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. 

Near "la Lieude", in the Lodève basin, more than a thousand of footprints are distributed in a twenty of trackways which amounts to 220 m length. They have been found on calcareous siltstone level in the B site named also "Réserve Naturelle Volontaire". This last is located in the Saxonian summit dated Upper Permian. "La Lieude" tracks are described by using statistical methods then they are compared with others from the world Permian. What allows to distinguish 4 following ichnotaxa: Lunaepes ollierorum nov.ichnosp., Merifontichnus thalerius nov. ichnogen. and nov. ichnosp., Planipes brachydactylus nov.ichnosp. and Brontopus circagiganteus nov. ichnosp. All these traces are attributed with possibility or probability to Therapsida or to Therosauria, except Brontopus circagiganteus nov. ichnosp. that could be due to Caseamorpha. All these animals whose sizes have been estimated between l and 5 m lived probably in a playa environment.The biological and sedimentological data from "la Lieude" footprints levels compared with informations provided by the tracks orientations, suggest the following scenario. Animals coming from the North have crossed a sandy channel bank with plants zones by directing to the South for the majority. Maybe, they were going to the lacustrine part of the playa, close to "la Lieude" footprints that they have just trampled on. 

A strong lophodonty, an extreme heterodonty, some hypsodonty and regular overlayings of coronal cement are prominent features of the genus Leptolophus (Palaeotheriinae = Palaeotheriidae s.s.). The histological pattern of the teeth unusually joins type II enamel prisms, characteristic of advanced ungulates, together with archaic features, such as an almost complete lack of Hunter-Schreger zonation and a weak expanse of peritubular dentine. The skull is narrow and slender, with an elongated ante-orbital facial region, a moderately notched nasal aperture, a rather elongated post-canine diastem, parallel zygornatic arches and a fairly dorsally located squamoso-mandibular joint.The functional analysis brings to light "ectolophodont" masticatory cycles with two phases, in which maximum power was applied, contrary to equíds, on hindmost teeth; likewise, skull accomodations to increasing height of the teeth are quite different. This study leads to the assumption that Leptolophus may have been light mammals, living in rather open surroundings, browsing on herbaceous plants or leaves cropped close to the ground. Moreover, it appears that it could have been some inadequacy of dental structures to the dietary, which leaded to quick wear of the teeth and to many enamel notches, but had been somewhat balanced by the early increase of hypsodonty, not induced in such a case by a biotop deterioration (as it will happen at the end of the Eocene). This ínadaptation might account for the short duration of the genus Leptolophus, whose the 3 species, L. stehlini, L. nouletí and L. magnus n. sp. are indeed confined in the level MP 16. Its geographical spreading (as far as known, South of western Europe) and the morphological pattern of its dentition suggest that this genus would have been related to early upper Eocene endemic spanish forms.





  

Pangolins have never shown a high taxic diversity and their fossil record is scarce. We report here the first discovery of a partial humerus from late Oligocene deposits in Pech du Fraysse (MP28, France). The new specimen from Pech du Fraysse is described and compared to various extant and extinct species of pangolins. It shows a suite of morphological features very similar to the humeri discovered in Saint-André (MP 26), Solnhofen (Burdigalien), and Saulcet (Aquitanian), attributed here to Necromanis franconica. The description of the specimen from Pech du Fraysse allowed us to discuss the systematics of Paleogene and Neogene pholidotans. The differences between PFY 4051 and N. franconica on the one side, and N. quercyi on the other side, might be sufficiently important to justify a generic distinction. A comparison with extant species showed that N. franconica was likely terrestrial and fossorial based on its humeral morphology. 

A detailed comparative study of a complete skeleton of Bachitherium and a cladistic analysis of the sub-order Neoselenodontia lead us to propose a cladogram and a new classification of this group. The Tylopoda are the sister-group of the Ruminantia, which are chiefly defined by the fusion of the cuboid and navicular. Within this infra-order, Amphimeryx is the sister genus of a tetraselenodont group, in which the Hypertragulidae are well-separated group from a monophyletic group defined by the loss of trapezium, fusion of capitatum and trapezoid, and the isolation of the hypoconid on lower molars. The most primitive genera of this group, Lophiomeryx and Iberomeryx still have an open trigonid on the lower molars, but this is lingually closed in Archaeomeryx, sister-genus of the higher Ruminantia which have fused metatarsals and more evolved milk teeth. We divide them into two pan/orders : Tragulina (including the recent and miocene Tragulidae, and the North-American Leptomerycidae), and Pecora, with reduced lateral metacarpals and a new crest (telocristid) on the lower premolars. Within the Pecora, the upper molars of Gelocus are more primitive than those of Bachitherium (a genus with many autapomorphies in the dentition) itself more primitive than the group Prodremotherium + Eupecora, with fused metacarpals. We consider the Eupecora (including several genera without frontal appendages) to be monophyletic. 

