A systematic analysis of an hipparionine horse assemblage from Maragheh, Iran is made. A brief orientation to systematic philosophy and informal superspecific characterizations of some Old World hipparionines is given as a background to this work. A character state analysis of skulls is made, and has revealed five distinct species. A character state and stratigraphic trend analysis of isolated check tooth and postcranial remains, with known provenance, is also made. These two combined analyses reveal that the most resolute discrimination of hipparionine species and their evolutionary relationships occurs when multiple character complexes of associated skulls, maxillary and mandibular dentitions are made. When this is not possible, skulls have provided the best basis for discriminating species and their evolutionary relationships. Traditional characters of isolated cheek teeth and postcranial remains are shown here to offer limited information content for hipparionine phylogenetic systematics. The systematic portion of this study includes a comprehensive description of cranial and postcranial remains, and has further corroborated the distinction of five species which belong to at least three superspecific groups including: «Hipparion» geltyi sp. nov., Group 1; Hipparion prostylum (s. l.), and Hipparion campbelli sp. nov., Group 3; «Hipparíon» aff. moldavicum and «Hipparion» ?matthewi, Group 2. These species stratigraphic ranges and evolutionary relationships are also given here and argued to be important for establishing future hipparionine geochronologic correlations between a number of Eurasian late Miocene provinces. 

Two steps can be distinguished in the history of the first invasion of western and south western Europe by the arvicolines. The first step corresponds to the installation of these rodents with the immigration of Promimonys inxuliferus Kowalski, then of Mimomys stehlini Kormos and of Mimomys gracilis (Kretzoi). The second is characterized by the establishment of a geographic differentiation in the arvicoline fauna between the south of France and Spain, from where are described new species of Mimomys (Mimamys cappettai, Mimomys septimanus, Mimonys medasensis), and the rest of France, where are found only elements already known from central Europe or England (Mimomys polonicus Kowalski, Mimomys pliocaenicus F. Major, Mimomys reidi Hinton, or forms very close to the latter). This geographic differentiation, which is very certainly the consequence of the division of Europe into distinct climatic provinces, one of them being the southern province comprising at least Spain and southern France, could result from a cladogenetic evolution of Mimomys stehlini and Mimomys gracilis after their immigration. The present work is also a contribution to the search for correlations between the diverse micromammal localities of the latest Pliocene (or early Villafranchian) and of the early Quaternary of Europe. 

The mammalian faunas of the Paleogene of Europe and their localities are reviewed with comments on problems of European stratigraphy (epoch, stage and substage limits) and on the possibilities of faunal migrations. Radiometric dating is discussed. A stratigraphic scale for the Paleogene is presented, as well as a refined system of sequential faunal levels. 

Aside from Madtsoiidae, anilioids, and Boidae that were studied previously, the middle Palaeocene of ltaborai (BraziI) has produced Ungaliophiinae ("tropidophiids"), booid-grade snakes incertae sedis, and a possible Russellophiidae (Caenophidia) that are described in the present article. This article is the third and final report on the snakes from the locality. The Ungaliophiinae (Paraungaliophis pricei gen. et sp. nov.) are rare whereas the booid-grade snakes incertae sedis (ltaboraiophis depressus gen. et sp. nov., Paulacoutophis perplexus gen. et sp. nov.) are more frequent. A single vertebra is referred to the Russellophiidae (Caenophidia) with reservation. An update of the whole fauna of snakes from ltaborai is provided. Hechtophis austrinus that was tentatively referred to the erycine Boidae is now regarded as a Boidae incertae sedis. Most snakes from Itaborai are known only from the locality. Astonishingly, only the ailioids Coniophis cf. C. precedens gives possible evidence of interchanges between South and North America. The fauna of snakes from Itaborai, as well as the other Palaeocene faunas of snakes from South America are distinct from those of the Cretaceous and the Eocene of South America; they appear to be more different from the Cretaceous faunas than from those of the Eocene. The fauna from Itaborai is the richest and most diverse assemblage of snakes from the Palaeocene worldwide; it shares only a few taxa with other Palaeocene localities. 

Sieving and washing of about 700 kg of sediment from the miocene site at Montpeyroux produced otoliths of 34 teleost species, of which four still occur in the present day fauna or are near to extant species. Among the fossil species, eight are new : Ilisha lerichei, «genus Clupeidarum ›› orbiculatus, Dipulus mediterraneus, Morone cornuta, Chanda nelsoni, Pomadasys steurbauti, «genus Sciaenidarum ›› barthassadensis and Paraplagusia roseni. The fauna found is typical for a tropical or subtropical very littoral, probably even estuarine environment; it was living in a period near the boundary between Lower and Middle Miocene, perhaps somewhat earlier. 

