Reassessment of the Morphology and Taxonomic Status of the Earliest Herpetotheriid Marsupials of Europe

The earliest Eocene locality of Dormaal (Belgium) has provided the oldest Cenozoic herpetotheriid marsupials of Europe. No herpetotheriid has ever been reported earlier than the Eocene in Europe, except for a questionable single upper molar from the Upper Cretaceous of the Belgian/Dutch border. The systematics of the herpetotheriids of Dormaal was formerly based on only a dozen dental specimens, which were assigned, after several revisions, to two species Peratherium constans and Amphiperatherium brabantense. Most importantly, these two species were considered at the root of most of the hepetotheriid lineages of the European Paleogene. Here we report a large sample of about 400 new dental remains that allow a better definition of both species as well as a testing of their systematic status. The evidence of significant morphological variability leads us to reconsider the diagnosis of Peratherium constans and to question the validity of Amphiperatherium brabantense. This study highlights that the primitive species Peratherium constans and Amphiperatherium brabantense are hardly distinguishable from each other, and therefore conclude that Peratherium constans was the only marsupial present at Dormaal. The important morphological variation exhibited by this herpetotheriid is similar to the variability observed in the type-species Peratherium elegans and in other fossil and extant metatherians. Consequently, our results suggest that several Amphiperatherium species from the Eocene could represent variants of the genus Peratherium. The question of the Amphiperatherium presence in Europe is therefore raised and a thorough discriminate analysis of both genera should be conducted in later works.     

A new genus of Bothremydidae (Chelonii,Pleurodira) in the Cretaceous of Southwestern Europe

Although the Bothremydidae are the most abundant and diverse group of turtles in the Upper Cretaceous fauna of Europe, the record attributed to the genus Elochelys was, until now, quite scarce. However, the revision of the specimens previously attributed to this taxon and the analysis of new material from the Spanish record significantly increase our knowledge of these turtles. Evaluation of the morphological variability, affecting many characters that have been considered diagnostic until now and comparisons with other members of the node Bothremydini, allow us to amend the diagnosis of the genus Elochelys, recognising Elochelys perfecta as a valid single species and reassigning its other representative, “Elochelys” convenarum, to the new genus Iberoccitanemys.     

Presence of Zanthopsis M’Coy, 1849 (Crustacea: Decapoda: Brachyura) in the Early Eocene of Iran,and comments on the different species and/or morphotypes and their palaeobiogeography

In the present work, we report the discovery of several carapaces of Zanthopsis sp. from the Lower Eocene of Central Iran, which represent the easternmost record of the genus. In addition, we discuss on the similarities of the different species and/or morphotypes of the genus, and conclude that the European species could be ascribed to the “Zanthopsis leachii complex”, until future works determine the validity of the species therein contained. In this sense, Zanthopsis jacobi Van Straelen, 1924, in agreement with Artal and Vía (1988), is herein considered a junior synonymous of Z. dufouri (H. Milne Edwards in d’Archiac, 1850). The presence of Zanthopsis in the lower Eocene of Iran suggests homogeneity of the brachyuran fauna on both sides of the Tethys Realm.     

Remains of sea turtles from the Ikovo locality (Lugansk Region, Ukraine; Middle Eocene)

Fragmentary remains of sea turtles (Cheloniidae sensu lato: Argillochelys sp., Puppigerus nessovi Averianov, 2005, and Cheloniidae gen. indet.) from the Ikovo locality (Lugansk Region, Ukraine; Lower Lutetian, Middle Eocene) are described. The genera Argillochelys and Puppigerus are recorded for the first time in Eastern Europe. The turtle assemblage from Ikovo is similar at the generic level to West European assemblages (Belgium, Great Britain) of approximately the same age (Lower-Middle Eocene). In the presence of P. nessovi, the Ikovo assemblage is similar to that from the Middle Eocene Dzheroi 2 locality (Uzbekistan).     

