The remains of Ruminantia are very rare at Montredon. No specific determination was possible. We have only one Micromeryx, one small cervid, one tragocere and one (?) gazella. The faunal spectrum is in good correlation with the general retreat of the cervids in the European upper Miocene, but in contrast it is not convenient for the bovids, which are in expansion at the same time. For them, we have to invoke the local ecological conditions.
  

Investigations of the Clinchfield Formation (middle Eocene, upper Bartonian) exposed at the Hardie Mine (Wilkinson County, Georgia, USA), produced 4,768 actinopterygian otoliths representing 14 taxa and increased the number of bony fishes threefold from the site. The somewhat limited richness was characterized by bonefishes, mud eels, conger eels, sea catfishes, cusk-eels, snooks, grunts, drums and croakers, and porgies. The assemblage had a relatively even distribution with Ophidiidae, Congridae, and Sciaenidae most common. Included in the otolith taxa was a new sciaenid genus and species, Eosciaena ebersolei, with unknown relationships to other Sciaenidae. The Clinchfield otoliths were compared to other middle and late Eocene in age otolith assemblages in Alabama, Mississippi, and Louisiana utilizing percentage similarity measurements. Analysis indicated that the Clinchfield otoliths were not greatly similar or greatly unlike the Moodys Branch and Yazoo Clay otolith assemblages. However, the Clinchfield showed little relationship to the slightly older Lisbon Formation in adjacent Alabama and is postulated to be related to global climatic and plate tectonic events. Biostratigraphically, the Clinchfield otolith taxa are essentially the same as the other formations except for the Lisbon, which has at least ten unique species. Abundances of Clinchfield otolith taxa indicate a possible sub-bioprovince in the eastern Gulf Coastal Plain. The Clinchfield otoliths indicate a tropical to perhaps subtropical, soft substrate, mainly normal marine to slightly reduced salinities, inner shelf (0–20 m) paleoenvironment with indications of proximal continental coastlines. This investigation represents an initial step in addressing the immensely understudied Paleogene otolith assemblages in Georgia.
  

Isolated elasmobranch teeth (sharks and rays) from the middle Eocene (Bartonian) Bandah Formation in the Jaisalmer District of Rajasthan, India are described. The remains improve our knowledge of the environment represented by this lithostratigraphic unit and the ecology preserved therein. Seventeen unequivocal taxa were identified, including Nebrius sp., Striatolamia aff. S. macrota, Brachycarcharias atlasi, B. lerichei, cf. Jaekelotodus sp., Carcharhinus mancinae, Rhizoprionodon sp., Physogaleus sp., Galeocerdo clarkensis, G. eaglesomei, Odontorhytis aff. O. pappenheimi, “Rhinobatos” sp., “Dasyatis” sp., Coupatezia sp., “Aetomylaeus” sp., “Rhinoptera” sp., and Ouledia aff. O. lacuna. Of these, “Aetomylaeus” sp., B. atlasi, C. mancinae, G. clarkensis, G. eaglesomei, cf. Jaekelotodus sp., Nebrius sp., Odontorhytis aff. O. pappenheimi, Ouledia aff. O. lacuna, and “Rhinoptera” sp. are reported from the middle Eocene of India for the first time. The Bandah Formation elasmobranch palaeofauna has close affinities to the Palaeocene-Eocene Tethyan/Paratethyan faunas of Africa, Madagascar, Asia, and Europe, and some taxa indicate a western hemisphere influence from North America. The Bandah Formation palaeofauna indicates that deposition occurred in a moderately shallow marine environment. The Bartonian age is primarily based on foraminifera but is corroborated by the presence of elasmobranch taxa that also occur in contemporaneous deposits elsewhere. The marine regression started during the early Palaeogene, and our study indicates that the sea completely withdrew from the Jaisalmer Basin after the deposition of the Bandah Formation. This event may have been synchronous with the middle Eocene uplift of the Himalayan-Tibetan Plateau. 

