The Laprade fauna is chronologically situated between those from Egerkingen and Lissieu and consequently, is close to the MP 14 reference-level of the European mammalian biochronological scale (Symposium of Mainz, 1987).
This new fauna is presently the oldest known in the Quercy phosphorites, formerly the Le Bretou fauna (MP 16) was considered as the oldest one. The Laprade fauna includes 21 species which belong in 7 mammalian orders (Marsupialia: Amphiperatherium bastbergense, Amphiperatherium goethei; Apatotheria: Heterohyus (Gervaisyus) pygmaeus nov. subgen., nov. sp.; Lipotyphla: Saturninia cf. mamertensis, Saturninia cf. intermedia; Chiroptera: Vespertiliavus lapradei nov. sp.; Rodentia: Protadelomys cf. lugdunensis, Elfomys nov. sp.; Primates: Nannopithex cf. filholi, cf. Pseudoloris or Pivetonia; Perissodactyla: ?Palaeotherium ?castrense, small-sized Palaeotherium sp., Plagiolophus sp., Anchilophus sp.; Artiodactyla: Dichobune cf. robertiana, Mouillacitherium cartieri, Tapirulus cf. depereti, Mixtotherium priscum, Pseudamphimeryx schlosseri, and Artiodactyla indet). Sixteen of these species are mentioned for the first time from the Quercy faunas.
The recognition of a new apatemyid, Heterohyus (Gervaisyus) pygmaeus nov. subgen., nov. sp., attests to an early origin of a lineage known in the Late Eocene. The emballonurid bat Vespertiliavus lapradei nov. sp. is presently the earliest record of this genus and family. This Auversian fauna leads to discuss the age of taxa showing archaic features. These taxa were defined on specimens collected in the Quercy during the last century and have never been found in the Quercy localities recently investigated. This fauna bears also evidence of a karstic filling episode older than those previously dated by fossils in the Quercy Jurassic. 

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. 

A lot of isolated teeth of a pteropodid fruit bat has been recently found within an assemblage of micromammals recovered from a karstic fissure filling named Lo Fournas-Il near the locality of Baixas (Pyrénées-Orientales, France). The fauna is Middle Miocene Serravallian age. The fossil fruit bat appears morphologically close to Rousettus; its size is that of a recent medium-sized fruit bat. While the fruit bats are very poorly known as fossils, this discovery shows that one of their recent types of dentitions was perfectly established by Middle Miocene times, and supports the presumed long geologic story of the suborder. One of the major invasions of the Old World fruit bats, supposed originated from SE Asia, reached up to Europe. A suborder unit is added to the miocene fauna of this continent. 

The Mont-Hélène's fauna [Pyrénées-Orientales, France], includes 15 species of rodents with a new one, Occitanomys montheleni n. sp. among the 9 species of the Murids which are listed. The uncommon Cricetid, Blancomys neglectus, is well represented in the fauna. Peculiarities of the population referred to Slephanomys cf. donnezaniare discussed. The locality a fissure filling may be the oldest one of Tabianian age known in Southern France. The diversity of the Murids gives evidence of a subtropical climate and of a diversified environment which may be linked to the spreading of the coastal plain following the filling up of the Roussillon Neogene Basin. 

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.





  

The late Cretaceous Auca Mahuevo nesting site (Neuquén Province, Argentina) has produced a large number of sauropod eggs, many of them containing the remains of embryos. Research at this site has generated important information about the development of the embryos, the morphology and eggshell microstructure of the eggs, and the reproductive behavior of sauropod dinosaurs. Cranial features present in the embryos have allowed their identification as those of titanosaurian sauropods. Differences in the texture of the sediments that contain some of the egg-clutches have illuminated their nest architecture. Microstructural studies of eggshells have expanded our knowledge of their variability and the incidence of pathologies within a reproductive titanosaurian population. Maps showing the spatial distribution of eggs and clutches. the stratigraphic distribution of the egg-beds, and the sedimentological context in which they are contained, have provided the basis for several inferences about the nesting behavior of these dinosaurs. 

