An updated review of the middle Eocene avifauna from the Geiseltal (Germany), with comments on the unusual taphonomy of some bird remains

The brown coal mines of the Geiseltal in Germany were among the most important middle Eocene fossil localities and constitute the reference sites for the Geiseltalian stage of the European Land Mammal Mega Zones. Here, an updated review of the Geiseltal avifauna is given. Thirteen species are represented by diagnostic bones and can be referred to avian higher-level taxa, but various indeterminable albeit distinct fossils indicate a higher diversity of the Geiseltal avifauna. The majority of the Geiseltal fossils belong to taxa, which are well known from early to mid-Ypresian fossil sites of Europe and North America. Some of the species from the Geiseltal are larger than their early Eocene relatives, but it remains elusive whether this indicates an evolutionary size increase in some avian lineages or reflects different palaeohabitats of the involved fossil sites. The Geiseltal avifauna exhibits some taphonomic peculiarities that have not yet been addressed. Most bird fossils consist of isolated bones or partial skeletons and here it is hypothesized that at least some of the fragmentary specimens represent feeding remains of crocodilians. For the first time, medullary bone, which is indicative of breeding females, is reported for birds from the Geiseltal. The comparative abundance of the coliiform species Eoglaucidium pallas is remarkable, and either this species was extremely abundant in the Geiseltal ecosystem or it had ecological preferences, which favored its preservation.     

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.     

New hyaenodonta (Mammalia) from the middle Eocene of Myanmar

The middle Eocene Pondaung Formation in Myanmar has yielded a rich mammalian fauna including several Primate taxa. Hyaenodonta are known by the genera Kyawdawia, Yarshea, Orienspterodon, and two other indeterminate taxa. We describe here new material of Kyawdawia, including some morphological details, a new species of the hypercarnivorous genus Propterodon and an indeterminate species, different from those described earlier in Myanmar, and characterized by a reduction of m3 and would belong to a third lineage with the same evolutionary trend as Galecyon and the Limnocyoninae. The hyainailourines (Orienspterodon) and hyaenodontines (Propterodon) are recorded for the first time in Southeast Asia and these subfamilies appeared in quasi the same time in Europe illustrating probably a profound change in the carnivorous fauna among Laurasia.     

Taxonomic description of in situ bee pollen from the middle Eocene of Germany

The middle Eocene Messel and Eckfeld localities are renowned for their excellently preserved faunas and diverse floras. Here we describe for the first time pollen from insect-pollinated plants found in situ on well-preserved ancient bees using light and scanning electron microscopy. There have been 140 pollen types reported from Messel and 162 pollen types from Eckfeld. Here we document 23 pollen types, six from Messel and 18 from Eckfeld (one is shared). The taxa reported here are all pollinated by insects and mostly not recovered in the previously studied dispersed fossil pollen records. Typically, a single or two pollen types are found on each fossil bee specimen, the maximum number of distinct pollen types on a single individual is five. Only five of the 23 pollen types obtained are angiosperms of unknown affinity, the remainder cover a broad taxonomic range of angiosperm trees and include members of several major clades: monocots (1 pollen type), fabids (7), malvids (4), asterids (5) and other core eudicots (1). Seven types each can be assigned to individual genera or infrafamilial clades. Since bees visit only flowers in the relative vicinity of their habitat, the recovered pollen provides a unique insight into the autochthonous palaeo-flora. The coexistence of taxa such as Decodon, Elaeocarpus, Mortoniodendron and other Tilioideae, Mastixoideae, Olax, Pouteria and Nyssa confirms current views that diverse, thermophilic forests thrived at the Messel and Eckfeld localities, probably under a warm subtropical, fully humid climate. Our study calls for increased attention to pollen found in situ on pollen-harvesting insects such as bees, which can provide new insights on insect-pollinated plants and complement even detailed palaeo-palynological knowledge obtained mostly from pollen of wind-pollinated plants in the dispersed pollen record of sediments. In the case of Elaeocarpus, Mortoniodendron, Olax and Pouteria the pollen collected by the middle Eocene bees represent the earliest unambiguous records of their respective genera.     

