This paper is mainly devoted to the systematics of rodents from two middle Eocene (Bartonian) localities: Chéry-Chartreuve and Rocourt-Saint-Martin (Aisne, Eastern Paris Basin). These two localities are stratigraphically located slightly above the Auversian sands. The two faunas, which comprise 11 and 8 taxa, respectively, are very different in their composition. That of Rocourt-Saint-Martin shows strong similarities with that of the geographically very close locality of Grisolles, referred to the MP16 mammalian Reference level. The very distinct fauna of Chéry-Chartreuve includes a new species of Ailuravinae, Ailuravus nov.sp, and some teeth of the theridomyid Protadelomys, which represent archaic elements in the fauna. The most abundant species of the locality represents a new genus of primitive Theridomyidae. The presence of some teeth belonging to a new species of large Remyinae, Remys nov. sp., of Elfomys engesseri HOOKER & WEIDMANN, and a population of small dimensions referred to the genus Estellomys allow a correlation with Les Alleveys (Switzerland), with however some differences that would indicate an older age for Chéry-Chartreuve. Situated at the base of the "Marinesian" from the Bassin de Paris, this fauna is unquestionably different from those referred to the MP16 reference level and could represent an older level for which the macrofauna remains very poorly known. Conversely, the comparison of rodents from La Livinière II with those present in MP16 faunas, especially those of Robiac (Gard), shows a great similarity between both localities. This casts doubts on whether to keep this La Livinière II faunule to define the current MP15 reference level, while the biostratigraphical position of Pontils 26 (Spain), previously referred to this level, is reconsidered. Chery Chartreuse could be a good candidate for a new definition of the MP15 reference level. 

Fourteen distinct phyletical lineages which belong at least in three families: Ischyromyidae ALSTON, 1876, Gliridae THOMAS, 1896 and Theridomyidae ALSTON, 1876, have been identified after the study of more than 3600 rodent dental remains from about twenty Early and Middle Eocene european localities. A systematical and phylogenetical revision of these rodents has been achieved. Nearly all the specific and generic diagnosis are emended. Several new combinations and synonymies are proposed. Four new species and two new genera, Euromys nov. (Ailuravinae) and Hartenbergeromys nov. (Microparamyini), are named and described. Euromys nov. gen. is known by three distinctive ypresian (MP 7 to MP 10 european reference levels) chronospecies. This new lineage is thought to be the direct ancestor of Meldimys MICHAUX, 1968 and Ailuravus RUTIMEYER, 1891. A new species of the genus Plesiarctomys BRAVARD, 1850, Pl. lapicidinarum from Condé-en-Brie (MP 8-9 reference level), allows to relate the Plesiarctomys lineage to the Pseudoparamys MICHAUX, 1964 one. The taxa Sparnacomys HARTENBERGER, 1971, Pantrogna HARTENBERGER, 1971, and Corbarimys MARANDAT, 1989 are erected to genus rank; the last one is not thought to be an Ischyromyidae. A new chronospecies of Pantrogna, P. marandati nov. sp. from the locality of Prémontré (MP 10 reference level), is described. This lineage is at the origin of two others, namely Masillamys TOBIEN, 1954, including M. mattaueri (HARTENBERGER, 1975) nov. comb. (MP 10 reference level), and Hartenbergeromys nov. gen., known from MP 10 (H. hautefeuillei nov. sp.) and MP 11 (H. parvus TOBIEN, 1954) reference levels. The phylogenetical position of Hartenbergeromys nov. gen., at the origin of the european family Theridomyidae, is discussed. The systematical and phylogenetical status of two probable Paramyinae, "Paramys" woodi MICHAUX, 1964 and an unnamed genus and species, are discussed. New populations of the primitive Gliridae Eogliravus HARTENBERGER, 1971 and of the primitive Theridomyidae Protadelomys HARTENBERGER, 1968, are described and assigned to previously known species.

  

The earliest North American rodents occur in basal Clarkforkian beds of the Fort Union Formation at Big Multi Quarry near Bitter Creek, northern Washakie Basin, Sweetwater County, Wyoming, and in closely correlative Fort Union beds formerly accessible in the Eagle Coal Mine near Bear Creek, northern Clark's Fork Basin, Carbon County, Montana. Two new species of early Clarkforkian rodents, Paramys adamus and Alagomys russelli, are described from Big Multi Quarry. Paramys adamus is represented by virtually complete upper and lower dentitions, which demonstrate that this species is one of the most primitive North American paramyids yet discovered. These specimens form the basis for a reevaluation of the content and stratigraphic range of P. atavus, which is known with certainty only from Bear Creek. Alagomys russelli is the first North American record for the enigmatic rodent family Alagomyidae, otherwise known from ?late Paleocene-early Eocene localities in Mongolia and China. Phylogenetic analysis of dental and gnathic traits suggests that Alagomyidae form the sister group of all other undoubted rodents. At least two rodent clades, alagomyids and basal paramyids, seem to have invaded North America from Asia at the beginning of Clarkforkian time, but only the paramyids persisted to undergo a significant evolutionary radiation in North America. 