First fossil record of the genus Nageia (Podocarpaceae) in south China and its phytogeographic implications

The first fossil record of the genus Nageia Gaertner from the Tertiary of southern China is reported. Nageia hainanensis sp. nov. is described from the Eocene Changchang Formation of Hainan Island. The extant species of Nageia are widely distributed in southeastern Asia, from northeastern India to southeastern China and southern Japan, and southward through Indochina to the Malay Archipelago and New Guinea. The fossil evidence of Nageia has been discovered from the Lower Cretaceous of the Far East of Russia and Japan. The distribution of both modern and fossil species of Nageia indicates that this genus could have originated in the northeastern part of Asia in the Early Cretaceous and spread to south China at least in the Eocene. Discovery of Nageia fossil record in south China provides important evidence for the geological and phytogeographic history of the genus.     

The “miacids” (Carnivoraformes,Mammalia) from the Early Eocene locality of Le Quesnoy (MP7, France); first occurrence of Vassacyon in Europe

We describe here “miacid” taxa from the Early Eocene Paris Basin locality of Le Quesnoy (Oise, France). We describe the new species Vassacyon taxidiotis, the first European record of this genus. The other “miacids” identified from Le Quesnoy are Miacis latouri and Gracilocyon solei. The P4 of G. solei is described here for the first time. Its morphology (e.g., wide protocone, short postmetacrista) supports a close relationship with Miacis rundlei from Abbey Wood (MP8 + 9, England). The latter species is therefore classified as Gracilocyon rundlei. Three new tooth positions are known for Miacis latouri: P4, p4 and m2. They support its reference to Miacis. These specimens imply that the European species is more basal than the North American species. The fauna from Le Quesnoy shares with Dormaal the presence of Miacis latouri and Gracilocyon solei, but the “miacid” fauna from Le Quesnoy also contains Vassacyon taxidiotis. The presence in Le Quesnoy of the two former taxa supports a reference to MP7 level of the French locality. The presence of three distinct genera in European localities show that the “Miacidae” were diversified in Europe, as previously observed in North America. The genera Gracilocyon, Miacis, and Vassacyon probably dispersed from Europe to North America during the Palaeocene/Eocene boundary.     

Présence du genre PaléocèneAdapisorex (Lipotyphla,Mammalia) dans les Sables d'Orp-le-Grand (Heersien) a Maret en Brabant (Belgique)

According to the classical stratigraphical conceptions, the Belgian Heersian was classified in the Landenian, and assigned to the basal Lower Eocene. It is composed of marine Sands of Orp-le-Grand, which contain a riche selacian fauna, also of Gelinden Marls covering and interpenetrating the Sands.Among the vertebrate remains recently discovered at Maret en Brabant in the marl facieswithin the Sands was a small tooth which represents the first mammalian remain from the Heersian, and attests the occurence of the insectivorous genus Adapisorex, hitherto known from the paleocene localities of Cernay-les-Reims and Walbeck. The specimen from Maret en Brabant reopens the question of the age of the Belgian Heersian and supports its classification within the Paleocene, according to paleobotanical data and indications obtained from the selacian fauna.     

Deux nouvelles tortues de l'Éocène inférieur de Saint-Papoul (Aude, France)

Two new turtles from the Lower Eocene of Saint-Papoul (Aude, France). Among the fossil turtles from of the Lower Eocene of Saint-Papoul (Aude, France), two new taxa of freshwater Testudinoidea are described. The first is a new species of the genus Palaeoemys, known before in the Lutetian of Germany. The second taxon is a new genus, also described on material of Pépieux-Cesseras and Eygalayes. To cite this article: S. Hervet, C. R. Palevol 2 (2003).     

Early Paleogene decapod crustaceans from the Sulaiman and Kirthar Ranges,Pakistan

A set of Paleocene and Eocene decapod crustaceans is described from the Sulaiman and Kirthar Ranges of Pakistan. The fossil crabs Proxicarpilius planifrons Collins and Morris, 1978 and Pakicarcinus orientalis (Collins and Morris, 1978), already known in the Eocene of northern Pakistan, are reported for the first time in the Paleocene of southern Pakistan, enlarging the stratigraphic and the palaeobiogeographical ranges of these species. The callianassid genus Calliax de Saint Laurent, 1973 is reported for the first time in the Paleocene of southern Pakistan; this is the oldest record for the genus.     