Italian Cenozoic crocodilians: taxa, timing and palaeobiogeographic implications

Crocodylian remains are collected in 39 fossil-bearing localities but only in seven localities specimens with reliable taxonomic attributions, at least to genus level have been collected. Three species have been reported from the early Lutetian Purga di Bolca site: Pristichampsus cf. Pristichampsus rollinati, Asiatosuchus sp., Hassiacosuchus sp. (=Allognathosuchus sp.). The three crocodilians discovered at Purga di Bolca have been reported also from Geiseltal and Messel (Middle Eocene, Germany). Bolca at that time was part of a Tethysian archipelago and no mammals have been found there till now. Crocodilians and turtles clearly arrived from the European mainland across a marine water barrier. Among the other fossiliferous localities of Veneto, very interesting is the Monte Zuello site, of late Middle Eocene age, yielding a longirostrine crocodilian, Megadontosuchus arduini, a tomistomine species. Tomistomines are known in contemporaneous sediments of both Europe and Africa, but the European forms Dollosuchus and Kentisuchus seem the closest taxa. Remains of Oligocene age have been collected in Veneto and Liguria, but the fossils discovered in the second region are teeth or fragmented bones. The fossil crocodilians of Monteviale (Veneto), of Early Oligocene age, have been assigned to two species but they have been recently all identified as Diplocynodon ratelii, known from several European sites of Late Eocene, Oligocene and Miocene age. This species arrived in the Monteviale area from the European mainland across a narrow sea. Several crocodilian fossils of Miocene age are very fragmentary or represented by isolated teeth. In the Middle and Late Miocene of Sardinia, a well-established species, Tomistoma calaritanum is present. Remains of Tomistoma of the same age have been reported in some localities in Tuscany, Apulia, Sicily and Malta. In the Mediterranean area, the genus is known from European and African sites (of older age). The colonisation of Europe by this genus is the result of a dispersion from Africa (or less probably from Asia). During Late Miocene Sardinia and Tuscany belong to the same palaeobioprovince characterized by the Oreopithecus-Maremmia fauna. In Tuscany, a crocodilian identified as Crocodylus bambolii is present in the late Miocene site of Monte Bamboli. If the generic attribution of this form is correct, its ancestors must have arrived from Africa. Another fossil assemblage of Late Miocene age characterizes the Apulia-Abruzzi palaeobioprovince (Hoplitomeryx-Microtia fauna) and testifies complete isolation between the two palaeobioprovinces. In this last area, remains of Crocodylus sp. have been collected in coastal sandstones at Scontrone (Abruzzi) and in several fissure fillings of Gargano of slightly younger age. The ancestors of this species arrived from Africa while no African elements are present among the mammalian fauna. The dispersion of the genus Crocodylus in the Italian palaeoislands may have taken place once, with allopatric differentiation of the two populations (Tuscany-Sardinia and Apulia-Abruzzi) or twice with independent colonisation of each area.     

Herpetofaunas from the Upper Paleocene and Lower Eocene of Morocco

Two Upper Paleocene and one Lower Eocene localities from Morocco (Ouarzazate basin) have yielded terrestrial assemblages that stand among the rare herpetofaunas from the Paleogene of the African Plate. The collections include one of the rare frogs and the only lizards known from the Paleogene of Africa. One of the two Upper Paleocene localities, Adrar Mgorn 1, has produced an indeterminate anuran and the most diverse assemblage of squamates from the Mesozoic and Cenozoic of Africa. It has yielded the earliest known scolecophidian snake and the earliest Gekkonidae, amphisbaenians, Tropidophiidae, and perhaps Boidae from Africa. Moreover, a specimen represents either the last sphenodontian or the earliest acrodontan lizard from this continent. One of the amphisbaenians represents a very distinct new taxon, Todrasaurus gheerbranti gen. and sp. nov. Indeterminate scincomorphans, lacertilians, Madtsoidae, and Aniliidae are also present. The fauna from the Lower Eocene is less diverse than that from the Upper Paleocene, but some taxa are common to both levels. Contrary to nearly all other Paleogene herpetofaunas from the African Plate, these Paleocene and Eocene assemblages include taxa that were terrestrial, not aquatic.     