The status of the giant bird taxa Liornis floweri and Callornis giganteus from the Santa Cruz Formation (late Early Miocene) of Patagonia, first described by Ameghino (1895) is reassessed on the basis of a re-examination of the type material at the Natural History Museum, London. Liornis floweri, which lacks a Pons supratendineus on the tibiotarsus and has an unbifurcated Canalis interosseus distalis on the tarsometatarsus, is clearly a brontornithid and is considered as a junior synonym of Brontornis burmeisteri. Ameghino’s replacement of Callornis by Eucallornis is unjustified. Callornis giganteus is a chimera based on a phorusrhacid tarsometatarsus (probably belonging to Phorusrhacos longissimus) and a brontornithid tibiotarsus. The latter can be considered as the lectotype of Callornis giganteus, which may represent a small morph of Brontornis burmeisteri or a distinct taxon. It is referred to here as Brontornithidae indet. The tarsometatarsus described by Dolgopol de Saez (1927a,b) as Liornis minor and considered by her as a gracile brontornithid apparently has a bifurcated Canalis interosseus distalis and should therefore be placed among the Phorusrhacidae. 

Ornithocheirid pterosaurs were the largest of the toothed pterodactyloids and had a worldwide distribution, although their fossil record is fragmentary, with the exception of the north-eastern Brazilian Crato and Santana Formations (Aptian, ?Albian, Early Cretaceous). With Istiodactylidae, they were also the only toothed pterosaurs that survived into the Cenomanian (Late Cretaceous), becoming extinct at the end of this period. Here we report on an ornithocheirid metacapus from the Late Cenomanian laminated limestone of north-eastern Mexico discovered about 120 km north-west of Ciudad Acuña, northern Coahuila at the south banks of Rio Bravo. The specimen comprises a fragmentary distal syncarpal, a crushed but complete metacarpal IV, two fragmentary preaxial metacarpals and a possible fragmentary terminal left wing finger phalanx. It represents the geologically youngest known ornithocheirid worldwide. We suggest that ornithocheirid pterosaurs may have become extinct because of massive sea level fluctuations during the mid to late Cretaceous that may have obliterated their breeding sites on coastal plains and low lying islands. 

The tooth morphology of the Thalattosuchia (marine crocodilians from the Jurassic and the Early Cretaceous) is analysed. The Callovian from Poitou and the Kimmeridgian from Quercy have yielded many remains of Metriorhynchus, Steneosaurus and Machimosaurus. These remains allow us to study the variations of tooth morphology during ontogenic growth, tooth replacement and the location of the teeth. We have defined different tooth types for these genera. In Metriorhynchus, the two tooth types defined do not coincide with the two groups recognized in the Callovian (broad-skulled and narrow-skulled metriorhynchids) but reflect the prey preferences of these forms. In Steneosaurus and Machimosaurus the five tooth types deñned are in agreement with the main taxa known from the Bathonian to the Early Cretaceous. This study allows to precise the function and the prey preference of the Thalattosuchia during the Jurassic and the Early Cretaceous.

  

The dentition of selachians is characterized by an often very pronounced heterodonty involving a great morphological diversity. Despite this fact, the dentitions of selachians can be grouped in a rather reduced number of dental types corresponding to trophic adaptations: grasping, tearing, cutting, crushing, grinding and grasping-grinding type. The numerous exemples of convergence and parallelism that can be observed in fossil selachians and between Recent and fossil ones is the result of this reduced number of dental types. These dental specialisations allow to try a reconstruction of the way of life of fossil forms. 

Bats are among the most numerous and widespread mammals today, but their fossil record is comparatively meagre and their early evolution poorly understood. Here we describe a new fossil bat from dental remains recovered from late Early Eocene sediments at Prémontré, northern France. This 50 million-year-old bat exhibits a mosaic of plesiomorphic and apomorphic dental features, including the presence of three lower premolars, a single-rooted p3, short p4 with metaconid, myotodont lower molars and a tall coronoid process of the dentary. This combination of features suggests it is an early member of Vespertilionidae, today’s most speciose and geographically widespread bat family. The Prémontré bat has bearing on hypotheses about the origins of vesper or evening bats (Family Vespertilionidae), as well as crown-group chiropterans.