Lalieudorhynchus gandi gen. nov. and sp. nov. is a new caseid synapsid from the Permian of the Lodève Basin, Occitanie, France. This new taxon is represented by a partial but well-preserved postcranial skeleton, and is characterized by the following apomorphies: a transverse section of the sacral and anterior caudal neural spines with a very thin keel-like process anteriorly, a slender dorsal tip of the dorsal and caudal spines, a narrow distal end of the first sacral rib, a fossa on triceps process of metacoracoid, and a very large distal tarsal 1 of same width than the astragalus, with nearly all sides being shallowly concave.
The skeleton corresponds to a sub-adult individual that was excavated from the La Lieude Formation dated as Roadian-Capitanian (Guadalupian). A sedimentological and taphonomical analysis of the type locality, together with preliminary osteohistological observations, suggest that this new French caseid was rather aquatic, as already hypothesised for other large forms.
A phylogenetic analysis of caseids is performed to test the position of this new taxon and to better understand the evolution of the clade: interestingly, Lalieudorhynchus gandi gen. nov. et sp. nov. is closer to the NorthAmerican “Cotylorhynchus” hancocki than to the other French caseids Ruthenosaurus and Euromycter from the Artinskian of the geographically closer Rodez Basin. These two last caseids document the Artinskian radiation of the clade, which remained diverse until Olson’s extinction. Caseids survived, as Lalieudorhynchus is one of the youngest representatives of the clade, and may have used novel ecological strategies to access their vegetarian food sources.

  

The present work is devoted to the study of the Cretaceous and Tertiary fishes (teeth of Selachians, Actinopterygians and Dipnoans) collected during a recent expedition in Niger. The Maestrichtian localities have yielded a new genus and a new subspecies of Selachian: Igdabatis sigmodon nov. gen., nov. sp. and Lamna biauriculata nigeriana nov. subsp. The locality of Sessao, which has been attributed to the Thanetian by means of the study of the fish, has furnished by screen-washing an interesting fauna wherein six new species are described: Raja Iouisi, Dasyatis sessaoensis, D. sudrei, D. russelli, Hypolophites thaleri and Ceratodus casieri. Comparison of these faunas with contemporary faunas of Africa has brought out a certain endemism in the Iullemmeden Basin during the late Cretaceous and the early Tertiary. 

Of the ten cricetid species from the Oligocene of Western Europe, attributed until now to the genus Eucricetodon, only four prove to be utilizable - E. atavus, E. huberi, E. praecursor, E. collatum - to which it is possible to add two forms newly described: E. huerzeleri and E. quercyi. The evolullon of the genus Pseudocricetodon is also the subject of new observations. The study of the dental morphology allows us to distinguish in these two genera three lineages beginning in the middle Oligocene:

- Lineage P. montalbenensis-P. thaleri (from Montalban to Goderet), of small size, without
increase in size.
- LineageE. atavus-E. infralactorensis (from Hoogbulsel to Estrepouy), of middle size, with a regular increase in size.
- Lineage E. huerzeleri-E. haslachense (level of Montalban to that of Estrepouy), of large size, with an increase in size.

Two other forms are equally represented in these loealitles: Heterocricetodon aff. helbengi and Melissiodon quercyi. It has been possible to attribute a precise age (zone of "Cournon") to the last species, which has been defined by Schaub (l925) from material in the Old Quercy collections. The genera Pseudocricetodon, Eucricetodon, ? "Cricetodon", Leydimys, Eumys, differentiated at the beginning of the middle Oligocene in Europe, Asia and North America, seem to derive from a common ancestral group. The place of origin of this group could be situated in Asia. 

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.   

Evidence of 350 Lower Cretaceous Dinosaur footprints is pointed out in El Bayadh area. Their preliminary study allow to distinguish four trackway assemblages which reveal vertebrate bipedal presence forms of tri-and tetradactylous Dinosauroïds (Assemblages 1-3) and quadrupidal Sauropoïd (Assemblage 4).

The analysis of their footprint biometric features will attribute the quadrupidal Sauropoïd form to Brontopodus ichnogenus which is weIl known in the Jurassic and Cretaceous periods. In retum and despite their age, the dinosauroïd forms were approached, temporarily, to Grallator and Eubrontes types.

The occurrence of the dinosaur traces (Theropoïd and Sauropoïd) constitutes, in the Lower Cretaceous, an important first step of the knowlege of the marshy Reptilian fauna which takes over, from the begining of the Secondary Era, a wide paleogeographie area on the Southem Tethyan margin. 

Tetraceratops insignis is known from a single, crushed skull from the Lower Permian of Texas. Its unique proportions and osteological details gained central meaning in the question of the origins of Therapsida since this early synapsid has been determined as the oldest and less derived therapsid. Apart from Tetraceratops, the ‘mammal-like’ Therapsida and their sister, the pelycosaur-grade Sphenacodontidae, are separated by one of the longest ghost lineages in tetrapod fossil record. However, the minor, though well justified critique faced insistent publication regarding the therapsid hypothesis. A carefull re-evaluation of the holotypic skull reveals that therapsid traits cannot be supported, including a rejection of the formerly supposed adductor shelf in the temporal fenestra. Increased understanding of ‘pelycosaur’ character variation underlines a haptodontine-grade or, less likely, sphenacodontid position for Tetraceratops. 

