Diversité systématique des Didelphidae (Marsupialia) européens tertiaires

From the early Eocene to the middle Miocene,26 species of european Didelphidae are recognized. The diagnosis of 15 new species are stated (Peradectes louisi, russelli and multigniensis; Amphiperatherium brabantense, bourdellense, gothei, maximum, bastbergense and fontense; Peratherium matronense, sudrei, bretouense, lavergnense, monspeliense and perrierense). For the first time, the sub-family of Didelphinae is divided in two new tribes (Peradectini and Didelphini). In Europe, the story of this sub-family is briefly stated.     

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).     

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.     

Using extant patterns of dental variation to identify species in the primate fossil record: a case study of middle Eocene <Emphasis Type="Italic">Omomys</Emphasis> from the Bridger Basin,southwestern Wyoming

Patterns of extant primate dental variation provide important data for interpreting taxonomic boundaries in fossil forms. Here I use dental data from several well-known living primates (as well as data from selected Eocene forms) to evaluate dental variation in Middle Eocene Omomys, the first North American fossil primate identified by paleontologists. Measurements were collected from a sample of 148 omomyid dental specimens recovered from Bridger B localities in the Bridger Basin, Wyoming. Most of these specimens have not previously been described. Nonmetric traits were also scored for this sample. Lower molar coefficients of variation range from 4.01 for M₂ length (n = 80) to 6.73 for M₃ talonid width (n = 57). All of the nonmetric traits scored exhibit less than 100% presence in the overall sample, including traits previously described as representative of Omomys (e.g., P₄ metaconids present in 91%, n = 55; M² pericones present in 80%, n = 15). Dental traits also vary in a set of spatially restricted localities from the same fossil horizon and in a separate, single fossil locality (DMNH 868, P₄ metaconids present in 67%, n = 6). An increasing frequency in several premolar traits across time in these more restricted samples suggests an anagenetic change in Bridger B Omomys. However, this degree of morphological variability is consistent with that seen in extant primate species from single locations. Metric variation in this sample is comparable to that seen in other Eocene primates, such as new data presented here for the omomyid Arapahovius gazini from the Washakie Basin, southern Wyoming. Omomys metric variation is also comparable to that found in several samples of well-known extant primates from single localities (e.g., ring-tailed lemurs and gray–brown mouse lemurs). These metric data also correspond to the patterns of variability described in previously published studies of Omomys carteri. In sum, a single species interpretation (O. carteri) for this new Bridger B Omomys sample from southern Wyoming is affirmed, and this study illustrates the usefulness of dental data from extant primates for evaluating primate fossil samples.     

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.     

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.     

Description of a new species of Gastornis (Aves,Gastornithiformes) from the early Eocene of La Borie,southwestern France

The early Eocene locality of La Borie is located near the village of Saint-Papoul, in southwestern France. It consists of clay deposits that have yielded numerous vertebrate fossils, including remains of the giant flightless bird Gastornis. These remains were initially attributed to the species G. parisiensis, which is otherwise recorded from the late Paleocene and earliest Eocene of the North Sea Basin. New fossil birds collected in the La Borie clay pit in 2018 include an almost complete mandible of Gastornis. We describe a new species of Gastornis based on this mandible and we show that the previously described remains from La Borie must be assigned to this new species. The new species differs from other species of Gastornis in the morphology of the mandible, maxilla and quadrate. The morphological diversity of the genus Gastornis, which existed in Europe for at least 17 million years, is emphasized.LSID of publication: urn:lsid:zoobank.org:pub:10E7938B-C972-4127-94DC-169D35977B11.     

Phylogeny of the dragonfly genus Sympetrum (Odonata: Libellulidae)

The libellulid dragonfly genus Sympetrum has been recognized since 1833, but lacks any morphological synapomorphies to unite the taxon. Previous researchers have disagreed over which species belong in Sympetrum, bringing the monophyly of the genus into question. We use DNA sequence data from 6 genetic loci (16S, tRNA-valine, 12S, elongation factor 1 alpha, cytochrome oxidase subunit I, and the second internal transcribed spacer region) and 25 morphological characters (mainly genitalic) to test the monophyly of Sympetrum with Bayesian inference and maximum likelihood analyses. Under Bayesian inference, all Sympetrum species included in this study form a clade, which also contains the Hawaiian monotypic genus Nesogonia, often considered a close relative of Sympetrum. Phylogenetic analyses also reveal at least six strongly supported clades (treated as species groups) within Sympetrum, but relationships between these species groups remain unresolved or unsupported. Although the relationships between Sympetrum species groups remain unresolved, several species groups include taxa from multiple biogeographic regions/continents, and the species group sister to the rest of Sympetrum contains migratory species from the New World and Africa. This pattern suggests a complex biogeographic history in Sympetrum shaped by vicariance and dispersal. Preliminary estimates of the divergence dates of Sympetrum species groups outline a rapid radiation of the groups approximately 32-38 million years ago, possibly influenced by cooling and drying climates of the late Eocene and early Oligocene.     

A goose-sized anseriform bird from the late Oligocene of France: the youngest record and largest species of Romainvilliinae

We describe a new anseriform bird from the late Oligocene of Saint-André, Marseille, in southern France. Saintandrea chenoides, gen. et sp. nov. is the first avian species reported from the locality, which is well known for its mammalian fossils. The new species belongs to the extinct Romainvilliinae and represents the latest occurrence of the taxon, which was before only known from the late Eocene and early Oligocene of Europe. S. chenoides is also the largest species of Romainvilliinae and increases the known morphological diversity of the taxon. The identification of a goose-sized representative of the Romainvilliinae in the late Oligocene of Europe raises the possibility that some of the large late Paleogene or early Neogene Anseriformes with uncertain phylogenetic affinities also belong to this taxon.     

