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".

  

A new hypothesis for the phylogenetic relationships of recent anomalurids and graphiurids is proposed, based on information from evolutionary lineages of Paleogene European rodents, particularly Gliridae, and Eocene Algerian Zegdoumyidae. Differences in first occurrences, in paleogeography, and in infraorbital structure in glirids (protrogomorphy and pseudomyomorphy) and graphiurids (hystricomorphy) separate Graphiuridae from Gliridae (Graphiurinae is here raised to family rank). Similar considerations, and dental morphology, suggest that Anomaluridae (appearing in the late Eocene) and Graphiuridae (appearing in the Pliocene) are related to early Eocene Zegdoumyidae. 

Sainte-Néboule has yielded only 4 species of Rodents. But the Theridomyids (Blainvillimys rotundidens and Patriotheridomys altus) are very significative of the age of the locality: Ste-Néboule is lower than the marker level of Escamps 

The artiodactyls (Mammalia) from the latest Ypresian locality of Prémontré from the Paris Basin (niveau repère MP 10 in the lower Eocene of the Paris Basin) are described in this paper. Three species have been identified: 1) Diacodexis cf. varleti SUDRE et al., 1983; 2) a new species of Eurodexis ERFURT & SUDRE (E. russelli nov. sp.) defined after the revision of the species Messelobunodon? ceciliensis from the Lutetian beds of Geiseltal (Germany); and 3) Eurodexeinae indet., a probable ancestor of another form from the Geiseltal which was previously recorded as Homacodon? sp. (Erfurt 1993) and now named Parahexacodus germanicus. The two later forms are referred to the new subfamily Eurodexeinae (Erfurt & Sudre 1996). The analysis of these forms as weIl as comparative studies have led us to reconsider our previous conclusions regarding the content of the species Protodichobune oweni LEMOINE 1878 and some aspects of Ypresian diacodexid evolution. One can postulate that the divergence of E. russelli nov. sp. occurred during the first radiation of these primitive artiodactyls. Some other stem form with bunodont teeth such as Protodichobune and Aumelasia have also differentiated from Diacodexis. Like Eurodexis, these two genera persist during the middle Eocene. The absence of Protodichobune and Aumelasia at Prémontré is probably due to particular ecological conditions. 

Sirenians are among the common marine fossil remains found in the Faluns deposits of western France. We describe new material of a Dugongidae sirenian from the middle Miocene Savignean facies of La Morfassière quarry (Indre-et-Loire, northwestern France) that includes a well preserved and almost complete skull associated with its mandible, several vertebrae and ribs. The cranial remains exhibit features that allow to attribute them to Metaxytherium medium, a species recorded from the middle and early late Miocene of European and Mediterranean coasts. The discovery of an associated skull and mandible of this species is unusual in this area and deserves to be reported, mostly because its preservation contributes to a better knowledge of the variable structure of its skull and teeth anatomy. For the first time the body size and weight of M. medium can be assessed through regression equations based on skull measurements. The particularly good condition of the material can be explained by the calm and deep marine environment in which it was deposited.
  

Extensive sampling of the Siwalik deposits of the Potwar Plateau of northem Pakistan and from the Zinda Pir dome of central Pakistan has produced a fossil record of Miocene ctenodactylids that can be correlated with the paleomagnetic time scale. The early Miocene Prosayimys flynni (n. gen., n. sp.) is recognized as the first ctenodactylid in the Indian subcontinent. Prosayimys is ancestral to Sayimys. From the late early Miocene to the early late Miocene, there is an anagenetic succession of three species of Sayimys: S. cf. S. intermedius, S. sivalensis, and S. chinjiensis (n. sp.). Sayimys chinjiensis gave rise to the late late Miocene S. perplexus. A second lineage is represented by Sayimys minor, S. sp. A, and S. sp. B. 

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. 