This paper describes the first hexanchid tooth from the Tithonian (Late Jurassic) of New Zealand. For the moment, this tooth represents the earliest representative of the fossil genus Notidanodon in the world and one of the most ancient neoselachians in the Southern Hemisphere. Despite the perfect state of preservation of the unique tooth, the species is left in open nomenclature, pending the discovery of additional specimens. Few nominal species have been assigned to the genus Notidanodon. Four from Cretaceous deposits: N. antarcti Grande & Chatterjee, 1987, Notidanodon dentatus (Woodward, 1886), Notidanodon lanceolatus (Woodward, 1886), Notidanodon pectinatus (Agassiz, 1843) and only two from Paleocene: Notidanodon brotzeni Siverson, 1995, and Notidanodon loozi (Vincent, 1876). Considering the important morphological variations observed between some of these species, it seems obvious that the genus Notidanodon is not monophyletic and will need a revision in the future. 
  

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.





  

The two species of Sandgrouse from Quercy, Pterocles validus MILNE-EDWARDS and P. larvatus MILNE-EDWARDS, are ascribed to the genus Archaeoganga MOURER-CHAUVIRÉ which includes a third, very large species, A. pinguis. The Sandgrouse of Saint-Gérand-le-Puy, Pterocles sepultus MILNE-EDWARDS, is ascribed to a new genus, Leptoganga. This form appears in the Upper Oligocene of Quercy, in Pech Desse and Pech du Fraysse localities, and is still present in the Lower Miocene of Saint-Gérand-le-Puy and Paulhiac. Recent Sandgrouse live in semidesert or desert areas. The indications provided by mammal and bird faunas in the localities where sandgrouse were found, confirm that the paleoenvironment was open and arid. The morphological study of these fossils indicates that, in the Upper Eocene, the Pteroclidae were already completely individualized with respect to Charadriiformes. 

The study of three new populations of Plesiosminthuspromyarion from the "phosphorites du Quercy" and of material from "Auvergne" brings new data on european oligocene Dipodidae. They appear in Western Europe at the beginning of late Oligocene. Evolutionary trends of the group are drawn and particularly the emergence of morphotypes announcing P. schaubi, from the Coderet level, is revealed among the most recent populations of P. promyarion. Differences are attempted to be drawn between the three species : P. promyarion, P. myarion and P. schaubi by restudying the type-population of P. myarion from the aquitanian deposits of Chavroches (Allier) in comparison with two other populations from the same age and the same region. Relationships between early european, american and asiatic Dipodidae are discussed. 

Pleistocene bird fossils have been studied from nine localities on Crete. Part of this material was described earlier by the author (Weesie, 1982) and will not be treated here in extenso, the results will be incorporated. More than one third of the over 10,000 fossil bird bones available could be identified ; they were found to represent at least 65 bird species. The following species of the Pleistocene Cretan avifauna are new to the fauna of Crete : Branta ruficollis, Haliaeetus albicilla, Gyps melitensis, Aquila chrysaetos simurgh n. ssp., Ketupa zeylomensis, Aegolius funereus, Dendrocopos leucotos, Zoothera dauma, Turdus iliacus and Pyrrhula pyrrhula. The Pleistocene Cretan avifauna differs less from comparable mainland avifaunas than (fossil) avifaunas from oceanic islands do. Still, the Pleistocene Cretan avifauna has two qualities that are characteristic of island avifaunas : the almost complete absence of a group of birds (the Galliformes) and the presence of two endemic (sub)species : the giant eagle Aquila chrysaetos simurgh n. ssp. and the long-legged owl Athene cretensis (Weesie, 1982). The new subspecies is described in the present study.
These endemic birds of prey were found in association with their supposedly principal prey species (now extinct as well) : endemic mice for the owl and endemic deer for the eagle. Endemic mammals have been found in association with endemic birds of prey on many islands, not only in the Mediterranean. There is evidence that the size of endemic birds of prey becomes optimally adapted to their feeding on certain endemic mammals, especially rodents. Another characteristic of the Pleistocene Cretan avifauna is the great number of species of birds of prey. This appears to be a common characteristic of fossil avifaunas from caves on Mediterranean islands as well as from caves on the European mainland. However, we think that ecological conditions on Pleistocene Crete (especially the abundant presence of mice) helped to account for the high representation of birds of prey. Furthemore, the fossil avifauna enables us to draw some conclusions about the climate and vegetation on Pleistocene Crete : it is concluded that the climate was cooler than today and that Crete was largely covered with forests. Finally, the reasons for the extinction or disappearance from Crete of some bird species of the Pleistocene Cretan avifauna are discussed. 

