The giant bird Gastornis in Asia: A revision of Zhongyuanus xichuanensis Hou, 1980, from the Early Eocene of China

The type specimen of the gastornithid Zhongyuanus xichuanensis Hou, 1980, from the Early Eocene of Henan (central China) is redescribed and compared with European and North American representatives of Gastornis. Differences between the Chinese specimen (the distal end of tibiotarsus) and its European and North American counterparts are of a minor nature, and it is concluded that it can be referred to as Gastornis xichuanensis (Hou, 1980). Stratigraphic evidence suggests that Gastornis originated in Europe, where it is known as early as the Paleocene. Whether the Asian Gastornis dispersed from North America via the Bering land bridge, or directly from Europe across the Turgai Strait, remains uncertain.     

Intraspecific variation of Eolophiodon laboriense,a basal Lophiodontidae (Mammalia,Perissodactyla) from the early Eocene of Southern France

The Lophiodontidae is an emblematic and well-documented Eocene family of perissodactyls from Western Europe. However, after more than a century and a half of studies, lophiodontids still display a complex systematics associated with blurry intraspecific variation and a poorly known early radiation. The locality of La Borie, located near the city of Toulouse, France, has yielded numerous remains of Eolophiodon laboriense. This abundance of remains allows for the first time the study of the intraspecific variation of a basal lophiodontid. The variation has been investigated for dental and cranio-mandibular characters, notably dental polymorphism, size variation and sexual dimorphism. The intraspecific variation of E. laboriense is high with more than 20 polymorphic characters of the dentition, including many additional crests and conules. This dental polymorphism is similar to the one observed in the Bartonian lophiodontid Lophiodon lautricense. E. laboriense also displays an important degree of sexual dimorphism, with male specimens having broader and longer mandibles with larger canines than females. Despite this high intraspecific variation, the low size variation of teeth and the consistency of diagnostic characters strengthen the validity of the genus Eolophiodon and does not impact the previous lophiodontid phylogeny.     

New proviverrines (Hyaenodontida) from the early Eocene of Europe; phylogeny and ecological evolution of the Proviverrinae

We describe six proviverrine species from the Early Eocene of France. Three species are new: M inimovellentodon russelli sp. nov. from Mutigny [mammal palaeogene (MP)8 + 9], B oritia duffaudi sp. nov. from La Borie (MP8 + 9), and L eonhardtina godinoti sp. nov. from Grauves (MP10). We describe new specimens and propose new generic combinations for three species from MP10: Protoproviverra palaeonictides, Matthodon menui, and Oxyaenoides lindgreni. We also propose a new generic combination for the primitive Eoproviverra eisenmanni (MP7). Matthodon menui was previously considered as a possible oxyaenodontan, but the new fossils clearly support its reference to Hyaenodontida. Leonhardtina godinoti and Ma. menui are the oldest occurrences for these genera, which were previously unknown before the Middle Eocene. Moreover, the discovery of the proviverrine Mi. russelli in Mutigny implies that the Proviverrinae dispersed in Northern Europe between biozone Palaeocene‐Eocene (PE) III (Abbey Wood) and biozone PE IV (Mutigny). This also supports a homogenization of the European faunas during the Early Eocene. The dispersal is concomitant with the disappearance of the oxyaenodontans, arfiines, and sinopines (Hyaenodontida) from Europe. The proviverrines may have filled the ecological niches left vacant by the disappearance of the other carnivorous mammals. With 20 genera and over 30 species, proviverrines were successful in Europe. We performed the first phylogenetic analysis comprising almost all the Proviverrinae. Our analyses indicate that the Proviverrinae diversified greatly during the Early Eocene Climatic Optimum and show a general trend towards specialization throughout the Eocene. © 2014 The Linnean Society of London     

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.     

Paratritemnodon indicus (Creodonta: Mammalia) from the early Middle Eocene Subathu Formation,NW Himalaya,India, and the Kalakot mammalian community structure

A complete post-incisor upper dentition and a left mandible with P/3-M/3 of the proviverrine hyaenodontid (Creodonta: Mammalia)Paratritemnodon indicus, which was hitherto known by heavily worn P/3-M/3, are described from the uppermost Subathu Formation (Middle Eocene) of the Kalakot-Metka-Mohgala area, Rajauri District, Jammu and Kashmir.P. indicus is most closely related to the North American Early Eocene proviverrine,Tritemnodon; its relationships withProdissopsalis from the Middle Eocene of Europe andPropterodon from the Middle to Late Eocene of China are also close. In the Kalakot Eocene vertebrate community of the Subathu vertebrate bioprovince,P. indicus is the only land dwelling carnivorous mammal against more than 20 species of herbivorous mammals, a majority of which are larger than it. This is viewed as an imbalance in trie community and it reflects an immaturity of the ecosystem supporting the fauna. The scarcity of carnivore remains in the Kalakot mammalian fauna is not an artifact.     