The first lists of Insectivores (Erinaceidae, Talpidae and Soricidae) from the Pliocene beds of Southern France and North-East Spain are given in this paper. The material from twelve localities is studied. These localities are geographically situated in Languedoc (Celleneuve, Vendargues, Nîmes, Sète, Balaruc 2 and Seynes), in Roussillon (Terrats, Serrats-d'en-Vacquer, Château d'eau and Mont-Hélène) and in North-East Spain (Layna, Medas Islands and Puebla de Valverde). These faunas correspond to the Early, Middle and Late Pliocene. 1 to 8 taxa are identified in these localities and 14 specific taxa are presently listed for this period in this area. Two new specific taxa are described as Galerix depereti nov. sp. from all the Early Pliocene localities in the North-Pyrenean area and as Desmanella gardiolensis nov. sp. from Balaruc 2. For this small mammals, two faunal assemblages are recognized. The first one is dated from the Early Pliocene (F 1, 2 and 3 zones in Aguilar et Michaux) and is characterized by Galerix depereti and rare and little diversified Soricids. The second one is Late Pliocene in age (zones G 2 and G 3). The fossils of the genus Talpa are relatively abundant and the Soricids are diversified and very abundant. The Middle Pliocene (zone G 1) is a transitional period. ln these faunas, most of the insectivore genera are known from the European Late Miocene beds (8 on 10). This fact demonstrates a relative continuity between the invectivore faunas from the Late Miocene to the Early Pliocene. In conclusion, somme paleoecological considerations are suggested.
  

The first good skeletal remains of Homogalax protapirinus from the Wasatchian of the Bighorn Basin, Wyoming, indicate that this primitive tapiromorph was more plesiomorphic in many features than primitive equoids including Hyracotherium. Compared to Hyracotherium, Homogalax more closely resembles Phenacodonta (the closest outgroup of Perissodactyla for which postcrania are known) in various details of articular surfaces, muscle attachments, and proportions of the humerus, manus, and pes.Among known taxa, Homogalax most nearly approximates the plesiomorphic postcranial skeletal anatomy of Perissodactyla. 

Three new fossiliferous localities, two of karstic origin, Castelnou 3 and Font Estramar, respectively Late Upper Miocene and Lower Pliocene, and one of lacustrine origin, Thuir, Lower Pliocene, add data about the transition between Miocene and Pliocene faunas of rodents in southern France. An unexpected association of taxa was present in the late Upper Miocene, including between others, Myocricetodon, Hispanomys, Ruscinomys, Cricetus barrierei, Promimomys and a new species of Stephanomys, S. dubari nov. sp. Myocricetodon is still known in the Lower Pliocene. It is shown that the large field-mice known since the Late Upper Miocene belong to two different lineages, on one side, A. jeanteti, on the other side, A. gudrunae followed by A. gorafensis. Biochronological and biogeographical implications are discussed. 

Mammal-bearing localities have been discovered in the marine and lacustrine series of the middle Ilerdian (Lowermost Eocene) from Southem France (Minervois and Corbières). In the localities of Fordones, Monze, Fournès, and La Gasque, thirty mammal species have been identified. Among others, they include ischyromyid rodents (Microparamys and Pseudoparamys), paromomyid and adapid primates (Arcius and Donrussellia), new insectivores, condylarths, and a dyspternine pantolestid. These faunas provide new informations on the early Eocene Mesogean faunas of Rians and Palette. The assemblages of primates and rodents from Fordones support good  correlations with Palette which was recently placed near the standard-level of Dormaal (MP 7). In fact, Palette and Fordones could be even older than Dormaal. Consequently, there seems to be a relatively important temporal gap between the late Paleocene of Cernay and the Sparnacian of Dormaal. This gap could be partly filled with the Mesogean faunas of Palette, Fordones, and Silveirinha. On the basis of these new mammal faunas the marine middle Ilerdian is proved to be older than the Cuisian stage of the Paris Basin. With regards to the position of the Fordones fauna at the top of the NP 10 calcareous nannoplankton biozone, the westem European paleomammalogists Paleocene/Eocene boundary could be situated between the NP 9 and NP 10 biozones. 

