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".

  

The Montian locality of Hainin (Hainaut, Belgium) yielded about twenty teeth of Multituberculates. They are very peculiar forms, showing no affinities, at the generic level, with those hitherto known from North America, Asia and Europe. They are referred to the new taxa Boffius splendidus nov. gen., nov. sp., Hainina belgica nov. gen., nov. sp., and H. godfriauxi nov. gen., nov. sp. They expose some common features, such as the advanced type of first upper molar. possessing at least three complete rows of cusps. Because of this, and also of the upper premolar reduction, Boffius splendidus appears as the most specialized form within the Ptilodontoidea suborder.
Several other characters of Hainina seem to be less advanced, such as the great number of upper premolars and the simple cusp-formula of the first lower molar.
Till now, only H. godfriauxi has been recovered within the Thanetian fauna from Cemay-les-Reims, where it is very poorly documented. 

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. 

The intensive exploration of the Pliocene Hadar Formation, rich in hominid remains, led us to the discovery of several micromammals levels. ln some of them, rodents are very abundant. The stratigraphic repartition of these levels do not cover the whole fossiliferous series of the formation but takes place only in the sedimentary members from Sidi Hakoma and Denen-Dora (rancing from 3.1 - 3.2 MY to 2.8 - 2.9 MY, according to the recent geochronological data). During this gap of time, the species do not show morphological changes, what allowed us to gather, in the same taxa, forms of slighty different ages.
Two striking facts, giving a lot of indications, characterize these small rodents'faunas. First, we notice the domination of the Muridae, as well on a qualitative way (number of species) as on a quantitative one (number of individuals). Then, it appears that, until now, two genera of these murids were known only in the south-western asiatic regions. So, we can suppose continuous biotops between Africa and Indian Subcontinent before 3 MY. In this hypothesis, the hominids had already the possibility to leave their african « cradle ››. Finally, almost all studied genera are still represented at the present time. This fact, previously observed in Laetolil, Omo, Olduvai contributes to remove hope of establishing a biochronological scale based on rodents, in tropical zone. Nethertheless, that allows to try a reconstruction of the palaeoenvironnement, by using the principle of actualism.
  

A reconstruction of a multituberculate endocranial cast is made on the basis of a complete natural cast prepared from the skull of Chulsanbaatar vulgaris, and other less complete endocasts, all from the Late Cretaceous of Mongolia. The multituberculate endocast is of mammalian pattern but it has retained a therapsid-like lateral profile with a deep rhombencephalon and a shallow telencephalon. It is characterized by: a heart-shaped cross-section of the telencephalon; an extensive lissencephalic neocortex; a very prominent pons placed far anteriorly; a lack of cerebellar hemispheres, and very large paraflocculi. Its structure, very different from the brains of other mammals, suggests thats the Multituberculata branched very early from the main mammalian stock. This supports Simpson's (1945) idea that the Multituberculata should be placed in a subclass of their own: Allotheria MARSH. The endocast and braincase structure show that the Multituberculata had strongly developed senses of smell and hearing. The encephalization quotient of approx. 0.55 evaluated for Ch. vulgaris is relatively high for a Mesozoic mammal. 

The formic acid preparation of a nodule collected from the Lower Triassic of Spitsbergen, and containing a skull with some associated postcranial elements of Lyrocephaliscus euri (WIMAN), precises and enriches previous descriptions of the reputedly well-known trematosaur. lt permits more detailed descriptions of the occipital region, of the course of the carotid arteries and collateral vessels, and refutes the previous description of a stapedial foramen in trematosaurs. The discovery of pleurocentres associated with the other vertebral components confirms the primitive rhachitomy of trematosaurs. 

The oldest european marsupials are described from some specimens (isolated upper molars) recently found from the Hainin sediment (Middle Paleocene of Belgium). These fossils document a new species of the Peradectes genus. They give evidence of a much older occurrence of the marsupials in Europe than it was assumed. They allow us to postulate a didelphid dispersal from South America towards the western-holarctic area operating in two phases : the first one of the Peradectes genus at the end of the Cretaceous; the second one of the Didelphíni tribe at the end of the Paleocene. A central american crossing is likely for the first one,  whereas a transafrican way is tentatively argued for the second one. 

An ilium from the Upper Eocene (MP l7a) of Hordle, England, represents the first report of Rana from the Eocene of Britain. The ilium is similar to those of the water frog (Rana [ridibunda]) species group. 

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. 