The first New World occurrence of the Eocene birdPlesiocathartes (Aves: ?Leptosomidae)

Two new species of the Eocene avian taxonPlesiocathartes (Aves: ?Leptosomidae) are described. They are similar in size and overall appearance to other members of the genus, as well as to the extant Madagascan cuckoo-rollerLeptosomus discolor. So far, putative fossil Leptosomidae have been found only in the Eocene European de-posits of Messel, Geiseltal, Quercy, and London Clay. Both new species are from the Lower Eocene Green River Formation (USA) and thus the first New World representatives of this taxon.     

Crocodylian assemblage from the middle Eocene Ikovo locality (Lugansk Province,Ukraine), with a discussion of the fossil record and geographic origins of crocodyliform fauna in the Paleogene of Europe

The known fossil record of crocodyliforms in Europe during the Paleogene is significantly biased, in that the fauna of Western Europe is far better sampled and understood compared to that of Eastern Europe. We describe in detail all known crocodyliform remains from the middle Eocene (Lutetian) Ikovo locality in Ukraine. We conclude that at least two taxa were present: a moderate to large-sized Tomistominae indet. similar to the basalmost known tomistomines, and the small-sized basal alligatoroid cf. Diplocynodon sp. Despite its scarcity, this is the first basal alligatoroid material reported from Eastern Europe (as part of post-Soviet countries) and the easternmost record of diplocynodontines in Europe so far. An allegedly freshwater cf. Diplocynodon sp. contributes a rare faunal element to the vertebrate assemblage of the Ikovo locality, otherwise dominated by resident or facultative marine taxa. The fossil record and historical paleobiogeography of crocodyliforms from the Paleocene and Eocene of Europe are reviewed. As it has been already known, the middle Eocene fauna of crocodyliforms proves to be taxonomically diverse and complex. Its constituent lineages geographically originated in Asia or North America (Diplocynodontinae, Asiatosuchus-like crocodyloids, Planocraniidae), North America (derived alligatorines), Africa (Tomistominae), and Gondwana (ziphodont mesoeucrocodylians Iberosuchus and Bergisuchus), with possible subsequent speciation in Europe. We propose a novel hypothesis of Asian origins of European diplocynodontines, which will be explicitly tested in future studies. The revealed similarities between crocodylians and turtles from the Ikovo locality and those from Western Europe support the presence of a single Pan-European biogeographical zone during the middle Eocene, distinct from that of Asia.     

The taxonomy,chronostratigraphy and paleobiogeography of glyptosaurine lizards (Glyptosaurinae,Anguidae)