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. 

HGL50 is a latest Early-early Middle Eocene vertebrate-bearing locality located in Western Algeria. It has produced the richest and most diverse fauna of amphibians and squamate reptiles reported from the Palaeogene of Africa. Moreover, it is one of the rare faunas including amphibians and squamates known from the period of isolation of Africa. The assemblage comprises 17 to 20 taxa (one gymnophionan, one probable caudate, three to six anurans, seven ‘lizards’, and five snakes). Two new taxa were recovered: the anuran Rocekophryne ornata gen. et sp. nov. and the snake Afrotortrix draaensis gen. et sp. nov. The locality has also yielded the first confirmed anilioid snake, the first Palaeogene gymnophionan, and probably the first caudate from the Palaeogene (and possibly from the Tertiary) of Africa. The presence of a caudate at that time in Africa would be of particular interest; unfortunately, the available material does not permit a definitive identification. The fauna comprises Gondwanan and more specifically West Gondwanan vicariants, probably autochthonous groups and a Eurasian immigrant (assuming that the identification of the caudate is accurate). The fauna from HGL50 is clearly distinguished from the few other Eocene assemblages of Africa. However, if this results largely from differences in geological ages, geographic positions of the localities and mainly differences in environments took a part in the composition of the faunas. 

The mammalian fauna (represented only by isolated teeth) from the early Cretaceous of Morocco includes a number of non-'therian' specimens, characterized by the antero-posterior alignement of the main molar cusps. This assemblage is very heterogenous, including forms (among them Gobiconodon palaios sp. nov.) that can be related to Laurasian triconodonts and several forms so far known only in this part of Gondwana (among them Kryptotherium polysphenos gen. et sp. nov.). The latter, in particular, exemplifies the diversity of the mammalian dental morphology in the Mesozoic, often largely underestimated. In the last part of this paper, the 'therian' affinity of the 'amphilestids' is discussed, on the basis of the arrangement of molar cusps, the interlocking mechanism and the occlusal pattem. 

The middle Palaeocene of São José de ltaboraí (State of Rio de Janeiro, Brazil) has produced a rich and diverse fauna of boid snakes. It comprises six or seven species: Hechtophis austrinus gen. et sp. nov., Corallus priscus sp. nov., Waincophís pressulus sp. nov., Waincophis cameratus sp. nov.,"Boinae A", and "Boinae B". Moreover, two dentaries might pertain to either H. austrinus or "Boinae B", or even represent a distinct taxon. Hechtophis austrinus is assigned, with reservation, to the Erycinae. All other taxa are referred to the Boinae. The vertebrae of all taxa have paracotylar foramina, which raises the problem of the apomorphic or plesiomorphic nature of this feature. This fauna also raises the question of the presence of extinct erycine boids in South America, but it does not allow this question to be settled. 

The principles and practices employed in establishment and recognition of South American land mammal ages are reviewed along with previous and present concepts of distinguishing time, rock, and faunal units. Previous chronological arrangements of South American Tertiary land mammal faunas are appraised on the basis of recent geological and paleontological data. Twelve South American Tertiary land mammal ages are here recognized [from oldest to youngest, Riochican (middle to late Paleocene); Casamayoran (early Eocene); Mustersan (middle Eocene); Divisaderan (late Eocene); Deseadan (early [to middle?] Oligocene); Colhuehuapian (late Oligocene); Santacrucian (early Miocene); Friasan (middle Miocene); Chasicoan (late Miocene); Huayquerian (latest Miocene); Montehermosan (early to middle Pliocene); and Chapadmalalan (late Pliocene)]. As all except the Friasian were originally defined on the basis of Argentine faunas, these are discussed first and at length, and each is reviewed with discussion of type locality, stratigraphy, type fauna, and faunal correlations. Non-Argentine faunas are then discussed country by country in alphabetical order.

     A review is given of radioisotope dates obtained on volcanic rocks (i.e., basalts, tuffs) associated with mammalbearing beds in Argentina. Based on these age determinations and on correlation of the late Tertiary land mammals involved in the interchange between North and South America, a chronology of South American land mammal ages correlated with North American land mammal ages and European marine stages is proposed.