In recent years, the techniques of chemical preparing have permitted a rich paleontologic material to be obtained from the phosporitic sediment of Bouzigues (Hérault, France). The fauna of this locality is comprised of quite varied microvertebrates, amphibians, reptiles, birds, mammals. Twenty five species of the latter, belonging to seven orders, are today known from the site. Among them, the numerous rodents have allowed L. Thaler to chronologically situate this fauna in the Zone of Laugnac (<< late Aquitanian ›> of some authors).
The chiropterans are, with the rodents, the best represented of the locality's mammals. Three families comprise the bat fauna, with nearly complete dominance by one of them (Hippoxideridae) over the two others (Megadermatidae and Vespertílionidae)
Six forms are described, of which three are new species and one a new sub-genus.
Megaderma braillomi n. sp., an animal of rather large size, shows like the Miocene megaderms several evolved dental characters, translating the adaptation of these animals to a partially carnivorous regime. The Bouzigues species seems, however, to represent a particular lineage.
Hipposideros (Brachipposideros n. subgen.) dechaseauxi n. sp. and Hípposideros (Brachipposideros) cf. collongenris Depéret, small sized forms, belong to a group rather well represented in the late Oligocene and early Miocene of Europe, and not distinguished until now within the genus Hipposideros.
Hipposideros (Pseudorhinolophus) bouziguensis n. sp., is the most abundant mammal in the locality and, occuring at the Oligocene-Miocene limit, the last representative known of the subgenus Pseudorhínolophus, common in Europe from the middle Eocene.
However, beyond Neogene and Quaternary times, certain among the numerous living species of Hipposideros are close to Pseudorhinolophus and others to Brachipposíderos. 'This fact would in the future justify a global revision of the genus, on the basis of comparative anatomy of the squeleton and of the teeth.
The bat fauna of Bouzigues is completed by two small Vespertilionidae, rare forms, Myoris sp. I and sp. II.
  

We present here a new updated counting of the perissodactyls of Robiac, the type locality of the MP 16 level of the biochronological scale of paleogene mammals and that of the Robiacian stage of Eocene Land Mammals Ages in Western Europe.
The outcrop of Robiac consists actually of two 500m apart loci, Robiac-Nord and Robiac-Sud, considered of the same age according to the current discriminating power, and is dated from -38,7 MA after the last faunal, magnetostratigraphic and climatic calibrations.
It has yielded a very abundant and rich of 21 taxa perissodactyl fauna, topped by the giant Lophiodon lautricense, last representative of the family Lophiodontidae, of which it is the last proved deposit. The Palaeotheriidae are much diversified with 5 genera and 9 species of "Pachynolophinae", 3 genera and 10 species of Palaeotheriinae. Nine taxa have been defined from Robiac: Chasmotherium depereti n. sp., Palaeotherium castrense robiacense Franzen, 1968, the genus Leptolophus Remy, 1965 with the species L. stehlini Remy, 1965 and L. magnus Remy, 1998, Anchilophus (Paranchilophus) jeanteti Remy, 2012, Metanchilophus chaubeti Remy, 2012, Lophiotherium robiacense Depéret, 1917 and Pachynolophus gaytei n. sp.
The faunal Robiac cenogram with the associated flora testify to a hot, wet and forestal environment, likely corresponding to a short warming climatic phase; the broken up fossil bones should have been carried away and then gathered in swamp areas along the banks of a meandering river.
The swarm of mammals of Robiac, the richest of contemporaneous deposits, has been followed by a drastic drop in perissodactyl diversity at the MP 17A level; a crisis which could have originated in a renewal of the global Eocene cooling. Fons 4, the type-locality of this level, is largely scarcer in perissodactyls and its cenogram testifies to a less diversified fauna, with on the whole smaller species, that likely means a cooler and drier climatic environment; a new perissodactyl diversification occurred but later.
  

The present study describes in detail the anterior limb osteology of a molossid chiropteran of the genus Tadarida, from Céreste, a Stampian locality in the Apt-Forcalquier Oligocene basin already known for its fishes, plants and insects.