Le présent volume est l'aboutissement d'un projet né il y a presque cinq ans. En décembre 1991, l'un d'entre nous (MG) prenait des contacts en vue de proposer un symposium sur les mammifères fossiles, dédié à D.E. Russell, dans le programme du 4e Congrès de la European Society for Evolutionary Biology. Ce congrès, baptisé "Evolution 93", devait se tenir à Montpellier en août 1993. Son Comité d'Organisation, animé par F. Catzeflis, recherchait des organisateurs de symposiums. L'idée fut acceptée avec enthousiasme par le second d'entre nous (PDG), et le titre de notre Symposium fut précisé: " Palaeobiology and Evolution of Early Cenozoic Mammals - A Symposium in Honor of D.E. Russell". Le projet fut formellement accepté par le Comité d'Organisation en avril 1992. 

New records of terrestrial mammals from the Qasr el Sagha Formation, Fayum Depression, Egypt are reported, and the stratigraphic occurrences of these fossils noted. These include additional specimens of Moeritheríum, Barytherium, and anthracotheres, as well as the oldest record of a hyracoid in the Fayum.These Eocene mammals occur almost exclusively in the alluvial deposits of the Dir Abu Lifa Member of the Qasr el Sagha Formation and show close affinities to the faunas from the lower sequence of the Jebel Qatrani Formation. There is no evidence of a more marked faunal discontinuity between the Qasr el Sagha and Jebel Qatrani Formations than there is across any of the three major breaks in sedimentation that exist within the Jebel Qatrani Formation. The faunal similarities between fossils of the lower sequence of the Jebel Qatrani Formation and of the upper part of the Qasr el Sagha Formation is consistent with recent paleomagnetic dating that suggests that these rocks differ in age by only one to two million years. 

This study describes four taxa of Gliridae from the Oligocene mammal locality Gröben 3: Gliravus tenuis BAI-ILO, 1975, Bransatoglis micio (MISONNE, 1957), B. planus (BAHLO, 1975) and B. heissigi n. sp. Gliravus tenuis from Gröben 3 is somewhat more advanced than the type population found in Heimersheim. This confirms previous research suggesting that Gröben 3 should be dated earlier than Heimersheim (MP 24). The first documented occurrence of B. mício around level MP 24 was found in Gröben 3. An abundance of tooth material from B. planus in Gröben 3 makes it possible, for the first time, to observe evolutionary stages within this species from MP 21 until MP 28. B. heissigi n. sp. is restricted to level MP 24. This species is located between B. mísonnei (MP 20 - 23) and Microdyromys praemurinus (MP 25 - 28). Within the lineage Bransatoglis bahloi - B. misonnei - B. heissigi, a decrease in size is noticeable.

  

The fauna from three new rodent localities (Castelmaurou, Grépiac-carrière et Grépiac-rive gauche) from Oligo-Miocene molasses of the Toulouse area is described. The one from Colomiers is completed. 11 species belonging to 4 families (Cricetidae, Eomyidae, Gliridae, Sciuridae) are present. The Miocene localities of Grépiac-carrière and Colomiers are correlated with Balizac, La Brète, Lambert and Lespignan. Grépiac-rive gauche is just a little older than these sites. Castelmaurou is somewhat younger than La Milloque and belongs to Oligocene. 

The exploitation of new early Eocene localities in the Paris Basin has resulted in the collecting of  numerous mammalian remains, among which are about 300 isolated teeth representing the rodents. They belong, for the most part, to the paramyid group. Only the latest level of the early Eocene has yielded rodents belonging to the pseudosciurid group. The paramyids, the object of this study, are represented by at least 5 genera and 10 species; they are distributed among 4 clearly dilferentiated subfamilies : Paramyinae Simpson 1945, Pseudoparamyinae Michaux 1964, Ailuraviínae n. subf., Microparamyinae Wood1962.
It results from this study that the principal types of rodents in the middle and late Eocene of Europe are clearly related to the forms described here: Plesiarctomys Bravard 1850 is related to Pseudoparamys Michaux 1964, Ailuravus Rütimeyer1891 to Meldimys n. gen., Gliravus hammeli Thaler 1966 to Microparamys nanus (Theilard1927), and some Masillamys Tobien 1954 to Microparamys russelli Michaux 1964 and to M. sp. 1. Gliravus and Masillamys are the oldest representatives of the Gliridae and the Pseudosciurídae, respectively, families which will progressively replace the Paramyidae in the course of the middle and late Eocene.
Two stages can be recognized in the history of te European Paramyidae fauna during the course of the early Eocene: the older is characterized by the Mutigny fauna, the younger by the Cuis fauna.
The Mutigny fauna appears already rather diversified and does not completely correspond, point by point, to the homologous fauna of North America. A geographic differentiation seems to have been manifested rapidly, unless the fauna established in Europe was already a little different from that which established itself in North America. 