New generic proposal for the European Neogene large testudinids (Cryptodira) and the first phylogenetic hypothesis for the medium and large representatives of the European Cenozoic record

Fossil testudinids are known in Europe since the Eocene, with several taxa of medium size (from more than 0.3 m to less than 0.7 m) recognized in the Palaeogene record, most of them being poorly known. The size of several European Neogene taxa was larger (between 1 and 2 m). These large testudinids were relatively abundant and diverse, ranging from the early Miocene to the Pleistocene. However, there is a nomenclatural gap at the generic level for the Neogene forms, as their generally used assignment to the more primitive Eocene Cheirogaster cannot be sustained. This is because relatively little material has been assigned to the described species, and also because of the absence of a detailed study comparing all of the European taxa. Here, the European Cenozoic taxa are incorporated for the first time in a data matrix, so that a hypothesis on their phylogenetic relationships is justified. This study identified the large testudinids from the Neogene of Europe as belonging to a monophyletic clade, assigned to the new genus T itanochelon . The hitherto poorly understood ‘Testudo’ bolivari, proposed nearly a century ago but lacking diagnosis, is analysed in detail. It is recognized as the best‐represented large testudinid from the European record, and is identified as the type species of T itanochelon gen. nov. Its comparison with the other Neogene species allowed a detailed study of the new genus and an analysis of its phylogenetic relationships with the other European taxa. © 2014 The Linnean Society of London     

Fish otoliths from the Priabonian (Late Eocene) of North Italy and South-East France - Their paleobiogeographical significance

The study of the Late Eocene (Priabonian) otolith associations from Possagno, North-East Italy, and from the Synclinal d’Allons in Haute Provence, South-East France, allows for the reconstruction of a teleost fauna of 55 taxa, which is the most diversified assemblage presently known from the Upper Eocene Paleo-Mediterranean basin. Thirty-six taxa are identified at the species level, and five of those are new: “genus Alepocephalidarum” astrictus, “genus Lophiiformorum” canovae, “genus Agonidarum” sudans, “genus Uranoscopidarum” cochlearis and Aseraggodes laganum. In the Synclinal d’Allons, the otolith associations reflect a tropical to subtropical neritic environment with a few mesopelagic fishes. At Possagno, the associations indicate an environment that changed from one that was deep and exposed to the pelagic realm and then evolved to a more shallow sea with a well-diversified benthic life and less mesopelagic fishes. A paleobiogeographical analysis of all known data on Priabonian otoliths, worldwide, shows clearly a western Atlantic (Louisiana) and an eastern Atlantic-Paleomediterranean association. In the eastern Atlantic-Paleomediterranean association, the Aquitaine association not only differs from the Possagno-Allons association in function of its more distant geographical position, but also by its stronger oceanic character in the southern part of the basin, and by the occurrence in the north, of a very shallow water facies (Saint-Estephe Formation) that contains some taxa which are known nowhere else in the Priabonian. The Ukraine fauna is characterized by a high number of species, which have an Oligocene record in other European sites. The northern geographic location of Ukraine, combined with the good connections to both the North Sea Basin and the Turgai street can provide the explanation. Many Oligocene species (or their close relatives) probably already existed at Eocene times in more northern regions, but could penetrate only in more southern European seas since the strong cooling at the beginning of the Oligocene.     