The Causses" is a near 3400 km2 large plateau located in the south of France. Here the first dinosaur footprints where found in 1935. After this, this area has yielded an ever-increasing number of ichnites now in excess of 500 specimens. These latter, 15 to 50 cm long, tridactyl or tetradactyl footprints of generally biped animals, were discovered at the surface of Hettangian to lower Sinemurian dolomite layers within 4 distinct stratigraphic units. The 35 sites bearing ichnites are located on the plateau margin. For the first time, morphologic characters studied through descriptive statistic methods with the usual parameters and classical Student and Snédecor tests, allowed us, to divide the whole set of biped traces into 6 ichnospecies. Their definitions are further constrained by multivariate statistical results using Principal Component Analysis (PCA), Factor Analysis of correspondances (FAC) and Discriminant Analysis (DA). All have confirmed the morphologic observations. So that now, the following taxa are identified : Grallator variabilis, G. lescurei, G. sauclierensis, G. minusculus, Eubrontes giganteus, Dilophosauripus williamsi, cf. Moraesichnium, Orníthopus fabrei nov ichnosp. The more immediately visible differences relate to the interdigital II-IV divarication and the digit length ratio. To this panel, we must add Batrachopus deweyi and shapes suggesting Trisauropodichnus and/or Anomoepus. Among all ichnite associations described in the lower Liasic, the New England assemblage presents the most affinities with ours. It shows the ichnotaxa Grallator, Dilophosauripus, Eubrontes, Batrachopus without forgetting Ornithopus fabrei nov. ichnosp. which is close to Ornithopus gallinaceus from the Massachusetts and Connecticut basins. On comparing the present early Jurassic ichnofauna of the Causses with the ones of the Middle and Upper Triassic formations of the eastem border of the Massif Central (France), it appears that tridactyl footprints become more and more numerous and large from Triassic to Early Jurassic. In the Causses, these latest are prevalent but in Quercy (France), Poland, Italy, USA, they are also associated with Omithopoda, Thyreophora and Sauropoda ichnites. Footprint areas considered here were generaly under an arid climate. Animals that passed by were heavy and bulky possible Megalosaur trackmakers, and lighter and slender Coelophysids or Ceratosaurs. For all, these areas were pathways as the orientations of the trackways seem point out. The directions followed by these reptiles were without any important variation during the Hettango-Sinemurian stages. These areas were also used from time to time by Crocodilomorpha and may be tetradactyl (I-IV) bipedal avian Theropods. However, the number of such trackways in sites, sometimes substantial, should not lead us to overestimate the trackmakers populations. These last were probably relatively moderately abondant in this inter-supratidal swamp environment. In the Causses, ichnites are connected with former algo-laminated deposits (Algal mats) which were rapidly hardened by means of calcitisation of cyanobacteria. The result has been a moderate depth of footprints; autopodia disturbing only a few cm of the carbonate substrate. Other fossils have been discovered : invertebrates with thin bivalve and gastropod shells, crustaceans tests and plants. These latter suggest the existence of paleomangroves like environments but also continental vegetation periodically overruning the swamp environment during regression/transgression cycles. At these times, wooded parts of it, could become protecting, feeding, resting and nesting places.

  

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. 

The new fauna of Collet Redon (Bouches-du-Rhône, France) is dated by three rodents: Megacricetodon aff. bavaricus, Democricetodon affinis mutilus and Peridyromys cf. hamadryas. They correlate this locality with Oggenhof and Ohningen in Bavaria (Western Germany). As the radiometric age of Ohningen is estimated between 14 and 13 M.Y., these three localities are of Serravallian age. This datation brings a complete readjusment of the stratigraphy of the section of Collet Redon formerly described by Collot and Combaluzier. The marine deposits with underly the continental formation with the mammal fauna, are Burdigalian. The angular unconformity between the marine and the continental deposits gives evidence of an episode of emersion on the margin of a sedimentary basin, with deformation and erosion. Owing to the newly discovered fauna, this geodynamical event is clearly settled within the regional geographical and chronological context. Lacustrine and continental deposits of such an age were up to now unsuspected in this area. 