The small insectivorous placental mammals from Ste-Néboule are poorly varied. Two orders of insectivores are documented : the Proteutheria by Pseudorhyncocyon cayluxi (leptictids), the Lipotyphla by Saturninia gracilis and S. beata (nyctitheriids). The bats are only rhinolophoids and mostly hipposiderids. Sufficient populations allow us to restrict the original definitions of three previous species (Hipposideros schlosseri, Palaeophyllophora quercyi, P. oltina). From now, these species can be regarded as monophyletíc. 

Three erinaceomorph species are known from the early Middle Eocene of Grube Messel near Darmstadt, Germany, which are referred to the family Amphilemuridae. Pholidocercus hassiacus, Macrocranion tupaiodon, and Macrocranion tenerum showed extraordinary adaptations to their different life strategies, and several of their specializations are unknown among living insectivores. Pholídocercus was a well-defended robust animal with an opportunistic feeding strategy. Macrocraníon zupaiodon was a slender forest floor-dweller with saltatorial specializations to escape from predators; fishes were the preferred component of its omnivorous diet. Macrocranion tenerum exhibited a combination of both survival strategies, extremely elongated hind limbs for rapid and even ricochetal flight and a spiny exterior as an effective protective device; it was probably specialized for feeding on ants. Thus, closely related, omnivorous-insectivorous forest floor-dwellers could exploit the Messel ecosystem. 

Some new material of percrocutas from the late Miocene of Axios valley (Macedonia, Greece) is studied. They have been found in the locality of "Pentalophos 1" (PNT). The material has been described and compared with the known late Miocene percrocutas of Eurasia. This comparison indicates that it can be identified as Dinocrocuta gigantea (SCHLOSSER, 1903). A maxilla of a percrocuta, named ”Hyaena" salonicae, was found in the same area (Andrews, 1918). "Hyaena" salonicae is smaller than the PNT material. It is also compared with other material from Eurasia while its taxonomic and age problems are discussed. It belongs to Dinocrocuta and shows close relationships with D. robusta and D. senyureki; its age can be considered as late Vallesian-early Turolian. The age of the locality PNT is also discussed and a possible Vallesian age is proposed for it. 

A new carcharhinid shark species, Physogaleus hemmooriensis sp. nov., is described from the Lower Hemmoorian (Behrendorfian, late Burdigalian, early Miocene) of Werder, Lower Saxony, Germany. P. hemmooriensis also occurs in the Edegem and Antwerpen Sands Members of the Berchem Formation, Belgium, and in the Miste Bed, Aalten Member of the Breda Formation, The Netherlands, which have an early to middle Miocene age. In the Western Atlantic region, the taxon is present in the early Miocene Calvert Formation of Delaware, U.S.A, which is largely contemporaneous with the Hemmoorian. 