Biostratigraphie et Poissons fossilesde la formation de l'Argile de Boom (Oligocène moyen du Bassin Belge)

The Clay of Boom, Rupelian (R₂c of BelgianGeological Map) was sampled in five quarries of the type area (Sint-Niklaas, Steendorp, Schelle, Terhagen, Kruibeke) for otoliths and other fish remains.At the moment 65 species are known from this unit, of which 31 are represented in our samples by otoliths or teeth. The fish fauna of the Clay of Boom is essentially a marine fauna suggesting that the clay was deposited in a calm, rather deep part of the continental shelf.The Elasmobranch fauna has no biostratigraphic value, although 5 new species were identified: Pristiophorus rupeliensis, Raja casieri, Raja cecilae, Raja heinzelini and Raja terhagenesis.The Teleostean fauna is dominated by Gadidae.a typical Northern Atlantic group, and includes one species new to science «genus Gadidarum ensiformis. The dominance of Gadids reflects the progressive replacement of the Indo-Pacific fauna existing during the Eocene, by a more Atlantic one. Some of the 69 lithological subunits recognized on lithological features, are also characterized by different otolith associations and can be correlated in the different quarries sampled. Some of these (49, 41 and 35) are further more characterized by a high frequency of otoliths; therefore these can probably be localised in borings and used for correlation. The Teleostean - otoliths permit a local biozonation of the Clay of Boom. The upper part of the clay is limited below by bed 38 and characterized by the association Argentina parvula - «genus Gadidarum parvus (typical form), the middle part by «genus Gadidarum parvus (thick-set-form) and the lower part of which bed 30 forms the top is characterized by the association Raniceps tuberculosus - Trisopterus elegans. In this part of the clay otoliths are scarce.     

A new sawshark,Pristiophorus laevis,from the Eocene of Antarctica with comments on Pristiophorus lanceolatus

The highly fossiliferous Eocene deposits of the Antarctic Peninsula are among the most productive sites for fossil remains in the Southern Hemisphere and offer rare insights into high-latitude faunas during the Palaeogene. Chondrichthyans, which are represented by abundant isolated remains, seemingly dominate the marine assemblages. Eocene Antarctic sawsharks have only been known from few isolated rostral spines up to now, that were assigned to Pristiophorus lanceolatus. Here, we present the first oral teeth of a sawshark from the Eocene of Seymour Island and a re-evaluation of previously described Pristiophorus remains from Gondwana consisting exclusively of rostral spines. The holotype of Pristiophorus lanceolatus represents a single, abraded and insufficiently illustrated spine from the Oligocene of New Zealand. All other Cenozoic rostral spines assigned to this species are morphologically very indistinct and closely resemble those of living taxa. Consequently, we regard this species as dubious and introduce a new species, Pristiophorus laevis, based on oral teeth. The combination of dental characteristics of the new species makes it unique compared to all other described species based on oral teeth. Rostral spines from the Eocene of Seymour Island are assigned to this new species whereas those from other Cenozoic Gondwana localities remain ambiguous.

LSID urn:lsid:zoobank.org:pub:7177A373-527B-4315-85F6-25180DB5E087     

A new species of Platydracus Thomson, 1858 (Coleoptera,Staphylinidae, Staphylininae) from the upper Eocene Florissant beds,Colorado, USA

Platydracus breviantennatus n. sp., is described and figured based on an impression fossil from the upper Eocene Florissant beds of Colorado, the United States. Based on the large and densely setose body, relatively small eyes, and robust mandibles (right mandible seemingly with one simple preapical tooth), tibiae and antennae, the new species is placed in the modern genus Platydracus Thomson, 1858. It differs from other species of Platydracus by its large body, small head, and distinctly short antennae.     

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.     

First record of Entelodontidae (Mammalia,Artiodactyla) from the late Eocene of Southeast Asia

We describe two entelodontid upper premolars that were recovered from the late Eocene of the Krabi coal mine in southern Thailand. The size and morphology of the material suggest that it can be referred to Entelodon aff. E. gobiensis, a species known from the late Eocene to the early Oligocene of northern Asia and southern China. The Thai material documents for the first time the southernmost occurrence of entelodontids in Asia during the Paleogene and also suggests that Eocene Southeast Asian mammal localities might potentially yield further entelodontid remains mostly associated with selenodont ungulates.     

