NEOGENE BIOGEOGRAPHY OF HYRACOIDEA (MAMMALIA) by T. K otsakis Members of the Order Hyracoidea (family Pliohyracidae) are very common in Palaeogene deposits of Africa but unknown from layers of other continents. But in the Neogene both Europe and Asia received immigrants. During Palaeogene time the subfamily Sagatheriinae is the dominant one. During Rusingan (African equivalent of Agenian and Orleanian) members of the genus Pachyhyrax, belonging to this subfamily, are found in East Africa (at Bukwa/Uganda, Rusinga/Kenya etc) (M eyer, 1978). Members of the same genus were discovered at Ad Dabtiyah and at Al-Sarrar, both belonging to the Dam Formation, in East Saudi Arabia, near the shores of Persian Gulf (H amilton, W hybrow and M cClure, 1978; T homas, 1982). Beside the fact that many rodents of these sites are of Asian origin (Sen, 1982a) the whole fauna indicates African connections. This is not surprising as free migration was apparently possible between Arabia and both North and East Africa during this time. Perhaps a Sagatheriine is present also in Gebei Zelten (Libya) (Savage and H amilton, 1973). But the subfamily who undertook a large eXpansion during Neogene was, by far, the subfamily Pliohyracinae. Following M eyer (1978) the most ancient findings of this subfamily come from the Rusingan of East Africa, both Lower Rusingan (a, Bukwa) and Late Rusingan (at Rusinga). This material is referred to the genus Meröehyrax. A more recent finding in Africa is that of Beni Mellal (Morocco—Middle Astaracian). The specimens discovered in this layer are classified as Parapliohyrax (G insburg, 1977). Out of Africa the most ancient findings of Pliohyracine hyracoids, belonging to the genus Pliohyrax, were these of the Aegean area, Lower Vallesian in age. They are collected at Melambes in the Island of Crete (Greece) and in Turkey at Esme-AkQakoy (Kuss, 1976; Becker Platen, Sickenberg and T obien, 1975a, 1975b). A migration of hyracoids from Africa to Aegean area at the moment (or moments) of the arrival (or arrivals) of Progonomys and Hipparion in Europe and N. Africa is possible but the rodents of Aegeid testify eXtinctions rather than migrations of African forms at the time of the boundary Astaracian—Vallesian (Sen, 1982b). Thus, a previous migration from Africa, cannot be excluded. In fact, by some palaeontologists (Leinders and M eulenkamp, 1978), the layer of Melambes is considered of the same age with the Cretan layer of Plakias, of Middle or Upper Astaracian age. The genus Pliohyrax is unknown from the very few layers of Upper Vallesian age of the Aegeid but is well represented or rather, less rare, in the Lower and Middle Turolian deposits of Greece and Turkey. At Balgikli Dere and perhaps at Kayadibi (Lower Turolian) and at Gar- kin, Pikermi, Halmyropotamos, Samos (Middle Turolian), remains of one or two species of the genus Pliohyrax were collected (M elentis, 1966; Becker Platen, Sickenberg and T obien, 1975a, 1975b; Solounias, 1981a, 1981b). 478 T. Kotsakis Fig. 1. Geographical distribution of the Hyracoidea during Neogene and Quaternary (excluding Sub-Saharan Africa) Lower and Middle Miocene: 1 Gebei Zelten; 2 Ad Dabtiyah; 3 Al-Sarrar; 4 Beni Mellal. Upper Miocene: 5 Melambes (?); 6 Esme Ak^akoy; 7 Can Llobateres; 8 Soblay; 9 Kayadibi; 10 Bal?ikli Dere; 11 Garkin; 12 Pikermi; 13 Halmyropotamos; 13 Samos; 15 Taghar; 16 Molayan; 17 Sor; 18 Baode. Pliocene: 19 Montpellier; 20 Jalalabad (?); 21 Yushe basin, zone II; 22 Cap Travertin; 23 Kvabebi. Pleistocene: 24 Nihewan; 25 Iraq e Zigan In the other side of Europe, at Can Llobateres in Spain, a well-known site of Lower Vallesian age, few fossils of a hyracoid assigned with doubt to are reported (G olpe Posse and C rusafont Pairo, 1981). Remains of a local species of Pliohyrax are discovered also in southern France, at Soblay, a layer of Upper Valle­ sian age (Viret, 1949). The Upper Astaracian—Lower Vallesian marco- and micro­ mammals discovered in the Iberian Peninsula are very different from those of Maghreb and we must conclude that a Betico-Rifain way of dispersion is highly improbable (Bruijn and M eulen, 1981; K otsakis, 1986). Thus, a migration of hyracoids from Southeast Europe is the most probable. Anyway, the scanty record of hyraxes in West Europe and their absence from the deposits of Central Europe indicate that these animals were very rare. Two Pliocene records of big hyracoids, perhaps representing the final stages of this Upper Miocene eXpansion, are also known. In the Upper Ruscinian or Lower Villafranchian (sensu A zzaroli, 1977=Villányán) layer of Kvabebi, in the Georgian SSR, a great number of hyracoid remains were collected. The specimens are attrib­ uted to an endemic Pliohyracine, Kvabebihyrax (G abunia and Vekua, 1966, 1974). Very probably this form is the last survivor of the eXpansion of Pliohyracines in the Near and Middle East. Also in France, a Pliohyracine of big size was found in the Lower Ruscinian of Montpellier (Viret and T henius, 1952). The relationships of this Neogene biogeography o f Hyracoidea.. 