Glyptosaurine lizards (Glyptosaurinae, Anguidae) are an extinct group of heavily armored lizards known from North America, Europe and Asia. Glyptosaurine lizards, taxa that possess fully developed tuberculated dermal armor, appear to have been established in North America by late early Puercan time (To3). “Proxestops,” a taxon distinguished by a combination of vermiculate and tuberculated osteoderm sculpturing, is considered to be a non-glyptosaurine, a sister taxon of the Glyptosaurinae. Known from only fragmentary remains, its wide chronostratigraphic distribution suggests that “Proxestops” is a form genus that, in all probability, represents more than one taxon, that ranges from the middle Paleocene to the early Eocene of North America. Moreover, the taxa Odaxosaurus piger, Parodaxosaurus sanjuanensis and “Proxestops” are best considered “proto-glyptosaurines”. “Melanosaurins” and glyptosaurins were well-established by the early Eocene, especially in North America, and are here documented by their type species and chronostratigraphic levels. Both tribes are present in Europe (MP7), too, but the record is not as estensive as that of North America. The North American taxon Gaultia silvaticus (Wa0) is transitional between a “melanosaurin” and glyptosaurin. Because it lacks the well-defined hexagonal osteoderms that characterize the Glyptosaurini, it is removed from that group and considered to be a “melanosaurin”. The “melanosaurin” taxon “Xestops” savagei (Wa4–Wa6) cannot be referred to Xestops (Br2) based on non-corresponding elements and because superficial similarity does not justify assignment to this taxon. Arpadosaurus sepulchralis (Wa6?), whose holotype is a fragmentary right frontal, is considered a subjective junior synonym of A. gazinorum, based on minor differences in the epidermal scale pattern that probably represent individual variation. “Glyptosaurus” agmodon (Wa6?), based on a partial right maxilla, cannot be referred to Glyptosaurus (sensu stricto), and the material upon which this taxon is based bears strong resemblance to material identified as cf. “?Paraglyptosaurus” yatkolai (Wa5–Wa6). “Glyptosaurus” rhodinos (Wa5) is based on an incomplete parietal, and its reference to Glyptosaurus is considered problematic. Eoglyptosaurus donohoei (Wa7) is probably valid and is re-established here. Glyptosaurus (sensu stricto) is known solely from the middle Eocene (Br2) by G. sylvestris. Dimetoposaurus wyomingensis (Br3) is removed from Xestops vagans because its synonymy was based on superficial similarities. Helodermoides tuberculatus, the largest and last glyptosaurin (Ch3), is restricted to the Chadronian of North America. Only the “melanosaurin” Peltosaurus granulosus (Or2–Or3), which includes the species P. abbotti, seems to have crossed the Eocene-Oligocene boundary, and appears to be largely restricted to the Orellan, but extended into the Arikareean. European glyptosaurines are also represented by both glyptosaurins and “melanosaurins” early in the Eocene (MP7). Placosauriops-like “melanosaurins” are known from Dormaal (MP7), and the glyptosaurin taxon?Placosaurus ragei occurs at the same level. “Placosauriops abderhaldeni” has been identified from the Grube Messel (MP11), but this assignment remains dubious because the species has not been adequately diagnosed, and the holotype species is from the Geiseltal (MP13), which is some 4.5 million years younger. Placosauriops weigelti (MP13) is the only valid species of this genus. Paraxestops stehlini (MP14) is not referable to the North American taxon Xestops, and its relationship to Placosauriops has not been studied. The late Eocene glyptosaurins Placosaurus estesi (MP17) and P. rugosus (MP18) are the last glyptosaurines known from Europe and appear to have gone extinct at the Eocene-Oligocene boundary, casulties perhaps of the “Grande Coupure”. Asian glyptosaurines are known solely from one species, Stenoplacosaurus mongoliensis, from the middle Eocene (Sharamurunian) of China. Glyptosaurines most likely originated in North America, diversified by late Paleocene time, and rapidly spread across the North Atlantic into Europe by the early Eocene. Both “melanosaurins” and glyptosaurins took a foothold in Europe by the early Neustrian, but the glyptosaurins, aside from one occurrence (Dormaal, MP7), were conspicuously absent for most of Neustrian through early Robiacian time. In North America, glyptosaurins diversified during the early and middle Eocene, while in Europe small “melanosaurins” were a prominent part of the paleoherpetofauna, and glyptosaurins are unknown for most of the Neustrian through the Geiseltalian, in both the fossilferous Lagerstätten of Messel and Geiseltal. Stenoplacosaurus is the only known glyptosaurin glyptosaurine from Asia, and its abrupt appearance during the late Eocene suggests the possiblity of a Beringian dispersal from North America into Asia.     

A late early to early middle Eocene mammal assemblage from Bayan Ulan (Inner Mongolia,China): Implication for the reassessment of the Arshantan Asian Land Mammal Age

Paleogene mammal localities of North China are particularly well represented in the Erlian Basin, Inner Mongolia. Among them, the locality of Bayan Ulan is most famous for its late Paleocene Gashatan fauna. However, the younger Arshantan fauna of the same site is not well known, since no extensive study has been done so far. Here, we present a small mammal assemblage based on dental and tarsal material from a new Arshantan collection retrieved from the red beds of the late early to early middle Eocene Arshanto Formation at Bayan Ulan. It consists of at least six different taxa: the basal lagomorph Dawsonolagus antiquus, the large pantodont Pantolambdodon sp., the tapiroid Schlosseria magister, and the rhinocerotoids Hyrachyus crista and Rhodopagus guoi nov. sp. The assemblage is dominated by perissodactyls, especially Lophialetidae and Hyracodontidae. For the first time, p4-m1 of Dawsonolagus antiquus, tarsal material from Pantolambdodon sp., and lower dentition and tarsals of Hyrachyus crista are described and illustrated. Unlike other described Arshantan faunas, the Bayan Ulan Arshantan mammal assemblage has been collected exclusively from a single locality, which contributes to the reassessment of the misunderstood Arshantan Asian Land Mammal Age.     