Evidence for a pre-PETM dispersal of the earliest European crocodyloids

Crocodyloid remains from the late Paleocene of Mont de Berru (France) hosted in the collections of the Muséum National d’Histoire Naturelle (Paris, France) and in the Institut royal des Sciences naturelles de Belgique (Brussels, Belgium) are described for the first time. This material, although fragmentary, can be clearly referred on a morphological basis to Asiatosuchus depressifrons (Blainville, 1855), a species previously reported from several Eocene Belgian localities thanks to abundant material including a nearly complete skeleton. The Paleocene material shares with A. depressifrons the number of alveoli involved in the dentary symphysis, the exclusion of the splenials from the symphysis, and the presence of a distinct depression on the jugal. The fossil remains from Berru represent the oldest European crocodyloid. Along with the alligatoroid Diplocynodon remensis Martin, Smith, de Lapparent de Broin, Escuillié and Delfino, 2014, previously reported from the same locality, the crocodyloid A. depressifrons indicates that these genera reached Europe before the Paleocene–Eocene Thermal Maximum. Although more complete remains from outside Europe are needed to refine phylogenetic hypotheses, according to the currently established fossil record the forerunners of diplocynodontids likely dispersed from North America, whereas those related to Asiatosuchus likely dispersed from Asia.     

A new species of Neochelys (Chelonii,Podocnemididae) from the Ypresian (Early Eocene) of the South of France

The known European record of the Gondwanan group Podocnemididae begins in the Early Eocene. Neochelys underwent a rapid diversification and was an abundant and diverse representative of this group during the Eocene of Europe from the Early Ypresian to at least the Bartonian. However, several of its species are poorly known, and the phylogenetic relationships among them are poorly understood. A complete shell from the Ypresian of Hérault (southern France) is described here. It is assigned to a new species, Neochelys liriae. The availability of characters of N. liriae is tested by comparison with the other European species, which appear to be well differentiated, and particularly with the neighbouring French taxa N. eocaenica and N. laurenti, of which the latter has been little known to this point.     

A new seabird (Aves, cf. Phaethontidae) from the Lower Eocene phosphates of Morocco

A proximal part of humerus from the basal Ypresian (lowermost Eocene) of the Ouled Abdoun Basin, Morocco, is described as a new genus and species tentatively assigned to the Phaethontidae (tropicbirds). This fossil possibly represents the oldest record of the Phaethontidae and markedly differs from Lithoptila, a contemporaneous Prophaethontidae from the same locality. This new taxon lived in a tropical climate and was probably an efficient flier with pelagic habits, like extant tropicbirds of the genus Phaethon.     

Diversity and paleoenviromental significance of Brazilian fossil Galictis (Carnivora: Mustelidae)

The evolutionary history of the genus Galictis in South America probably begins after the Great American Biotic Interchange. Two species are recognised: Galictis vittata and Galictis cuja. The latter are more frequently found in open areas in southern South America and the first occurs in humid forests from northern South America to Central America. Apparently, they do not occur in sympatry. Both are differentiate by the presence of a metaconid in the first inferior molar of G. vittata and for its bigger size when compared to G. cuja. The fossil record of Galictis is scarce, G. cuja is known by few specimens from Argentina, Chile and Brazil; G. vittata have only one record from Southern Brazil. The specimens related to this record were collected by Peter Lund and are housed at the Statens Naturhistoriske Museum. However, the specimens published by Lund are not fossils. Thus, it is presented here other unpublished specimens collected by Lund and housed at the same museum that we recognise as the first G. vittata fossils. Additionally, it is described here the first fossil record for G. cuja from the late Pleistocene of Brazil – an almost complete mandible recovered from sedimentary deposits from Central Brazil.     

Wood of Tapirira (Anacardiaceae) from the paleogene Clarno Formation of Oregon

A new species of fossil wood, Tapirira clarnoensis, is described from a probable late Eocene deposit in the Clarno Formation of Oregon known locally as the Nut Beds. The wood represents, to the knowledge of the writer, the earliest occurrence of Tapirira in the fossil record and the only occurrence of the genus outside its present neutropical range of distribution.     