479 species, attributed either to Pliohyrax or to Postschizotherium, with the Vallesian ones from the same area or with oriental elements of the Order Hyracoidea is a matter of question. During Upper Miocene Pliohyracines eXpanded their area also to Central and Eastern Asia. Remains of members of the genus Pliohyrax are collected in the Turolian deposits of Afghanistan and more precisely in the Lower Turolian site of Taghar and in the Middle Turolian site of Molayan (H eintz and Brunet, 1982; Brunet, H eintz and Battail, 1984). It is interesting to note that hyracoids never reached the Siwaliks in spite of the proXimity of the Afghan layers (Brunet, Heintz and Battail, 1984). The layer of Sor, in the SSR of Tadzhikistan, might be also of Turolian age. In this locality remains of a Pliohyracine, named Sogdohyrax by D ubrovo (1978), are discovered. During the last phases of Miocene, Pliohyracine hyracoids undertook an ex­ pansion further to east reaching China. The most ancient findings came probably from Baode (ShanXi) from sediments of Baodean age (Chinese equivalent of Turolian) and are assigned to the genus Pliohyrax (Tung Y oung-sheng and H uang Wan-po, 1974) but this attribution is criticized (D ubrovo, 1978). The other remains of Chinese hyraxes are classified within the genus Postschizotherium. Three species are described, but some fossils were bought in drugstores and therefore their geographic and stratigraphic position is uncertain (Koenigswald, 1966). Anyway remains belonging to the genus Postschizotherium are found in the second zone of Ylishe basin, of Jinglean ag$ (Chinese equivalent of Ruscinian) and in the Cap Travertin (Hebei-Henan), of Youhean age (Chinese equivalent of Lower Villafranchian or Villányán). The genus survives during the Lower Pleistocene at Nihewan and other localities (Li, Wu and Qiu, 1984). A radius discovered in the (probably) Upper Ruscinian site of Jalalabad in Afghanistan is assigned also to Postschizotherium (Heintz, Ginsburg and H arten- berger, 1978). At the moment of the discovery of this fossil the presence of Pliohyrax in the Turolian layers of the same country (Afghanistan) was unknown. At present four hypotheses are possible: 1) the hyracoid from Jalalabad represents the last Central Asiatic survivor of the eXpansion of Pliohyrax during the Turolian; it may be a relict population like Kvabebihyrax in Caucasus and perhaps the hyracoid of Montpellier; 2) it belongs to the genus Kvabebihyrax; 3) it belongs to the genus Sogdohyrax; 4) it is really a Postschizotherium coming back from China through Central Asia. Perhaps it may be also that the age of the layer of Jalalabad is older than Upper Ruscinian (interv. of D r. E. Heintz during the Congress). More material should be available for a solution of this problem. The history of the recent family Procaviidae begins with the genus Prohyrax in the Lower Miocene of Africa. This family includes also a few fossil species of the genera Gigantohyrax and Procavia from the Plio—Pleistocene deposits of the same continent (M eyer, 1978). A specimen referred to cfr. Prohyrax is reported from the Libyan layer of Gebei Zelten (M eyer, 1978). The recent species Procavia capensis Pallas, the only one of the Order living not only in Africa but also in Syria, Lebanon, Israel, Arabian Peninsula, is known from the Upper Pleistocene deposits of Near East (Heller, 1978). The mention of the genus Dendrohyrax as fossil in my Abstract (K otsakis, 1985) is due to a mere error of type­ writing. (For the Mammalian Stages—Ages of Europe, Africa and Asia see M ein, 1975; Savage and Russell, 1983; Ch. Li, W. W u, andZH. Qiu, 1984.) 480 T. Kotsakis REFERENCES A zzaroli A. 1977: The Villafranchian stage in Italy and the Plio-Pleistocene boundary. — Gior. Geol. (2). 41:61—79. Becker Platen J. D., Sickenberg O. and Tobien H. 1975a: Die Gliederung der känozoischen Sedimente der Türkei nach Vertebraten. Faunengruppen. — Geol. Jb. (B). 15.: 19—45. — 1975b: Vertebraten. Lokalfaunen der Türkei und ihre Altersstellung. — Geol. Jb. (B). 15. :47—100. Bruijn H. de and M eulen A. J. van der 1981: The distribution of land mammals in the Mediter­ ranean through the Neogene. Is there a fit with the paleogeographic and paleoclimatologic reconstructions based on data from marine basins? — Ann. géol. pays hellén. (Hors sér.) 4.:323—335. Brunet M., H eintz E. and Battail B. 1984: Molayan (Afghanistan) and the Khaur Siwaliks of Pakistan: an example of biogeographic isolation of Late Miocene mammalian faunas. — Geol. Mijn. 63. :31—38. D ubrovo I. A. 1978: New data on fossil Hyracoidea. — Paleont. Zhur. (3):97—106. Moskva (in Russian). G abunia L. K. and V ekua A. K. 1966: Peculiar representative of Hyrax in the Upper Pliocene in Eastern Georgia.