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.     

Die erste Wirbeltierfauna aus der miozänen Braunkohle der Niederrheinischen Bucht (Ville-Schichten, Tagebau Hambach)

This paper describes the first Miocene vertebrate fauna from the Ville Series of the Lower Rhine Embayment in western Germany. The fossil material was discovered in the vast Hambach open cast lignite mine ofRheinbraun. The numerous remains, mostly isolated teeth and disarticulated bones, were accumulated in a channel-fill within the main lignite seam. The fauna consists of chondrichthyans, teleosteans, amphibians, reptiles, and mammals which inhabited brackish-estuarine, lacustrine-fluviatile, and terrestrial environments. This is the first fauna to provide an insight into the diversity of vertebrates which lived in the forested peat swamps of the Rhenish Tertiary. Based on the rich association of more than 70 mammalian taxa, including about 30 rodent species, this late Orleanian fauna can be correlated with the upper part of the Mammalian Neogene biozone MN 5. Therefore the middle part of the main lignite seam (horizon 6C of the local lithostratigraphy) can be placed in the early middle Miocene. This means an age of 15.2-16.0 Ma (Langhian or Reinbekian of the stratigraphy of the NW German Tertiary). For some mammalian taxa (Lanthanotherium, Plesiosorex,Miopetaurista, Myoglis, Fahlbuschia, ‘Lartetomys’, Anchitheriomys, Dorcatherium), Hambach 6C is one of the richest European localities. In addition this assemblage contains some very rare mammals (‘Lartetomys’, Anchitheriomys, Orygotherium, Pliopithecus). The Hambach vertebrates are of great palaeobiogeographic importance as they represent the north westernmost outpost of terrestrial Miocene faunas in Europe.     

New notharctine primate fossils from the early Eocene of New Mexico and southern Wyoming and the phylogeny of Notharctinae

Previously undescribed notharctine primate fossils are reported from the early Eocene San Jose Formation, San Juan Basin, New Mexico, and the early Eocene Wasatch Formation, southern Wyoming. These collections include the most complete specimens yet discovered of the poorly known species Copelemur tutus and Copelemur praetutus; the first upper dentitions of Cantius angulatus and Cantius frugivorus from the type area of these taxa; and fossils attributable to two new notharctine species, Copelemur australotutus and Smilodectes gingerichi. These new fossils reveal that current ideas concerning notharctine phylogeny are incorrect. Two major, monophyletic clades are apparent within the subfamily: the tribe Copelemurini, consisting of the genera Copelemur and Smilodectes, and the tribe Notharctini, comprising the genera Cantius, Pelycodus, and Notharctus. Analysis of the paleobiogeographic distribution of the Copelemurini indicates that this clade was limited to more southerly regions of western North America during early Eocene time. Northward migration of more tropical habitats during the late Wasatchian and early Bridgerian in western North America, associated with an overall climatic warming trend through the early and middle Eocene, appears to have allowed several mammalian taxa, including Smilodectes, to extend their ranges northward during this time interval. Such taxa thus possess diachronous distributions and have been partly responsible for the long-standing confusion regarding the biostratigraphic correlation of early Eocene faunas from New Mexico with those from Wyoming. Based on several taxa which are also known from the Wasatchian of Wyoming, the age of the San Jose Formation appears to be middle Wasatchian.     