Sinopaninae and Arfianinae (Hyaenodontida,Mammalia) from the Early Eocene of Europe and Asia; evidence for dispersal in Laurasia around the Paleocene/Eocene boundary and for an unnoticed faunal turnover in Europe

Hyaenodontida are represented in Europe by three subfamilies: Proviverrinae, Arfianinae and Sinopaninae. Here, we review all the specimens of Arfianinae and Sinopaninae known to date in Europe and Asia. A new Galecyon species is erected: Galecyon gallus nov. sp. We discuss the taxonomic position of the two Asian hyaenodontidans Anthracoxyaena palustris and Arfia langebadreae; the genus Anthracoxyaena is synonymized with Arfia. The analysis of the European and Asian arfianines and sinopanines provides new data concerning the dispersals and faunal events that occurred during the Early Eocene in Laurasia. The Arfianinae and Sinopaninae appeared in Europe around the Paleocene/Eocene boundary (reference-level MP7). The sinopanines are widespread in Europe; they are known in Dormaal (Belgium, reference-locality of the level MP7), Rians, Soissons, Pourcy, Try, Le Quesnoy (France), and Abbey Wood (England). The analysis of the paleogeographic distribution of all Oxyaenodonta and Hyaenodontida at and after the MP7 supports the existence of two European provinces: the North Province and Mesogean Province. We show that the Arfianinae and Sinopaninae rapidly disappeared from Europe; they are unknown in Avenay (reference-locality of the level MP8+9) and younger localities. Their disappearance from Europe is synchronous with that of the Oxyaenodonta. These observations support the existence of a faunal turnover, which occurred between the reference-levels MP7 (Dormaal) and MP8+9 (Avenay). The hypothesis of a dispersal from Europe to North America during the Paleocene-Eocene transition for the Arfianinae and Sinopaninae is supported. Moreover, the study of Arfianinae supports a dispersal from Europe to Asia around the P/E boundary, followed by a short period of endemic evolution. However, our study does not support a close relationship between Arfia and the “Arfia-like South Asian Proviverrinae” (Kyawdawia, Indohyaenodon, Paratritemnodon and Yarshea).     

An ant-associated mesostigmatid mite in Baltic amber

Fossil mesostigmatid mites (Acari: Parasitiformes: Mesostigmata) are extremely rare, and specimens from only nine families, including four named species, have been described so far. A new record of Myrmozercon sp. described here from Eocene (ca 44–49 Myr) Baltic amber represents the first—and so far only—fossil example of the derived, extant family Laelapidae. Significantly, modern species of this genus are habitually myrmecophilous and the fossil mite described here is preserved attached to the head of the dolichoderine ant Ctenobethylus goepperti (Mayr, 1868). It thus offers the oldest unequivocal evidence for an ecological association between mesostigmatid mites and social insects in the order Hymenoptera.     

New material of Diacodexis (Mammalia,Artiodactyla) from the early Eocene of Southern Europe

Diacodexeidae are the first representatives of Artiodactyla in the fossil record. Their first occurrence is at the very base of the Ypresian (earliest Eocene, 56.0 Ma) with Diacodexis, a genus well diversified during the early Eocene in Europe, especially during the MP7–MP8 + 9 interval. However, most of European species are documented by scarce material, retrieved from single localities. In this work, we describe new Diacodexis material from ～MP7 and ～MP8 + 9 localities of Southern Europe, including material of D. antunesi from Silveirinha, considered as the most primitive European Diacodexis species, and material from three localities from Southern France (Fordones, Palette, and La Borie). The new material documents Diacodexis premolar morphology and deciduous dentition which bear potentially important phylogenetic information, as well as astragali, including a specimen from Silveirinha that constitutes the earliest occurrence of an astragalus of the genus Diacodexis in the European fossil record. Investigation of the enamel microstructure reveals that early European species had a simple enamel pattern with one-layered Schmelzmuster composed of ‘basic’ radial enamel only, instead of the two-layered Schmelzmuster (thin radial enamel + thick layer of Hunter-Schreger bands) observed on North American species and so far considered to represent the primitive condition within Artiodactyla. In accordance with previous studies, our observations highlight that Diacodexis gigasei from Belgium is morphologically closer to the North American species D. ilicis than to D. antunesi from Portugal. The latter species, together with D. aff. antunesi from Fordones, appears to be morphologically closer to the Asiatic taxa D. indicus and D. pakistanensis. Finally, we found numerous similarities between D. cf. gigasei from Palette and D. gigasei, a result that challenges the intra-European provincialism that characterizes the earliest Ypresian. Diacodexis gigasei could be one of the rare species shared by the northwestern and southwestern European bioprovinces.