Le Quercy est aujourd 'hui un vaste plateau calcaire, parcouru par un réseau karstique actif, profondément
entaillé par des vallées aux falaises abruptes, comme celles du Lot ou du Célé. Sur un sol peu épais domine la forêt de chênes, accompagnés de cornouillers, érables, genévriers. La faune est pauvre, peu diverse, et les nombreux chasseurs se satisfont de gibier d'élevage ...

Il y a trente cinq millions d'années environ, le paysage était bien différent. La période de l'Eocène supérieur, qui s'achevait, avait été chaude et humide, si l'on se réfère à la fois aux paléotempératures (calculées à partir de sédiments marins extra·européens) et aux restes fossilisés de végétaux typiquement tropicaux. 

    Two families are reviewed (Hyracodontidae and Rhinocerotidae) from the Paleogene of Mongolia. The following taxa are described from the former family:  Triplopus? mergenensis sp. nov., Prohyracodon meridionale CHOW, Prohyracodon? parvus sp. nov., Forstercooperia ergiliinensis GABUNIA & DASHZEVEG, Ardynia praecox MATTHEW & GRANGER, A. mongoliensis (BELIAYEVA), Ardynia sp., Urtinotheríum sp. (or Indricotherium) and Armania asiana GABUNIA & DASHZEVEG. All of them are from the Paleogene of the eastem Gobi Desert The hyracodontid Pataecops parvus RADINSKY is described from a new locality, Menkhen-Teg (Middle Eocene), in the Valley of Lakes.
    In the second family, a new species of Ronzotherium, R. orientale sp. nov. has been recognized from the Oligocene of Ergilin Dzo and Khoer Dzan of the eastem Gobi Desert. Ronzotherium sp. and Allacerops sp. have been described from the Oligocene of Khoer Dzan. The genus Ronzotherium, formerly known from the Oligocene of eastem Europe, has been reliably established in the Oligocene fauna of Mongolia. The genus Symphysorrachis BELIAYEVA, 1954, previously believed to be a junior synonym of Ronzotherium, is here resurrected.
    Descriptions are provided for key localities in the Eocene and Oligocene of Mongolia and the adjacent territories of northern China, containing fossil Hyracodontidae and Rhinocerotidae. In addition, the question of the Eocene-Oligocene boundary in these Mongolian and Chinese sections is discussed. Paleontological evidence has been used to correlate Eocene-Oligocene boundary layers in the eastem Gobi Desert (Mongolia) and Inner Mongolia (China). A brief discussion is provided on the phylogenetic affinities of the genera within the family Hyracodontidae. 

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. 

Valverde's cenogram method is used to analyse mammalian communities from the late Eocene to late Oligocene of southern France, mainly from the "Phosphorites du Quercy". Cenogram analysis involves plotting the size of each component species in a fauna on a semilog diagram in rank order, permitting fossil faunas to be compared with Recent ones. The configurations of Recent communities serve as models for establishing the general environmental characteristics of fossil mammalian faunas. This method of analysis applied to faunal sequence can reveal major and sudden ecological perturbations. The paleobiogeographical event (i.e. the mammalian immigration wave) at the Eocene-Oligocene Boundary in western Europe, known as the «Grande Coupure", is here shown to represent a drastic and sudden ecological change: late Eocene tropical environments in Europe deteriorated rapidly turning to subdesert or desert environments al the beginning of the Oligoccne. 

Previous chronological arrangements of South American Quaternary land mammal faunas are appraised on the basis of current geological and paleontological data. Three South American late Pliocene-Pleistocene land mammal ages are conventionally recognized, from oldest to youngest, the Uquian, Ensenadan, and Lujanian ; all are defined on Argentine faunas.