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. 

As a result of the enrichment of eotheriodont material by one of us (P.K.T.), these specimens (essentially Biarmosuchur and Eotitanosuchur) are reexamined and refigured. A reevaluation of their particularities supports the distinction of two families, for which new diagnoses are proposed. This leads us to discuss the affinities of these families, with respect to the sphenacodonts on one hand, and to the South African primitive theriodonts on the other (gorgonopsids and ictidorhinids). This study contains inherent paleogeographic consequences which are considered in conclusion. 

The anatomy of Norselaspis glacialis n.g., n.sp., a primitive kiaeraspidian from the Lower Devonian of Spitsbergen, is described on the basis of spécimens studied by grinding sections or prepared with dilute formic acid. This study yielded some new anatomical details, including the presence of a canal prolonging posteromedially the canal alloted to the facial nerve by Stensiö. This posterior prolongation of the « facial canal ›› into the posterolateral part of the labyrinth cavity is consistent with the hypothesis put forward by Allis, Lindström, Jefferies and Whiting, that this canal housed the glossopharyngeus nerve. Furthermore, in N. glacialis, the foramen usually referred to as the foramen for the œsophagus opens posteriorly into a cavity in the postbranchial wall, referred to here as the intramural cavity, and which is interpreted as having housed the heart. Consequently, the œsophagus probably accompanied the dorsal aorta through the aortic canal. Finally, the foramen generally interpreted as having transmitted the ventral afferent arterial trunk is here considered as having housed the hepatic vein, which emptied into the venous sinus of the heart. The ventral afferent arterial trunk may thus have passed through the former «œsophageal ›› foramen.
The problem of the position of the dorsal nerves in the Osteostraci is discussed, and it is suggested that the three foremost nerve canals opening into the oralobranchial cavity housed the maxillary ramus of the trigeminus, the facial nerve and the glossopharyngeus nerve respectively. The mandibular ramus of the trigeminus must have accompanied one of the two foremost nerves, but for the moment it is impossible to decide which.
The problem of the nature of the interbranchial crests of the Osteostraci is briefly discussed. Comparison with the branchial apparatus of the Petromyzontida does not support the hypothesis that the interbranchial crests are part of the branchial arches, incorporated into the endoskeletal shield. A different hypothesis is proposed, that the branchial skeleton of the Osteostraci was situated entirely inside the oralobranchial cavity, and was attached to the endoskeletal shield only by the ventromedial processes. The grooves classically allotted to the efferent branchial arteries would thus have housed extrabranchial arteries, branching off from the dorsal aorta, and irrigating the ventral branchial musculature.
A phylogeny and a classification of the kiaeraspidians are proposed. The evolution of this monophyletic group is characterized by, e.g., reduction of cornual processes, shortening of the abdominal division of the shield, subdivision of the lateral fields, and enlargement of the supraoral fossae.
The phylogenetic position of the kiaeraspidians within the Osteostraci remains uncertain. Their sister-group may be either the benneviaspidiens or the thyestidians, or Thyestes alone (in which case they would have to be included within the thyestidians). 

The genus Plagiolophus is documented, almost solely in Western Europe, from the middle Eocene up to the mid Oligocene (MP 12 to MP 25), i.e. more than for 15 MY. Seventeen species are now recorded whose two of them are new, P. ringeadei nov. sp. and P. mamertensis nov. sp. Some anatomical variations and the deflection of certain evolutionary trends justify the distinction of three subgenera, Paloplotherium, Fraasiolophus nov. and Plagiolophus s.s. The genus displays a wide range in size and weight (between 10 and 150 kg). The detailed description of the skull of several species is here given for the first time.
Despite important evolutionary drifts during this long time span, the dentition shows a great structural homogeneity, which renders difficult the determination of fragmentary specimens or isolated teeth. It is characterized by a great heterodonty; premolars are little molarized and present a certain regression through time with paradoxically some progress in the molarization. The hypsodonty increases: the first Plagiolophus are hardly less brachyodont than Propalaeotherium, and the last ones are nearly as hypsodont as Merychippus from the early Miocene. The upper molars change from a wide crown pattern, with an open occlusal surface, lightly oblique transverse lophs and rounded internal cusps, to a narrower pattern, with a frontally constricted occlusal surface and internal lophs aligned parallel to the ectoloph. The M3/3 become always longer.