  

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 litoptem Proterotheriidae are extinct endemic South American ungulates frequently used as an example of evolutionary convergence with the horses. They were considered to be exclusively Tertiary representatives with the youngest record being in the late Pliocene, before the appearence of the equids and cervids during the Great American Interchange. Two undoubted Pleistocene records in Argentina and the specimen here described from Uruguay, confirm the persistence of the proterotherids into that period. In the Quaternary, these ungulates are found outside the typical pampean region and probably were confined to a few northern and warmer more forested relictual microhabitats.

  

The middle Palaeocene of São José de Itaboraí (State of Rio de Janeiro, Brazil) has yielded a very rich and diverse snake fauna which includes Madtsoiidae, Aniliidae s.l., Boidae, Tropidophiidae s.l., Booidea incertae sedis, and Russellophiidae. The present article (part I) deals with Madtsoiidae and Aniliidae s.l. Madtsoiidae are represented by many vertebrae and a few skull bones. They comprise one new species assigned to the genus Madtsoia (M. camposi sp. nov.). However, the definition of the genus Madtsoia is unsatisfactory and the generic allocation might be provisional. A few elements, vertebrae only, belong to the Aniliidae s. l. Two taxa are referred to this latter group: Coniophis cf. C. precedens and Hoffstetterella brasiliensis gen. et sp. nov. The forthcoming part II will deal with Boidae.

  

Extensive collecting in previously known and new Middle Eocene rodent localities in the Subathu Group of the Rajauri and Reasi districts, Jammu and Kashmir (northwest Himalaya, India) has yielded over 500 isolated cheek teeth, numerous incisors and cheek tooth fragments, and two mandibular fragments with M/1-M/3. An analysis of this additional material and a restudy of that reported earlier from the Metka (Rajauri) area has revealed the presence of diverse Middle Eocene ctenodactyloid rodents in India. The assemblage comprises six genera and twelve species, viz.., Bírbalomys woodi, B. ibrahimshahi, B. sondaari, Basalomys vandermeuleni, B. ijlsti, B. lavocati, Chapattimys wilsoni, C. debruijni, Gumbatomys asifi, cf. Advenimus bohlini, cf. Petrokoslovia sp. indet. 1 and cf. Petrokoslovia sp. indet. 2. Most species of this assemblage are endemic to the Indian subcontinent and are referred to the Family Chapattimyidae. However, at least three taxa, viz.., cf. Advenimus bohlini, cf. Petrokoslovia sp.indet. 1 and cf. Pezrokoslovia sp. indet. 2 show close affinities with their Central and Eastern Asiatic contemporaries and are attributed to the Family Yuomyidae. The dentitions of most of the Middle Eocene rodents from India are described here in better detail thus providing more precise characterization than was available earlier. The taxonomie status of Basalomys, formerly a subgenus of Birbalomys has been raised to that of a genus. Basalomys vandermeuleni is here designated as the type species of Basalomys in place of B. ijlsti. The rodent assemblage from the Subathu Group is specifically similar to that from the coeval Kuldana Formation of Pakistan. The absence of Birbalomys ibrahimshahi and cf. Advenimus bohlini in Pakistan is related to sampling bias. The identifications of the Early Eocene rodents from Barbara Banda, Pakistan have been reexamined and the familial assignments of some of the Early and Middle Eocene rodents from Central and Eastern Asia have been discussed and reviewed in the light of recent discovery of early rodents in Mongolia.

  

A comparative bone histological study of the three known genera of the endemic European ornithopod dinosaur family, Rhabdodontidae, is presented here in an ontogenetic context. Investigated specimens were assigned to different ontogenetic stages based exclusively on the histological indicators of osteologic maturation during diametrical bone growth; an entirely size-independent method as opposed to most previous studies. Qualitative comparison of bone histology of corresponding ontogenetic stages and elements among the three valid rhabdodontid genera, Mochlodon, Zalmoxes, and Rhabdodon, revealed some consistent patterns. Genus specific histological differences within Rhabdodontidae are most expressed between Rhabdodon and the Mochlodon-Zalmoxes clade. These indicate a prolonged phase of fast growth and a less constrained cyclicity in the growth dynamics of Rhabdodon, as opposed to the slower and more regulated growth strategy reflected in the bones of Mochlodon and Zalmoxes. These genus specific differences are consistent with the phylogenetic interrelation of the genera and are most probably related to the pronounced differences in body size. However, when compared to other ornithopods, most detected histological features in rhabdodontids do not seem to reliably reflect either phylogenetic relations or body size. A notable common feature of all rhabdodontid genera irrespective of body size is the ontogenetically early onset of cyclical growth and secondary remodelling; a pattern that more resembles the condition found in derived ornithopods than that described in more basal taxa which are closer relatives of rhabdodontids. The recognition of taxon-specific histological patterns as well as patterns indicative of ecological and thereby functional traits clearly requires more accurate, preferably quantitative evaluations.   