Middle Eocene Pelagornithidae and Gaviiformes (Aves) from the Ukrainian Paratethys

Abstract: We describe avian remains from Novopskov, a new middle Eocene marine locality in Ukraine. The fossils constitute the most substantial collection of Palaeogene bird bones from Eastern Europe and contribute to a better knowledge of the Paratethyan seabird fauna. Most of the specimens belong to Pelagornithidae (bony‐toothed birds), and two species of very different size can be distinguished. The larger of these is tentatively referred to Dasornis sp., the smaller to Odontopteryx toliapica. The specimens include skeletal elements that were not described for Palaeogene bony‐toothed birds and document previously unknown morphological differences between Palaeogene and Neogene Pelagornithidae. It is argued that the purported crane Eobalearica tugarinovi, from the middle Eocene of Kyrgyzstan, is probably also a bony‐toothed bird. A new genus and species of small Gaviiformes, Colymbiculus udovinchenkoi, is described, which is the earliest fossil record of a loon from Europe, preceding the next oldest specimens by more than 10 myr. The Ukrainian fossils document profound differences between middle Eocene and extant marine avifaunas of Europe, and whereas the middle Eocene Paratethyan avifauna appears to have been similar to that of the North Sea with regard to pelagornithid diversity, the absence of prophaethontids and relative abundance of Gaviiformes may indicate faunistic differences concerning the remaining seabirds.     

New specimens of the Middle Eocene fossil mousebird Selmes absurdipes Peters 1999

Described here are an excellently preserved new specimen of the Middle Eocene mousebird Selmes absurdipes from the type locality Messel (Hessen, Germany) and first records of this species from the Geiseltal (Sachsen-Anhalt, Germany). These specimens allow the recognition of several previously unknown osteological features, and thus more detailed comparisons with other fossil and recent Coliiformes. Selmes absurdipes, which previously has been known only from a single poorly preserved skeleton, most closely resembles the Lower Eocene species Chascacocolius oscitans in its overall morphology. The species shares a derived character with each of the two coliiform families, the extinct Sandcoleidae and the Coliidae, and at present it cannot be assigned to either of these families with certainty.     

Fossil water‐penny beetles (Coleoptera: Psephenidae: Eubrianacinae) from the Eocene of Europe,with remarks on their phylogenetic position and biogeography

Abstract: A species of water‐penny beetles is recorded from larval instars from the Middle Eocene Messel pit fossil site in Germany. This species clearly belongs to the psephenid subfamily Eubrianacinae, but its precise systematic affinities remain unclear. It is the second fossil species of this taxon recorded from Europe. The holotype of the first fossil species, Eubrianax vandeli Bertrand and Laurentiaux, 1963, is lost. The high number of fossil specimens from Messel allowed discussion of stratigraphic and spatial occurrence of the eubrianacine species in the Messel pit, but its larval ontogeny could not be unravelled. Because of the high number of fossils and their wide occurrence, it is inferred that the species from Messel gen. sp. 1 was an autochthonous faunal element of the Eocene Lake Messel, which might indicate that some parts of the former Lake Messel had a shore area with stones. The analysis of the phylogenetic position of both Eocene eubrianacine species showed that their phylogenetic placement cannot be resolved because preservational influences limit the evaluation of characters. The historical biogeography of Psephenidae and Eubrianacinae is analysed and discussed. The fossil record shows that psephenid beetles have fossil members occurring outside their current distribution range, so interpretations of their biogeography based only on extant members can be misleading.     

New carnivoraforms from the early Eocene of Europe and their bearing on the evolution of the Carnivoraformes

Two new mammalian carnivoraform species, Uintacyon hookeri sp. nov. and Quercygale smithi sp. nov., are described from the early Eocene of Europe. U. hookeri sp. nov. is recorded in Mutigny (MP8 + 9, PE IV), Avenay (MP8 + 9, PE V), Brasles, Condé‐en‐Brie (MP8 + 9) and Cuis (MP 10), while Q. smithi sp. nov. comes from Mutigny and Mancy (MP10). Because the two species are not recorded in earliest Eocene localities such as Dormaal and Le Quesnoy (MP7, PE I), it is proposed that they dispersed after the main phase of the Mammal Dispersal Event. U. hookeri sp. nov. supports the existence of terrestrial connections with North America, while Q. smithi sp. nov. implies possible faunal exchange with Asia. This evidence for the evolution of the Carnivoraformes supports: (1) a rapid decrease in their diversity after the Mammal Dispersal Event; and (2) the existence of a mammal turnover event in Europe during the early Eocene. The discovery of a new species of Quercygale, which is generally considered as the closest carnivoraform to the crown‐group Carnivora, shows that the genus had already lost the M₃ by the early Eocene and supports an important, but very poorly known, radiation of the carnivoraforms at least during the earliest early Eocene.     