A comparision with the Miocene forms T. srehlini from Saint-Géraud localities and T. sp. from Württemberg, also with the recent forms T. teniotis and Eumops perotis, does not show any clear morphological differences between the Tertiary and Recent Tadarida, indicating a rather noticeable anatomical stability, not exceptionnal indeed among Chiropterans. The Céreste fossil exhibits however slightly primitive wing proportions if compared to the Saint Gérand Aquitanian species.

Several remarks deal with the peculiar relationships between the ecology of the molossids and their kind of fossilisation, frequently associated with sedimentary facies of the lacustrine type.
  

The biometric and morphologie variability of the cheek teeth in the end-of-the-phylum species Lophiodon lautricense Noulet studied in this note, reposes on the observation of about 800 teeth. These were revealed to be little variable in absolute dimensions. The considerable morphologie variability in the upper premolars permitted the problem of the molarization process to be taken up. An hypothesis concerning the order of eruption of the cheek teeth is formulated based on an examination of a large number of milk dentitions. In conclusion, it is suggested that reservations be held on the value of dental characters classically used in systematics for the group under consideration. 

The rodent-fauna (Cricetidae and Gliridae) recently found at Castell de Barbera (Spain) is similar to those from the other locslities of the Valles - Penedes - Can Ponsic 1 and Can Llobateres - : same composition and similar evolutionary level of the different species. On the other hand this fauna is different from those of Upper Vindobonian and Vallesian localities of the Calatayud - Teruel area. Castell de Barbera has an intermediate chronological position between the localities of Anwil (Switzerland) and Can Ponsic 1. It is still not possible to validate or invalidate the initial attribution based on absence of Hipparion of Castell de Barbera to the Upper Vindobonian. 

Until now Aplodontidae of the European Oligocene have been documented by four species only. The phylogenetic relations remained obscure. as the distribution of only one species has been known in some detail. New material made it possible to define the stratigraphic range of two of the already existing species (Plesispermophilus angustidens, Sciurodon cadurcense) and to follow their development during the Oligocene beginning with the event of the « Grande Coupure ››. Sciurodon remained nearly without change until the end of the Middle Oligocene. Plesispermophilus angustidens split into two distinct phyletic lines, one of which (P. macrodon n. sp.) reaching considerable size, is represented till the beginning of the Upper Oligocene (Pech de Fraysse, Gaimersheim). The other line leads to Plesispermophilus ernii (basal Upper Oligocene of Burgmagerbein 1. terminal Upper Oligocene of Coderet). Besides the already known forms a new small-sized species (P. atavus n. sp.) is described, which by its primitive features closely resembles the genus Plesispermophilus. Two other small-sized species already known from the Upper Oligocene (? P. argoviensis) and Lower Miocene (? P. descedens) seem to be closely related to the new species. It cannot be decided whether they are descendents of this line or have developed independently, because of their poor fossil record.
Comparison of the evolutionary modalities in the different phylogenetic lines reveals general trends. the most striking of which is the complication of the tooth pattern by the development of additional crests. In the lower molars the cusps diminuate in size and are more and more transformed into ridges. ln addition new connection between the crests appear. in the upper molars, the « selenodont » shape of the teeth becomes more and more dominant, and in the two main evolutionary lines of Plesispermophilus the metaconulus becomes duplicated. A further evolutionary trend is the size increase of the premolars compared to the molars, which is even more pronounced in the Miocene Aplodontidae.
Phylogenetic relations between the primitive Plesispermophilus and certain « prociurines ›› of Northern America as well as between Plesispermophilus (P. angustidens) and more progressive forms of the Upper Oligocene (P. ernii, P. macrodon n. sp.) can be documented. In this light, the taxonomic distinction between Prosciurinae (bunodont) and Allomyinae (selenodont) sensu Rensberger 1976 can be shown to be artificial, because it separates forms from each other, which are evidently closely related. Consequently the separation into two subfamilies has been abolished. 