Late Miocene Carnivora from Chad: Herpestidae,Viverridae and small-sized Felidae

The small-sized feliforms (Viverridae, Herpestidae, Felidae) from the Late Miocene fossiliferous area of Toros-Menalla, Chad, are described. The Viverridae are represented by dental and postcranial remains of two species: a new, large-sized viverrid, Sahelictis korei n. gen. n. sp., which is characterized by a more trenchant dentition than in Viverra spp., and an indeterminate species similar in size to Viverra howelli. The Herpestidae are represented by a subcomplete mandible with partial dentition assigned to Herpestes sp., similar in size to the smallest individuals of the extant Herpestes naso and H. ichneumon. Felids are known from two, possibly three small-sized species. Fragmentary dental and postcranial remains indicate the presence of one or two species of the size of the golden cat (Profelis aurata). A partial skeleton of a wildcat-sized species assigned to Felis sp. is also described; this record is by far the earliest record for the genus in Africa.     

Footprints of Giant Birds from the Upper Eocene of the Paris Basin: An Ichnological Enigma

In 1859 the French geologist Jules Desnoyers reported the discovery of vertebrate footprints in the Late Eocene gypsum of the Paris region. Although they attracted some attention at the time, those footprints were never illustrated or described in detail, and the present whereabouts of the specimens seem to be unknown. Several types of footprints were referable to animals known by skeletal elements from the gypsum, but some were not. Among the latter were tridactyl footprints of very large birds, which Desnoyers tentatively attributed to the giant ground bird Gastornis, which had been discovered in the Lower Eocene of the Paris region a few years earlier. Gastornithids are now known from the Paleocene to the Middle Eocene, but no skeletal remains of giant birds have yet been found in the Upper Eocene of Europe. The tracks of giant birds from the gypsum of the Paris region are thus an example of fossil footprints without known osteological counterparts, and the identity of the trackmakers remains an enigma.     

The late Eocene hyracodontid perissodactyl Ardynia from Saint Jacques,Inner Mongolia,China and its implications for the potential Eocene–Oligocene boundary

Saint Jacques is a classic Oligocene fossil locality in China, which was first investigated by P. Teilhard de Chardin and E. Licent in 1923. All the fossil mammals previously reported from the area are Oligocene in age. Here we report some new material of the hyracodontid Ardynia from two lower horizons in the Saint Jacques area. These new specimens represent two species: Ardynia praecoxMatthew and Granger, 1923 and A. ordosensis n. sp. A right lower jaw of Ardynia praecox with complete i1-c shows that i2 is the largest. The new species, Ardynia ordosensis, is characterized by a moderately hypsodont M1 with a pillar-like “crista” and the metaloph connecting the ectoloph distal to the metacone, characters that are more similar to those of A. altidentata than to any other species in Ardynia. The stratum bearing Ergilian Ardynia extends the deposits at Saint Jacques down to the upper Eocene. In combination with early Oligocene microfossils from the strata overlying the upper Eocene Ardynia-bearing layers, it is reasonable to conclude that the Saint Jacques section may bracket, or likely document, the Eocene–Oligocene transition.     

Studies on Cenozoic crocodiles: 8.Bergisuchus dietrichbergi Kuhn (Sebecosuchia: Bergisuchidae n. fam.) from the Middle Eocene of Germany, some new systematic and biological conclusions

Bergisuchus dietrichbergi Kuhn is redescribed as a member of its own family, Bergisuchidae n. fam., within the Sebecosuchia. At present, two fragmentary specimens ofBergisuchus are known: the holotype ofBergisuchus, a partial rostrum, comes from the lower part of the Middle Eocene of the Messel pit, near Darmstadt; and a recently discovered fragmentary mandible from the middle part of the Middle Eocene of Geiseltal in an opencast coal-mining pit near Halle (Saale).Bergisuchus shows similarities with species ofSebecus which are known from the Paleocene to Miocene from South America. All palaeoecological and biomechanical data support the hypothesis ofBergisuchus as a small, mainly terrestrial crocodile, which did not build its nests near either of the fossil localities from which it has been collected.     

Subfamily Limoniinae Speiser, 1909 (Diptera,Limoniidae) from Baltic Amber (Eocene): The Genus Elephantomyia Osten Sacken, 1860

A revision of the genus Elephantomyia Osten Sacken (Diptera: Limoniidae) from Baltic amber (Eocene) is presented. Four species—E. baltica Alexander, E. brevipalpa Loew, E. longirostris Loew, and E. pulchella Loew—are redescribed and documented with photographs and drawings. In addition, two new species of the genus are described: Elephantomyia bozenae sp. nov., and Elephantomyia irinae sp. nov. All these fossil species are placed within the subgenus Elephantomyia. A key to the extinct species of Elephantomyia is provided, and the genus’ ecological pattern and evolutionary aspects are discussed.     