Le groupe des Mustélides « Ischyrictoïdes » développe des tendances évolutives qu’engendrent des formes de Carnassiers prédateurs robustes et de grande taille. C’est une évolution comparable à celle que l’on trouve chez Plesiogulo, qui, lui, a une origine énigmatique (Petter, 1963). Chez les Mustélides on peut signaler deux lignées assez nettes : celle du sous-genre nominal des Laphyctis, plus conservatrice, plus mustéliforme, renfermant les espèces Ischyrictis (Laphyctis) mustelinus VIRET et L. zibethoides BLAINVILLE ; celle de type félinoïde caractérisée par L. Ginsburg (1961) avec la forme type Ischyrictis (Hoplicitis) florancei du Vindobonien inférieur de Pontlevoy-Thenay et I. helbingi VIRET, 1951 du Vindobonien supérieur de la Grive Saint-Alban. De plus rappelons que I. vorax DEHM et I. comitans DEHM du Burdigalien de Wintershof-West sont sans rapport avec les espèces du Miocène supérieur (Ginsburg, o.c.).

Récemment G. Petter (o.c.) a montré la difficulté qu’il y avait à envisager que l’Ischyrictis de Can Llobateres qu’elle décrivait puisse être le descendant de Ischyrictis helbingi de La Grive : la forme plus récente est évidemment de plus grande taille, mais le développement très marqué et la position avancée du protocône paraissent constituer un obstacle à une telle filiation.

Donc on signalait jusqu’ici deux lignées de Mustélides « ischyrictoïdes » toutes deux de grande taille : d’une part I. zibethoides de Sansan, d’autre part I. helbingi de la Grive. Or à notre point de vue nous considérons actuellement qu’il existe très probablement trois phyla d’Ischyrictis. Le premier serait représenté par les formes burdigaliennes déjà nommées I. vorax et I. comitans ; il se caractériserait par la perte précoce de la P₁ inférieure et la forme spéciale de la canine inférieure. Ce premier ensemble pourrait être rassemblé dans le sous-genre Laphyctis. Le reste des ischyrictoïdes de type conservateur, pour ce qui est de la région des prémolaires et de la forme de la canine, pourrait être subdivisé en deux sous-genres : le sous-genre Ischyrictis désignant les formes les plus mustéliformes avec I. mustelinus et I. zibethoides d’une part, et le sous-genre Hoplicitis pour les formes félinoïdes avec I. florancei et I. helbingi. 

This study examines the microfauna from Djebel Qafze Cave, focusing on mammalian remains, with Microtus guentheri as the dominant species and a notable absence of Cricetinae (hamsters). The richest layers (XVIa, XVII, XVIII) suggest accumulation by raptors, particularly owls, given the high presence of nocturnal species like voles and shrews. Two significant species, Arvicanthis cf. ectos and the newly described Rattus (? Mastomys) nazarensis, are highlighted. Reptiles (gekkos, Ophisaurus) and birds are also present, while hedgehogs and lagomorphs are absent. The findings underscore the role of avian predators in fossil deposition and refine the chronology of regional Pleistocene fauna. 

Tarsals that can be confidently attributed to Paschatheríum dolloi from Dormaal (Belgium) are described. The astragalus is short; its broad neck is set at an angle of 30 degrees to the trochlea. The trochlea is pulley-shaped and proximally extended. There is no astragalar foramen. The medial tibial facet extends distally in a cup deeply excavated in the neck and buttressed. The sustentacular facet extends toward the navicular facet but is not fully confluent with it. The calcaneum bears a proximo-distally extended proximal facet for the astragalus. A relatively large peroneal tubercle projects from the body, and is situated distally. Functionally, the trochlea indicates extensive flexion-extension movements of the foot. Calcaneo-astragalar facets indicate sliding and rotation between the two tarsals. The inclination of the navicular facet suggests frequent foot inversion. The peroneal tubercle reflects good muscular capacities for foot rotation. Overall morphology is interpreted as a scansorial adaptation similar to that of sciurids.Comparisons are made with the tarsals of Macrocranion vandebroeki from Dormaal and Hyopsodus paulus from the Bridgerian of Wyoming. Some similarities between the astragali of Paschatherium and Macrocraníon (trochlea) are interpreted as convergences for rapid locomotion. However absence of mobility at the lower ankle and midtarsal joints in M. vandebroekzi suggests that this species was cursorial, as is known for M. tenerum from Messel. Similarities in the calcanea of Paschatherium and Hyopsodus are probably the result of close phylogenetic relations, and confirm the placement of Paschatherium in the hyopsodontids. The differences in the calcanea of Paschatherium and Macrocranion underline that Paschatherium is distinct from erinaceomorph insectivores. The differences in astragalar morphology between Paschatherium and Hyopsodus show that a marked adaptive divergence separates the two genera. We speculate about the common occurrence of a deep tibial cup (“cotylar fossa”) and a pulley-shaped trochlea in Paschatherium and hyracoids, suggesting that an adaptive scenario similar to that having led to Paschatherium (scansoriality) might explain the acquisition of the peculiar hyracoid tarsal characters; such a scenario contradicts the concept of Pantomesaxonia. Other peculiar characters of Hyopsodus suggest that hyopsodontids might be given more consideration in the search for hyracoid (and tethythere) origins. 