The rodents (Cricetidae, Gliridae, Sciuridae) found in lacustrine, brackish marine and karstic sediments of Miocene age in Languedoc, assign the position of the different localities in the scale of "niveaux repères" used by mammalogists. Some detailed stratigraphical studies bring several correlations between this continental biochronological scale and the marine scale ; the most important results are the Aquitanian age of the "niveaux repères" of Coderet and Paulhiac, the Burdigalian age of Laugnac, Estrepouy, Vieux-Collonges, La Romieu and Sansan and the Langhian or Lower Serravallian age of La Grive M. The correlations between the Tethys and the Central Paratethys for the Lower Neogene profit also of these results, since the locality of Neudorf Spalte 1, 2 (Czechoslovakia) is shown to be younger than Sansan (France). The paleontological study has also several geological inferences for the Miocene of Languedoc ; with the calibration of this Miocene, we know quite precisely that the Lower Miocene is chiefly a time lacustrine sedimentation, and also that the marine Miocene sedimentation ends early in the Miocene Period, in Langhian or lower Serravallian times. 

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

The complex taxonomic history of the paramyid rodent genus Ailuravus is reviewed. It has been described as Hyracotherium, as a creodont carnivore and as a lemuroid primate - errors at the ordínal level that are most unusual for a rodent. The genus is a member of the poorly known subfamily Ailuravinae, probably derived from some European Early Eocene species of Paramys. Aíluravus was a large arboreal paramyid with highly rugose cheek teeth, very well developed hypocone, and a remarkably weak lower incisor. It was tropical to subtropical. Three named species are recognized, A. macrurus from the Lutetian of Messel; the genotype, A. picteti, from Egerkingen, Buchsweiler and the Geiseltal, slightly later in the Lutetian; and A. stehlinschaubi, new name, from the Bartonian of Mormont-Eclépens and Robiac. One or more unnamed species are present in the Ypresian of Cuis. The species are close to a phyletic sequence. No later representatives of the genus are known. The late Eocene to earliest Oligocene North American paramyid Mytonomys, whose relationships have been obscure, is tentatively referred to the Ailuravinae. 

The recent excavation of the Upper Miocene locality of Montredon (Hérault) provides a rich material of Mammals. The present study deals with rodents. The new quarry is 10 meters above the classical Depéret's Quarry. No differences have been noticed between the rodent populations coming grom these two sites. The description of two new species : Hispanomys mediterraneus and Muscardinus heintzi is given and the dormouse Eliomys is recorded there for the first time. The new zonation of the Miocene given by Aguilar (1982) and the previously established correlations between marine and continental chronological scales give Tortonien age (estimated between 9 and 10 M.Y. old) to the site of Montredon, which belongs to the Mammal zone D 2. 

The Montredon local fauna yielded very rare bats, represented by damaged isolated teeth. Only a few documents are available for this period of the European Neogene. ln this poor state of knowledge, the material represents three undetermined species, a supposed molossid and two vespertilionids. 

A new middle Miocene hipposiderid bat is described from a limestone deposit on Riversleigh Station in north-western Queensland. Hipposideros (Brachipposideros) nooraleebus n. sp. is the first record of this subgenus from anywhere in the world outside of France. The palaeoecological setting of the fossil bats appears to have been a relatively quiet, sunny lime-enriched tropical pool that contained tortoises, crocodiles and fish. It is possible that the bats were washed into the pool from an adjacent cave.
The Riversleigh bat most closely resembles the French Burdigalian (early middle Miocene) bat H. (B.) aguilari. It is also possible that it may have been closely related to the original Australian hipposiderid stock that ultimately gave rise to the endemic monotÿpic Rhinonycteris aurantius. The disjunct distribution of species of H. (Brachipposideros) suggests that representatives of this subgenus will be found in at least tropical southern Asia. 

The numerical importance of the Condylarths in the Cernaysian fauna is discussed. The Condylarth family, Tricuspiodontidae, is reviewed in the light of new material and its close relationships to the Phenacodontidae is suggested ; one new species is recognized : Tricuspiodon sobrinus. European Arctocyonidae are reviewed and the recentclassification of Van Valen is briefly commented on. Also,  the arctocyonine Landenodon is described for the first time in Thanetian (Late Paleocene) sediments ; two new species are proposed : T. lavocati and T. phelizoni.