The dental enamel displays horizontal Schreger-bands with imprecise limits occupying only the middle part of the enamel layer. The dentine is remarkable by its high rate of pericanalicular dentine. The crown cementum, lacking in earlier forms, increases to the point where it fills the occlusal valleys of the
teeth.

The masticatory musculature shows a increasing prominence of the temporal, with probably an important role devoted to the pterygoid muscles in lateral movements related to a two-phase type of chewing.

The evolution of the dentition, of the masticatory musculature and of the repartition of masticatory forces indicate that the Plagiolophus have known different diets through their long evolutionary history; at first browsers they became mixed feeders and finally grazers. Their relatively long neck allowed these animals to reach different vegetal layers. The strength of the nuchal crests also suggests that they were able to have strong backwards movements of the head to pull up their food.

This evolution of diet seems related to the slow degradation of environmental conditions attested during this period in western Europe, with the generalization of more open landscapes, increasing aridity and more marked seasons.

Besides, a remodeling of the face is ontogenetically and along time observed, in relation with the evolution of the masticatory apparatus and especially with that of the mandibular lever arm. The postcanine diastemata become longer in the course of evolution; the free extremities of the nasals are always relatively long which contradicts the hypothesis according to which Paloplotherium may have had a trunk. At last the lineage Fraasiolophus can be distinguished by the presence of a deep malar fossa, probably related to a strong development of the maxillo-labialis superior muscle.

The orbit is always large and tends to increase in size, which indicates a good development of the vision and its increasing role in the life relations. A peculiar type of epitympanic sinus could have been used as a resonance chamber insuring a certain amplification of sounds before their transmission to the eardrum. The endocranial cast reveals a relatively large brain with an advanced degree of gyrencephaly. Beside the role eventually played in food research and social relations, these neurophysiological abilities, also related to an advance in cursorial fitness, could have contributed to the survival of these animals facing the predation pressure of the first fissipede carnivores and the competition with new immigrant herbivores after the "Grande Coupure".
On the basis of some shared apomorphies with the Pachynolophinae, which prevent from considering the latter as Equidae (molarization of the premolars, reduction of the premaxilla dorsal apophysis, peculiar epitympanic sinus, splitting of the jugular process), the hypothesis of an autochthonous origin of Plagiolophus issued from a form near Propalaeotherium, is once again proposed and discussed. Finally, intra-generic relationships are taken into consideration. 

This paper proposes a systematic revision of the Oligocene Mongolian Ctenodactylidae, on the basis of abundant material obtained by screen/washing operations in stratified localities of the Ulantatal area (Inner Mongolia) (UTL1, 2, 3, 4, 5, 7, 6 & 8). A Chinese-German team has collected several thousands of isolated rodent teeth, and a number of fragmentary jaws. A new genus is identified (Alashania nov. gen. tengkoliensis nov. sp.), and eight former species are reevaluated, Karakoromys decessus, Tataromys sigmodon, T. minor, T. plicidens, Yindirtemys ulantatalensis, Y. bohlini, Y. deflexus, with several synonymies. A new Yindirtemys species is described: Y. shevyrevae nov. sp. and another one close to that: Y. aff. shevyrevae nov. sp. Four new species, which are rare in the localities, remain in open nomenclature because they are not well-represented. Yindirtemys differs from the other genera by the permanence of crescentic structures, while the other genera show a general reduction of the trigonoid area (= anterior valley). We define a range of size variation for each well documented population. Although the dental morphology shows a wide range of variation, given that transitional morphologies occur in a single locality, it is possible to provide a clear definition for most species. We show that dental patterns of the different genera can be derived from the pattern of Karakoromys. As a number of Tataromyinae have been determined in several localities from China, Kazakhstan and Mongolia, usually on the basis of scarce material, or surface collections, the present study would be used to re-evaluate their attribution inasmuch as the taxa are now placed in the Oligocene stratigraphy. The diversity of sizes and forms reflects the adaptive radiation of the family during the Oligocene, within a forested environment where the vegetation was probably abundant. 

The hipparions from the Dytiko localities of the lower Axios valley (Macedonia, Greece) are studied. The material comes from three localities Dytiko-l, 2, 3 (DTK, DIT, DKO), which are situated near the village of Dytiko, about 60 km northwest to Thessaloniki. Three species have been determined, the medium-sized H. mediterraneum, the small-sized H. matthewi and the very small-sized H. periafricanum. The determined Hipparion species, their morphological characters and their comparison with the other Axios valley material indicate a Late Turolian age for the Dytiko localities.