Stratigraphic studies in the Aktau Mountains bordering the Dzhungarian Alatau Range in southeastern Kazakhstan have included mapping of Tertiary lithostratigraphic units, documentation of fossiliferous deposits, correlation of sections, etc. These investigations have led in turn to revised interpretation of the Tertiary geology of the area. The Tertiary sequence in the Aktau Mountains is represented by three lithostratigraphic units (in ascending order): (1) the middle Eocene Akbulak Formation; (2) the Oligocene Aktau Formation with a lower member including white quartz sands that contain fossil mammals, and an upper member including red-colored clays and sandstones, brick red clays, an anhydrite and gypsum clayey horizon, and bright brown-red clays; and (3) the upper Oligocene-Miocene Chul'adyr Formation with a lower member of greenish and yellowish conglomerates and gritstones, a middle member including grayish and yellowish sands and gritstones, and an upper member including brown and red clays and carbonate- and anhydrite-rich clays. The Aktau and Chul”adyr Formations represent separate cycles of sedimentation. Mammalian biostratigraphy and biochronology of the three vertebrate faunas in Aktau Mountains are reviewed. The mammalian fauna from white sands of the lower Aktau Formation is small but includes Ardynia and is thought to be early Oligocene in age. The mammalian fauna from conglomerates and gritstones of the lower member of the Chul”adyr Formation is also small but includes Paraceratherium and is thought to be late Oligocene in age. The mammalian fauna from sands of the middle member of the Chul'adyr Formation is extensive, with micro- and macrofauna attributed to Neogene mammal zones MN4 to MN 6, indicating a latest early Miocene to earliest middle Miocene age (Orleanian-Astaracian). Most genera of middle Chul”adyr mammals are known from the middle Miocene Shanwang faunas of China and from the Castelnau-d”Arbieu faunal assemblage (MN4-MN6) of southwestern France. 

The scientific career of D.E. Russell began with a Pliocene fauna from Oregon, and then turned in the direction of European Paleogene mammals. Field work followed by study of the mammals that were collected, firstly in the Paleocene and later in the early Eocene, greatly rejuvenated learning in this field. Syntheses on the Northwest European Tertiary basin and on European marnmals and stratigraphy came next. Research on the Eocene of Asia was carried out jointly with Gingerich on Pakistan and with Dashzeveg on the faunas of Mongolia. An important synthesis on the entire Paleogene of Asia, joint with Zhai, followed. Field work in Africa with Sigogneau-Russell led to the discovery of Mesozoic mammals there. A synthesis of mammalian paleofaunas of the world was written with Savage, and a similar synthesis of Cenozoic vertebrate faunas is currently being prepared. These achievements reflect the perennial importance of field work, numerous collaborations with both amateurs and professionals, and the human qualities of this author.
  

The recently discovered, very rich, Pliocene locality of Layna (Soria, Spain), has already yielded 30 species of mammals. Hipparion fissurae, described here is more dolichopodic than other Hipparion. It is related to Hipparion crusafonti from Villaroya (Villafranchian), but more slender between other characters.
  

The entelodonts of the Quercy phosphorites are essentially known from isolated teeth. The skull remains attributed to Entelodon magnum by Kowalewsky in 1876 and the type left mandible of Enteladon depereti RÉPELIN, 1918 do not belong to this family. From a morphologic point of view, the teeth from Quercy are quite comparable to those from the type localities of E. magnum and E. deguilhemi. In the absence of distinctive anatomical criteria between E. magnum (small species from the level of Ronzon) and E. deguilhemi (large species from the level of Villebramar) they are referred, according to their size, to one or the other of these two species. However, some of them are intermediate in size. Their existence speaks in favor of a close relationship between E. magnum and E. deguilhemi and confers a real stratigraphic interest to these entelodonts of Western Europe. Finally, some anatomical characters (notch on the mesial border of P4, presence of a paraconid joined to the metaconid, trigonid higher than talonid) considered until now as peculiar to the North American Archaeotherium and not yet cited in its European contemporary Entelodon have been brought to light on the teeth from Quercy.

  

New Dendromus species (Rodentia, Muroídea) from Langebaanweg (Pliocene, South Africa). Stratigraphical and paleoecological consequences.

Two new species of Dendromus are described from the Langebaanweg site which precises the evolutionary trend among this genus in South Africa and gives further paleoenvironmental indications. Two evolutionary stages are described: D. darti nov. sp. shows low-crowned molars with bunodont cusps and its more closest relative would be D. melanozis from the Cape region. On the contrary, D. averyi nov. sp. is more lophodont and is better related with the modem D. melanotis. Both species are at a less evolved stage than the Dendromus sp. from Laetolil Beds at Laetoli. The Langebaanweg deposits cannot still be dated by biostratigraphy but they clearly cannot be older than the basis of Pliocene times. The association of Dendromus and Mystromys in the same levels indicates a grassland environment with woodland patches as well as probable swamps. 

The locality of St.-Victor-la-Coste (Gard) has yielded, rather abundantly, teeth of two glirids hitherto very poorly known: Glirudinus praemurinus (Freudenberg) and Glirudinus glirulus (DEHM). It has permitted, moreover, new views on the evolution of Peridyromys murinus (POMEL). Study of these forms confirms the late Oligocene age of the fauna, without allowing, however, further precision.