     The Uquian is based fundamentally and historically on the fauna from the Uquía Formation in Jujuy Province, northwestern Argentina. Important known formations in Argentina yielding Uquian Age faunas include the sub-surface Puelche Formation (or Puelchense) near the city of Buenos Aires, and the Barranca de Los Lobos and Vorohué Formations between Mar del Plata and Miramar, Buenos Aires Province. A tentative subdivision is propos-ed for the Uquian into three subages based on knowledge of the Mar del Plata-Miramar sequence, from oldest to youngest, the Barrancalobian, Vorohuean, and Sanandresian. In Argentina the Uquian is presently marked by the first known record of Scelidodon, Hydrochoeropsis, Ctenomys, Canidae, Ursidae, Gomphotheriidae, Equidae, Tapiridae, Camelidae, Cervidae, and the last known record of Thylatheridium, Thylophorops, Dankomys, Eumysops, Pithanotomys, Eucoelophorus, Hegetotheriidae, Sparassocynidae, and Microtragulidae.

The Ensenadan Age is based on the fauna from the Ensenada Formation near the city of Ensenada, Buenos Aires Province. In Argentina the Ensenadan is marked by the first known record of Lomaphorus, Neothoracophorus, Plaxhaplous, Cavia, Lyncodon, Lutra, Galera, Smilodon, Dicotyles, Lama, Vicugna, the last known record of Orthomyctera, and the only known record of Brachynasua.

     Typícal beds of late Lujanian Age in Argentina consist of fluvial deposits occupying stream channels, and shallow basins, often incised into beds of early Lujanian (i.e. Bonaerian of early workers) and Ensenadan Age. The Lujanian Age is based on a fauna from beds along the Rio Luján, about 65 km west of the city of Buenos Aires, Buenos Aires Province. The Lujanian in Argentina is marked by the first record of Equus, Chlamyphorus, and Holochilus, and the last record of Megatherioidea, Glyptodontoidea, Arctodus (=Arctotherium), Smilodon, Litopterna, Notoungulata, Proboscidea, Equidae, Morenelaphus, and Palaeolama.

   These land mammal ages are often difficult to recognize in other South American countries. The compositions of South American Pleistocene faunas vary with the environment. Some taxa were widely distributed in fossil deposits throughout the continent, but their occurrences need not reflect synchroneity. This is a result of changing climates and habitats in time. Consequently, proposed intracontinental correlations need confirmation based on magnetostratigraphy and a radioisotope time scale. Paleontologic characterizations of these land mammal ages (i.e. first and last record, and guide fossils) are useful for much of Argentina, but extensions to most of the other parts of South America are at best tenuous.

The majority of known non-Argentine Pleistocene faunas are believed to be Lujanian in age. Possible non Argentine early Pleistocene (Uquian) faunas include Ayo Ayo and Anzaldo in Bolivia, and Cocha Verde in southern Columbia. A possible middle Pleistocene (Ensenadan or early Lujanian) fauna is the Chichense of Ecuador. Paleomagnetic and radioisotopic date (MacFadden et al., 1983) clearly indicate that the greater part of the Tarija fauna (Bolivia) is Ensenadan in age.

  The end of the Pleistocene and beginning of the Holocene in South America is marked by extinction of nearly all large mammalian herbivores and their specialized large predators. Radiocarbon age determinations suggest that large scale extinctions of megafauna occurred between 15,000 and 8,000 yrs. B.P. (years before present). 

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.
  

Les Pistes de Vertébrés du Stormberg Supérieur ("Trias terminal à Rhétien"), ou Quthingien

Si les zones du Stormberg inférieur se sont révélées contenir de nombreuses traces, surtout dans les faciès dits "Molteno moyen et supérieur", représentant apparemment la base du Keuper, il est frappant de voir pratiquement l'ensemble de cette grosse faune "Molteno" disparaître avec la fin de cette période, que nous avons appelée le "Maphutsengien".

Dès les premières zones du Stormberg supérieur, que nous nommons le"Quthingien" la zoocénose et la phytocénose, en même temps que les données d'ensemble manifestées par l'environnement, sont modifiées. Nous ne verrons plus guère de dépôts marécageux à flore riche et variée, parfois même luxuriante. Les fougères elles-mêmes ont disparu. Elles sont remplacées par de maigres plantes, aux feuilles très souvent filiformes qui paraissent témoigner d'un climat continental. Le sol est devenu de plus en plus rouge, avec des variations latérales beaucoup plus accusées. Les fleuves amenant des galets des monts du "Grand Sud" ont tari. La faune va en subir les conséquences. Certaines des espèces se révèleront sautillantes ou coureuses, pour un grand nombre plus légères et pour la quasi-totalité d'apparence carnivore ou entomophages, les phytophages devant se contenter d'un régime ingrat,difficile ou à tout le moins irrégulier,les dépôts le montrent.