Elucidating the affinities and habitat of ancient, widespread Cyperaceae: Volkeria messelensis gen. et sp. nov., a fossil mapanioid sedge from the Eocene of Europe

The sedges (family Cyperaceae) are an economically and ecologically important monocot group dating back at least to the Paleocene. While modern genera are mostly unknown before the Oligocene, several extinct taxa are recognized as the earliest sedges. Their affinities have been unclear until now, because they are found as isolated, often abraded fruits or endocarps. Exceptionally preserved sedge fossils from the Middle Eocene of Messel, Germany yield more characters for identification. Fossil cyperacean infructescences with in situ pollen are recognized for the first time and show features of the early-divergent mapanioid sedges. Pollen resembles that of tribe Hypolytreae. Comparisons with extant taxa suggest the closest affinities with Hypolytrum and Mapania. However, the Messel fossils represent a distinct taxon, Volkeria messelensis gen. et sp. nov. Without the additional characters of infructescence and pollen, the Messel fruits would have been placed in the extinct genus Caricoidea, a typical Eocene sedge that was widespread across Eurasia. Similarities of fruit structure suggest that Caricoidea was also a mapanioid sedge. Mapanioid sedges are found today in tropical wet forests and swamps, a distribution suggesting that early sedges occupied a similar habitat, unlike many modern sedges, and were not precursors to open grassland vegetation.     

New chiropterans from the middle Eocene of Shanghuang (Jiangsu Province,Coastal China): new insight into the dawn horseshoe bats (Rhinolophidae) in Asia

Until recently, the fossil record of Paleogene bats in Asia primarily included extinct families (i.e. ‘Eochiroptera’) from the early Eocene of Vastan in India and from the middle‐late Eocene of the Liguanqiao and Yuanqu basins in central China. Here, we describe a new fauna of Chiroptera from the middle Eocene Shanghuang fissure fillings of China. The fauna includes abundant material referred to a new rhinolophid (Protorhinolophus shanghuangensis gen. and sp. n.), one specimen of a possible rhinopomatid and several indeterminate rhinolophoids. This new bat assemblage constitutes the earliest record of extant families of microbats in Asia. Because it lacks representatives of ‘Eochiroptera’, this Shanghuang bat fauna indicates significant turnover in Asian bat communities. The dental pattern of P. shanghuangensis shows a mosaic of primitive and derived features (‘Eochiroptera’ vs Rhinolophidae dental characteristics), suggesting that this taxon occupies a basal position among the Rhinolophidae. Rhinolophids were already well diversified at the end of the late Eocene in Europe. Interestingly, many dental characteristics of Protorhinolophus are also found in a primitive rhinolophoid taxon, Vaylatsia, from the middle Eocene to late Oligocene of Europe, supporting a close relationship between these taxa. These affinities testify to the widespread Eurasian distribution of rhinolophoids during the Eocene and are consistent with a westward dispersal of the group from eastern Asia to Europe owing to the greater antiquity of Protorhinolophus.     

Amphibians and squamates from the middle Eocene of Namibia,with comments on pre-Miocene anurans from Africa

Three middle Eocene localities (Silica North, Silica South, Black Crow) recently discovered in Namibia have produced terrestrial faunas that rank among the few known from the period of insulation of Africa (Aptian-early Miocene). Collectively, the three localities have yielded anuran amphibians (one pipid frog, the earliest assemblage [three taxa] of ranoid frogs in Africa, one indeterminate family) and squamate reptiles (an amphisbaenian ‘lizard’, a snake that likely represents a colubroid, and two indeterminate ‘lizards’). These Eocene faunas suggest that ranoids, colubroids and African pipids are autochthonous to Africa. However, whereas pipids are vicariants inherited from West Gondwana, ranoids and colubroids (if really autochthonous) originated in Africa from unknown stems. Silica North and Silica South correspond to aquatic environments, permanent fresh water being present in the first locality; the environment of Black Crow was drier.