It is concluded that South America was an island continent through most of the Tertiary Period (ca 65 to about 3 Ma). As a result, the land mammal fauna of South America developed in isolation and was dominated by autochthonous endemic groups. Toward the end of the Tertiary (i.e., middle Miocene) a unique faunal balance had been achieved by the descendants of the ancient inhabitants (notoungulates, litopterns, condylarths, astrapotheres, edentates, marsupials) and of later (late Eocene) waif immigrants (caviomorph rodents, platyrrhine primates). A prominent feature of this mammal fauna was the combination of carnivorous and omnivorous marsupials with native placental herbivorous ungulates, subungulates, and edentates.

Sometime during the late Miocene, a limited but important interchange of mammalian taxa between North and South America took place. Procyonids (raccoons and their allies), a group of North American origin, first appear in South America in strata of Huayquerian Age, while members of the extinct South American ground sloth families Megalonychidae and Mylodontidae first appear in North America in early Hemphillian time. These groups dispersed along island arcs before the appearance of the Panamanian land bridge in the Pliocene (ca 3.0 Ma). Cricetine rodents, a group of North American origin, are first known in South America in strata of Montehermosan Age. The known taxa are too advanced and diversified to be considered the first of this group to invade South America. lt is believed by some workers that these rodents arrived before the Montehermosan, possibly in the late Miocene or earlier, by waif dispersal from North America.

The isolation of South America ended with the appearance of the Panamanian land bridge, which provided a direct, dry land connection between the two Americas. Across this portal an extensive interchange of terrestrial faunas occurred, and the fossil record documents an intermingling of these long-separated land mammals faunas.

      The beginning of this interchange by land route in South America is marked by the appearance of mammals which evolved from North American emigrants in the Chapadmalal Formation of Argentina. These include a mustelid (Conepatus), a tayassuid (Argyrohyus), and four genera (Akodon, Dankomys, Graomys, Reithrodon) of cricetine rodents. The appearance of this contingent of northern animals favors the existence of the Panamanian land bridge by this time. Likewise, a large number of terrestrial vertebrates of South American origin appear in North America in beds of late Blancan Age date around 2.7 Ma. Among the mammals are Neochoerus, Erethizon, Glyptotherium, Glossotherium, Kraglievichia, and Dasypus. 

Morphological descriptions are given of Eocene mammals from the locality Toru Ajgyr (NEKyrgyzstan) that were excavated in 1997 and 1998 in a cooperation between the Martin-Luther-University Halle (Germany), the Zoological Institute in St. Petersburg (Russia) and the Seismological Institute in Bishkek (Kyrgyzstan). The species found belong mostly to perissodactyls, as Lophialetes sp., Teleolophus sp. and brontotheres. The primitive ungulate family Olseniidae is represented by a complete foot skeleton of cf. Olsenia sp. In addition, postcranial materials of Gobiatherium mirificum (Dinocerata) and of artiodactyls have been collected and are described herein. Based on mammals, the locality is part of the Asian Land Mammal Age Arshantan and is stratigraphically equivalent with the Bridgerian Land Mammal Age in North America and with the lower and middle Geiseltalian of the European Middle Eocene. 

This paper provides a revision of the early and middle Eocene European rodents previously referred to as Ischyromyoidea, including taxa considered to be at the origin of the Theridomorpha. The use of an accurate dental terminology and a better understanding of the size and shape of their infra-orbital foramen (i.o.f.) led us to a substantial revision of this group, which allowed to better characterize them and to appreciate their variability. On these bases, phylogenetic analyses (cladistic and standard Bayesian
approaches) of early Ypresian to late Priabonian European rodent species were undertaken in order to highlight the root of the early Theridomorpha and its content. In this paper, the phylogeny was established based on 343 characters (338 dental) through 45 early Paleogene taxa using both cladistic and bayesian analyses. The ingroup included on one hand a few North American genera (Reithroparamys, Microparamys, and Acritoparamys) and European ones (Eogliravus, Ailuravus, Corbarimys, Meldimys, Euromys, Plesiarctomys, and Pseudoparamys) considered until now as being related with the North American superfamily Ischyromyoidea. On the other hand, it included genera close to the root of the Theridomorpha (Sparnacomys, Pantrogna, and Hartenbergeromys) and early Theridomyoidea (Masillamys, Protadelomys, and some Pseudosciuridae). The phylogenetic results obtained via the two
distinct reconstruction approaches are consistent in virtually all relationships. The proposed systematics here derives from these phylogenetic results. This phylogenetic context led us to change the suprafamilial, familial, subfamilial or generic attribution of several species. Characters of Theridomorpha, like the obliquely developed postprotocristid allied with the occurrence of a metalophulid I, have been found in genera previously considered as Ischyromyidae (Pseudoparamys, Euromys, Sparnacomys, Meldimys, Pantrogna, and Hartenbergeromys) as well as the large i.o.f., when preserved (Pseudoparamys, Hartenbergeromys, and Masillamys). Based on these morphological observations and new phylogenetic considerations, the content of the Theridomorpha clade is here enlarged, thereby extending back the first theridomorph radiations to the early Eocene. Aside, a new taxon (Reinomys rhomboides gen and sp. nov.) is described from Avenay. In addition, a new genus, Auroremys, is created for the species subita (Comte et al., 2012) from Chery-Chartreuve. 