C'est dans ces conditions que s'inaugure notre Etage nouveau,quelque peu discordant sur les zones A/5, A/6 ou A/7 du Stormberg inférieur (Maphutsengien). Le Stormberg supérieur (ou Quthingien) commence avec le paléopaysage remarquable dit de Moyeni, que nous allons maintenant étudier, typologiquement, avec ses homologues du même âge. Quelques 38 types d'animaux tous nouveaux vont défiler à nos yeux lors de la zone de base de cet Etage, ou zone B/1.

L'on nous avait proposé d'intituler ce Ile Tome de la série : "La grande Dalle de Moyeni et ses homologues. Paléo-spectacles, scènes et paysages animaux au Lesotho à l'approche du Trias finissant". Nous avons préféré garder le sous-titre plus haut, peut-être plus prosaïque.

Un llle Tome est en préparation : "Les développements ultérieurs et terminaux de la faune du Gondwana".

  

Redécouvertes en 1968 dans le bois du Bretou, les deux poches à phosphorites qui ont livré cette faune sont parmi les plus méridionales du plateau quercynois. De faible profondeur, huit mètres, les cavités sont de petite taille et font partie d'un complexe de sept fissures réparties dans les bois du Bretou. Les autres trous n'ont livré aucun fossile, soit parce qu'ils avaient été intégralement vidés de leur sédiment, soit que des venues d'eau en interdisent l'accès.
Dans les deux poches fossilifères, proches de quelques mètres, le sédiment que l'on rencontre en placage sur les parois présente un aspect particulier. Il s'agit en effet d'une brèche très indurée, où la calcite domine, et contenant assez peu de sédiments sidérolithiques. Les ossements contenus dans cette brèche sont assez nombreux, très souvent brisés. Les dents isolées y sont relativement rares. Exploitées séparément, les deux poches ont livré des faunes identiques, aucune différence chronologique ne pouvant être décelée entre elles. Les formes rencontrées dans une seule des cavités sont rares mais la poche la plus riche est celle dénommée Le Bretou 2.
L'intérêt de la faune récoltée réside dans son ancienneté. On sait en effet que H.G. Stehlin avait signalé dans les collections du Quercy des restes attribuables à Lophiodon lautricense, mais la provenance exacte de ces fossiles reste inconnue. Au Bretou si nous n'avons pas rencontré à ce jour le Lophiodon, il n'en demeure pas moins que l'âge de la faune récoltée doit être proche de celui de la faune de Robiac. 

A short review of the genera and species of snakes from the Quercy's phosphorites described by former authors is followed by the study of specimens recently collected. Most of these latter specimens belong to the Aniliidae, Boidae, Colubridae and to the Scolecophidia; the precise systematic position of some of them is not defined yet. The following genera and species are described: Eonilius europae nov. gen. and nov. sp., Platyspondylia lepta nov. gen. and nov. sp., Coluber cadurci nov. sp. and Dunnophis cadurcensis nov.  sp.; a species described by de Rochebrune (Palaeopython filholi) is revised. One of the most important conclusions of this study is that the Colubridae appear as early as the Upper Eocene. The stratigraphic repartition of these snakes shows that the rich Upper Eocene fauna is followed by the very reduced fauna of the Lower Oligocene; then the oligocene «Grande Coupure» corresponds to a very important faunistic renewal. The fauna remains poor and little diversified during the beginning of the Middle Oligocene after which there is a new faunistic explosion. 

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. 

Remarkable association of a small infraorbital foramen, similar to that in recent Heterocephalus, and of a strong muscular print on the dorsal anterior part of the zygomatic plate and on the premaxillary. Several anatomical structures to be compared with those of Heterocephalus suggest relationships with this genus. Richardus supports the ancestrality of the hystricomorph character in the bathyergids