An upper giraffid premolar without any indication about its origin is preserved at the Montpellier University among numerous fossils from the ruscinian formation of Montpellier. It can be related to Samotherium, of the Upper Miocene in Eastern Europe, North Africa and Asia, or more probably to Bramatherium  or Hydaspitherium of the Pliocene of South East Asia. The sedimentological study of the matrix shows a calcareous background, which may indicate that this tooth does not come from the Montpellier formation. 

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

Is hereby described an impressive lchnoiauna belonging to the Lower to Middle Ludian of the Gard (S. France). The slab, already cleaned over a length of 18 m, is located near the top of the Potamides aporoschema lacustrine limestone (Lower Ludian, Euzet zone). It is therefore older than the Célas sandstone deposit, and still more than the Melanoides albigensis and M. acutus marly limestone corresponding to the Upper Levels of the Ludian stage. Although biostratigraphically older than the La Débruge and Montmartre zone, the biotope shows already a sampling of very tall Artiodactyles, Perissodactyles and Carnivorous. One of the most « majestic ›› Artiodactyles, Anopolotheriipus lavocati, nov., points out a huge size type. To mention also among the Ichnotypes described, 10, the big Perissodactyle Palaeotheriipus similimedius, nov., and the big Carnivorous Hyaenodontipus praedator, nov. 

The study of dental variations in Landenodon from Dormaal (early Eocene, Belgium) shows that only one
species is present in this locality : L. woutersi QUINET, 1968 (Condylarthra, Arctocyonidae, Arctocyoninae). A high morphological variability is described, especially concerning the metaconid of P/4, the paraconid of M/2, and some characters of M1/ which have been found to be independant of each other. This variability is discussed. 

The genus Microstonyx was found only in the north eastern part of Spain : M. antiquus, referable to the
Eppelsheim type, from the pyrenean basins ; M. antiquus, more progresive, from the Vallès Basin : M. major-erymanthius group, from the Teruel-Ademuz Area and from Pieru (Penedès Basin).

  

The Arvicolid evolution results in an increase of the dental structure complexity. The M3/ differenciation seems to characterise the tribe subdivisions, that of M/1 being variable from one to another lineage. The phyletic relationships of fossil lineages are discussed from a paleogeographic point of view. 

The rodent fauna of Beni-Mellal is characterized by the abundance of ctenodactylids and cricetids. The latter are represented by four distinct species, among which a new form. Dakkamys zaiani nov. gen., nov. sp. is described. A detailed morphological analysis shows that, contrary to that which had been established before, « Cricetodon ›› atlasi Lavocat, 1961, is not closely related to any European form known; this species is attributed, in consequence, to the new genus Mellalomys. A simple biometric analysis has shown that the genus Myocricetodon Lavocat, 1952, is represented in this locality by two distinct species. The systematic homogeneity of the Beni-Mellal cricetids is also demonstrated: they can, as a matter of fact, all be referred to the subfamily Myocricetodontinae. The definition of this subfamily is completed. The sciurids and glirids are also reviewed in the light of new systematic and biogeographic information established ln Europe. A new species of Atlantoxevus from the early Pleistocene of Morocco, A. huvelini nov. sp., is described. It is probably the descendant of A. tadlae from Beni-Mellal. Biogeographic analysis leads one to consider this fauna as the result of geographic isolation in the Maghreb since the late Oligocene or the early Miocene. In particular no direct European influence can be discerned. Stratigraphic considerations resulting from the discovery of new localities in North Africa lead to the confirmation of the ante-Vallesian age of this fauna and to its parallelism with the faunas of La Grive in Western Europe and Fort Ternan in East Africa. The peculiar geologic nature of this locality is discussed.
  

La poche à phosphorite de Sainte-Néboule (Lot) a livré au cours des récentes campagnes de fouilles effectuées dans le cadre de la RCP 311 une douzaine de dents ou fragments de dents de paléothéridés à rapporter à 3 taxons.