Seventeen species of cricetid rodent are recognized and described from lower and middle Siwalik deposits in the Potwar Plateau of Pakistan. These species are grouped in three categories, characterized as subfamilies (e. g., Megacricetodontinae, Myocricetodontinae, and Dendromurinae); an additional and more abundant category of rodents from these deposits, the Democricetodontinae, is excluded from this study, and will be described in a later study. Fifteen of the species are new, and four new genera are described. The Siwalik cricetid taxa are : Megacricetodon aquilari, n. sp.; Megacricetodon sivalensis, n. sp.; Megacricetodon daamsi, n. sp.; Megacricetodon mythikos, n. sp.; Punjabemys downsi, n. gen. & n. sp.; Punjabemys leptos, n. gen. & n. sp.; Punjabemys mikros, n. gen. & n. sp.; Myocricetodon sivalensis, n. sp.;  Myocricetodon sp.; Dakkamyoides lavocati, n. gen. & n. sp.; Dakkamyoides perplexus, n. gen. & n. sp.; Dakkamys asiaticus, n. sp.; Dakkamys barryi, n. sp.; Dakkamys sp.; Paradakkamys chinjiensis, n. gen. & n. sp.; Potwarmus primitivus, n. gen.; and Potwarmus minimus, n. gen. & n. sp. This diverse record of middle Miocene small mammals illuminates a profound radiation of cricetid rodents in southem Asia, the effects of which were felt in Europe and Africa as well as the rest of Asia. 

A fossil fish fauna, based on 5533 otoliths, from the La Paillade locality at Montpellier is described and figured. The otolith-bearing marls correlate to mammal zone MN l (Aguilar, 1982), and thus represent the earliest Miocene. The fish fauna consists of 30 taxa belonging to 20 families. Two species are new: Dussumieria sittigi and Liza gaudanti. The predominant faunal element is the Lesueurigobius vicínalis-species complex, composing 73% of all investigated otoliths. The palaeoecological analysis reveals a marine to euryhaline fish fauna living under tropical to subtropical conditions in the transition zone littoral - sublittoral. Water depth probably was more than 10 m. The scarcity of pelagic físhes suggests that the habitat was either a sheltered bay and/or far away from the open sea. Furthermore, some genera represented in the La Paillade fish fauna presently live exclusively in the Indopacific realm. Their presence strongly supports a broad connection between the Indian Ocean, the Mediterranean, and the Paratethys Seas during the Early Miocene (Aquitanian). From a palaeobiogeographical point of view, faunal relationships were found between the La Paillade fish fauna and both the Paratethys fish fauna and the fish fauna from the deposits in the Upper Rhinegraben and the Mayence and Hanau Basins (Germany).



  

A new Propalaeotherium species, clearly distinct from the genus Eurohippus, is described. It is characterized by having a similar size as P. voigti from the German Geiseltal localities (MP 11 to MP 13 reference-level), but differs in several features suggesting a slighty more derived morphology. It presents indeed less brachyodont crowns with less prominent and less elevated cingula, slightly larger relative surface of premolars, and a more marked metaconid splitting on cheek teeth. This new species is unknown from other European localities except the nearby Saint-Martin de Londres locality which has been considered older than the MP 13 level. 

The description of original material of anthracothere and proboscidean in the new locality of Bir el Ater 3 from East Algeria, and a thorough review of early Libycosaurus remains of Bir el Ater 2 allows us validating L. algeriensis as the smallest and earliest species of Libycosaurus and probably the earliest migrant of the genus from Asia. The presence of a Tetralophodon in the Neogene Nementcha formation might represent the earliest occurrence of the genus in Africa. These original fossil remains allow us to discuss the age of the Neogene part of the Nementcha formation close to the Serravalian/Tortonian boundary.