa is rather incomplete, and There are, however, differences between the re- does not allow for exact timing of the events. It cent and Miocene patterns, in that the present does suggest however, again majar dispersals Holarctic extends into , whereas this into Africa. The latest Miocene dispersal of hip- was not the case in the Early and Middle Miocene. pos of African origin into Europe has received PICKFORD& MORALES(1994) claimed thatthe bound- much attention, but is limited to the south of ary zone between the Ethiopian and Palaearctic Europe. Correctingfor the emblematic dispersal bioprovinces shifted north several times during of the hippo to , the resulting the Miocene, which would sugg8St that there are picture is of a change of the balanceof immigra-

468 tion: first the majority of dispersals were into the Palaeotropic, later into the Palaeoarctic and then ALFéREZ, F., MOLERO,G., MALDONADO,E. & IÑIGO,C . again into the Palaeotropic. Though there exist (1988): Los restos fosiles mas antiguos de Orycter- differences with the results of PICKFORD& Mo- opidae (Tubulidentata, Mammalia) hallados en Eur- RALES(1994) in the details of timing of the events asia. - Coloquio Homenaje a Rafael Adrover " Bio- and the dominant direction of dispersal at a par- eventos y Sucesiones faunísticas en el Terciario ticular time, both studies support the view that Continental Ibérico". Sabadell, 14-16/12/1988. Re- the balance of faunal exchange changed through sumenes: 7; Sabadell. the Miocene. ANDREWS, ~, HARRISON, T., DELSON, E., BERNOR, R. L. & Global sea level has been relatad to global MARTlN, L. (1996): Distribution and biochronology of climate and the accumulation of ice masses on European and southwest Asian Miocene catarrhines. - In: BERNOR,R. L., FAHLaUSCH,V. & MITTMANN, H.-W. continents, in particular on , but also to (eds.): The Evolution of Westem Eurasian Neogene tectonic activity (PARTRIDGEet al. 1995). Whereas Mammals Faunas, 168-208; New York, Chichester global sea level reacts immediately to climate, the (Columbia University). relationship may change gradually over longer BARRY, J. C. & FLYNN, L. J. (1990): Key biostratigraphic periods. Tropical soils in Germany, with ages events in the Siwalik sequence. -In: LINDSAY, E. H., ranging 17-15 Ma, have been interpreted to be FAHLBUSCH,v: & MEIN, ~ (eds.): European Neogene Mammal Chronology. - NATO ASI Saries, (A) 180: caused by the greenhouse effect, causad by high 557-571; New York, London (Plenum). atmospheric CO2concentrations (SCHWARZ1997). BENDA, L. & MEULENKAMP,J. (1990): Biostratigraphic cor- Oxigen isotope records suggest relatively warm relations in the Eastem Mediterranean Neogene. 9. global climates during the Early and initial Middle Sporomorph assocíations and event stratigraphy of Miocene, however, becoming progressively colder the Neogene. - Newsletters during the Middle Miocene (MILLERet al. 1995). on Stratigraphy, 23 (1): 1-10; Berlin, Stuttgart. This occurred in the period MN 6-9 (14-10 Ma), BoNIS, L. DE, BOUVRAIN, G., GERAADS, D. & KOUFOS, G. when the balance was shifting in favor of disper- (1990): New remains of Mesopithecus (Primates, sals towards Africa. There is evidence for a glaci- Cercopithecoidea) from the Late Miocene of Mace. ation in 11 Ma ago (HELLAND& HOLMES donia (Greece), with the description of a new spe- cies. - Journal of Vertebrate , 10 (4): 1998). 473-483; Chicago. As mentioned above, seasonality is an impor- BRUUN, H. DE; DAAMS, R., DAXNER-HóCK, G., FAHLBUSCH, tant determinant in climate. large areas of Europe V:, GINSBURG,L., MEIN, ~, MORALES, J., HEIZMANN,E., may have had a sufficiently warm and humid and MAYHEW, D. F:, MEULEN, A. J. VAN DER, SCHMIDT- not very seasonal climate during part of the Mio- KlTTLER, N. & TELLES ANTUNES, M. (1992): Report of cene, to allow the right vegetation to grow for the RCMNS working group on fossil mammals, Rei- folivorous and frugivorous animals. Another limi- sensburg 1990. - Newsletterson Stratigraphy, 26 tation related to seasonality is the reproductive (2/3): 65-118; Berlin, Stuttgart. BRUUN, H. DE & HUSSAlN, S. T. (1984): The succession of cycle that in the Palaearctic coincides with the rodent faunas from the Lower Manchar Formation seasonal cycle (PICKFORD& MORALES1994). 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