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. 

The genus Cryptomeryx SCHLOSSER, 1886, inusited during a long period, has been discovered in Lower and Middle Oligocene localities of the Quercy region (South-West France). This new material, as well as specimens from the old collections referred to Cryptomeryx, are described; their study, allows us precising the definition of the genus, and confirming its allocation to the Tragulidae family. The type species of the genus, Crypmmeryx gaudryi (= Lophiomeryx gaudryi FILHOL, 1877), occurs in several localities at the base of the Middle Oligocene (Itardies, La Plante 2, Roqueprune 2, Soulce, Herrlingen 1). The new species C. matsoui n. sp. has been defined in the older locality of Mas de Got (top of Lower Oligocene). It is possible that the species Pseudamphimeryx decedens STEHLIN, 1910 pertains to the same genus. Also to the Tragulids must be referred the monospecific genus Iberomeryx (I. parvus GABOUNIA, 1964) from Upper Oligocene of Benara (Georgie, URSS), with which Cryptomeryx is related. These genera are not direct ancestors of Miocene tragulids; their occurrence in the Western European Oligocene results from a first immigration wawe of the family. These Tragulids are one of the most archaic groups of Ruminants. They are probably derived from a primitive stock which had acquired in Asia the selenodont condition of the dentition. 

The description of new material from three classic middle Eocene localities of western Europe permits the addition of details to the systematics of primitive Pseudosciurids. The cranial anatomy of Protadelomys cartieri (STEHLIN and SCHAUB) from Egerkingen is described here and compared to that of the Adelomyines from the late Eocene, until now the only ones known. The morphologic and biometric study of the dentition of P. cartieri compared to that of P. alsaticus n. sp. from Bouxwiller and P. Iugdunensis n. sp. from Lissieu, forms respectively older and younger than P. cartieri, permits the evolutionary tendencies of the group to be demonstrated and shows that notable differences in age exist between these localities. This ensemble of forms can constitute a valuable guide lineage in the establishment of a fine stratigraphy of the period. Other less well known lineages are present at Egerkingen along with P. cartieri. They can be related to genera that have been noted int he late Eocene. In conclusion, a criticism of recent zonation proposals, made by divers authors, completes this article. 

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. 

The discovery in the Turolian (Late Miocene) of Dytiko 3 (Macedonia, Greece) of a complete skull with mandibles and cervical vertebrae, atlas and epistropheus, is a very important contribution to the knowledge of the subfarnily Chalicotheríinae. After the description, the comparison with other specimens of Miocene chalicotheres permits the revival of the generic name Macrotherium with a new species M. macedonicum. This genus is mainly characterized by a short snout and an inflated cerebral skull. It coexists during the Miocene with Chalicotherium. A cladistic analysis leads to conclusion that the species which has been described from the Early Middle Miocene of Rusinga must be identified as the type-species of a new genus: Butleria.





  

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. 

Based on material from 30 localities, morphologic dental, cranial and biometric analyses have permitted the characterization of two parallel Issiodoromyine lineages, and also the definition of diverse species representing several evolutive stages. Thus it is that new lineages complete the contribution made by the Theridomyinae and Cricetidae and permit, for the Quercy in particular, additional precision in the biochronologic succession of the localities. One of the lineages is limited to the genus Pseudoltinomys LAVOCAT; the other evolves from the genus Elfomys HARTENBERGER to the genus Issiodoromys BRAVARD in GERVAIS. The latter is affected by profound anatomical changes due to a functional modification of the mastication apparatus. These changes seem to be able to be put in relation with the aridification and cooling of the climate at the end of the Eocene. At the end of the middle Oligocene, a new chewing structure is achieved. It is found in diverse living rodents that inhabit a rather arid steppe environment (Cavia, Pedetes, Ctenodactylus). To these supposed nearby ecologic conditions, these rodents have responded in a convergent fashion. It is possible to attribute to the extreme specialization of Issiodoromys its incapacity to adapt to the new climatic crisis of the end of the Oligocene. The arrival of immigrants may be considered as another cause of its disappearance at this time, complementary or not with the first.