New discoveries on the giant hedgehog Deinogalerix from the Miocene of Gargano (Apulia,Italy)

During the Miocene, the Abruzzo/Apulia region (Italy), isolated from the continent, was the theatre of the evolution of a vertebrate ecosystem in insular context. During the Late Miocene, the protagonists of this ecosystem called “Mikrotia fauna” show a high endemic speciation level, presenting spectacular giant and dwarf species of mammals and birds. Deinogalerix is one of the most uncommon forms of this peculiar fauna. It is the largest Galericinae that ever lived. From Gargano (Apulia, Italy) deposits (also called “Terre Rosse”), Deinogalerix has been, since its discovery in the 1970s’, the subject of two publications in which various morphotypes were described. Its presence is also attested at Scontrone (Abruzzo, Italy). In 2005, the new Miocene fissure “Mikrotia 013” (M013) was found in the Gargano area at Cava Dell’Erba by a team of the Università degli studi di Torino. The study of the micromammals, and in particular the presence of an archaic form of Mikrotia, attests that M013 is the oldest fissure known to date. All the Deinogalerix remains come from the smallest morphotype ever found. The morphological features (other than size) allow us to describe the new species Deinogalerix masinii. It presents a large variability of dental and mandibular features expressed in mosaic patterns within the material: the robustness of the upper and lower P3 and P4, and the characteristic trigonid of m1 of the genus; the archaic morphology of the maxilla and the mandible; the abrading pattern of the teeth are unique for Deinogalerix. These new remains allow us to update the knowledge of Deinogalerix. The suit of characters testified that D. masinii nov. sp. is an archaic form on the evolutionary trend toward the most derived representatives of the genus. The affinities of Deinogalerix with the Parasorex group are confirmed, but the time of immigration in Gargano remains debated.     

Community Sampling and Integrative Taxonomy Reveal New Species and Host Specificity in the Army Ant-Associated Beetle Genus Tetradonia (Coleoptera,Staphylinidae, Aleocharinae)

Army ant colonies host a diverse community of arthropod symbionts. Among the best-studied symbiont communities are those of Neotropical army ants of the genus Eciton. It is clear, however, that even in these comparatively well studied systems, a large proportion of symbiont biodiversity remains unknown. Even more striking is our lack of knowledge regarding the nature and specificity of these host-symbiont interactions. Here we surveyed the diversity and host specificity of rove beetles of the genus Tetradonia Wasmann, 1894 (Staphylinidae: Aleocharinae). Systematic community sampling of 58 colonies of the six local Eciton species at La Selva Biological Station, Costa Rica, combined with an integrative taxonomic approach, allowed us to uncover species diversity, host specificity, and co-occurrence patterns of symbionts in unprecedented detail. We used an integrative taxonomic approach combining morphological and genetic analyses, to delineate species boundaries. Mitochondrial DNA barcodes were analyzed for 362 Tetradonia specimens, and additional nuclear markers for a subset of 88 specimens. All analyses supported the presence of five Tetradonia species, including two species new to science. Host specificity is highly variable across species, ranging from generalists such as T. laticeps, which parasitizes all six local Eciton species, to specialists such as T. lizonae, which primarily parasitizes a single species, E. hamatum. Here we provide a dichotomous key along with diagnostic molecular characters for identification of Tetradonia species at La Selva Biological Station. By reliably assessing biodiversity and providing tools for species identification, we hope to set the baseline for future studies of the ecological and evolutionary dynamics in these species-rich host-symbiont networks.     

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.     

Lizards of the family Arretosauridae Gilmore, 1943 (Iguanomorpha, Iguania) from the Paleogene of Mongolia

The family Arretosauridae is an enigmatic group of Asian iguanians, which contains one species, Arretosaurus ornatus Gilmore, 1943 from the Late Eocene of Inner Mongolia (China). In this work, the fossil family is classified within the taxon Iguanomorpha (Iguanidae sensu lato) and supplemented with three new forms from the Middle Eocene and Early Oligocene of Mongolia and the previously described Hemishinisaurus latifrons from the Late Eocene of China, which was originally assigned to Anguimorpha (Xenosauridae). New data characterize Arretosauridae as an important element of the Paleogene herpetofauna of Asia.     

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.