The first French tragulid skull (Mammalia,Ruminantia, Tragulidae) and associated tragulid remains from the Middle Miocene of Contres (Loir-et-Cher,France)

The Faluns Auger quarry (Contres, France) is famous for its Langhian mammal fauna (MN5) and tragulid remains have been identified as Dorcatherium sp. New excavations provided a fragmented skull of a tragulid, dental remains, and a metapodium which we describe here. Based on morphological and morphometrical characters, these specimens are attributed to Dorcatherium crassum. CT-scans provide insight in the petrosal bone and bony labyrinth. We prove intra-population variability of the p4 and intra-specific variability of the bony labyrinth. Nevertheless, we can demonstrate that the bony labyrinth is useful for systematics. Thus, we confirm the presence of D. crassum in the Pontlevoy–Thenay and Savigné-sur-Lathan Faluns Basins as reported already prior to our study. Yet, we cannot confirm evidence for Dorcatherium naui. D. naui specimens reported from these basins may belong to another species. D. naui should be absent from the Faluns before MN7, such as in the rest of Europe.     

A Review of the Fossil Record of the Genus Itea (Iteaceae, Saxifragales) with Comments on its Historical Biogeography

Itea is a genus of about 20 species of trees and shrubs that are today native to southeastern North America, eastern Asia, and eastern Africa. In this paper, I review the fossil record of Itea, which is based on four types of fossils: diporate, psilate pollen attributed to Itea or the dispersed pollen genus Iteapollis; carpofossils representing fruits and seeds attributed to Itea europaea; flowers preserved in amber and assigned to Adenanthemum iteoides; and leaf impressions attributed to Itea. The distributions of these fossils indicate that Itea was present in western North America from the early Eocene to Miocene, in eastern North America beginning no later than the early Miocene, and in western Eurasia from the late Eocene to Pliocene. Only one datapoint is known from eastern Asia; it is early Miocene in age. Based on the fossil record, it can be inferred that Itea crossed between continents over both the Bering Land Bridge and North American Land Bridge, and that it reached Africa from Europe via Anatolia. Thus, it is predicted that the sole extant North American species, I. virginica, may be most closely related to the sole extant African species, I. rhamnoides. The potential application of Itea fossils to calibrating phylogenetic trees generated from molecular sequence data is also discussed.     

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.     

EOCENE FLORAL EVIDENCE OF LAURACEAE: CORROBORATION OF THE NORTH AMERICAN MEGAFOSSIL RECORD

Numerous megafossils of Lauraceae have been reported from the early Tertiary of North America, but the subfamilial affinities are usually not well understood due to the great morphological variability found in extant taxa. The flowers of Androglandula tennessensis gen. et sp. nov. Taylor, from the Middle Eocene Claiborne Formation, are six-parted, pedicellate, bracteate, and have stamens with paired basal staminal glands. The flowers have ethereal oil cells and paracytic stomates throughout. The fossil species has affinities with the subtribe Cinnamomineae, and this supports suggestions that the Middle Eocene climate of the southeastern U.S. was subtropical. The existence of this fossil, and reports of the subtribe from the Eocene of Europe, indicate a South American-North American-European-southeast Asian paleodistribution suggesting that extinction in North America and Europe was the cause of the tribe's current disjunct distribution.     

Size analyses of the coccolith species Biscutum constans and Watznaueria barnesiae from the Late Albian “Niveau Breistroffer” (SE France): taxonomic and palaeoecological implications

The size variability of the coccolith species Biscutum constans and Watznaueria barnesiae has been studied in 50 samples of the Late Albian “Niveau Breistroffer” black shales (Col de Palluel section, SE France). For each species, length and width of the total coccolith and the central unit have been measured in 60 specimens per sample. In addition, ellipticity and central unit length/total coccolith length ratio have been calculated. This study aims to improve our understanding of the taxonomic concepts of B. constans and W. barnesiae, and to test whether the size changes correspond to palaeoceanographic changes interpreted from other proxies. Two morphotypes (varieties) were differentiated for each of the two taxa studied. B. constans includes morphotypes with a narrow (B. constans var. constans) and a wide central unit (B. constans var. ellipticum). Between these two morphotypes no significant size differentiation has been observed. They seem to represent end members of a size continuum, relating to both the total coccolith- and central unit size distribution. The two morphotypes of W. barnesiae differ only in the absence (W. barnesiae var. barnesiae) and the presence of a narrow central opening (W. barnesiae var. fossacincta). No significant size differences have been observed between these two morphotypes, and both show similar distributions of the measured characteristics. We recommend that the studied morphotypes of both B. constans and W. barnesiae should not be assigned to different morphospecies. Throughout the studied black shale succession the measured parameters show mostly statistically insignificant, short-term fluctuations, but significant long-term trends have been observed for B. constans. These show a trend to smaller forms with a narrower central unit in the upper part of the succession. This change coincides with a cooling trend, as indicated by a nannofossil based temperature index. Therefore the two morphotypes of B. constans are interpreted to represent ecophenotypic varieties rather than two different morphospecies. No clear relationship has been recognized between the size of W. barnesiae and the palaeoenvironmental conditions.     

Linking Seed Photosynthesis and Evolution of the Australian and Mediterranean Seagrass Genus Posidonia

Recent findings have shown that photosynthesis in the skin of the seed of Posidonia oceanica enhances seedling growth. The seagrass genus Posidonia is found only in two distant parts of the world, the Mediterranean Sea and southern Australia. This fact led us to question whether the acquisition of this novel mechanism in the evolution of this seagrass was a pre-adaptation prior to geological isolation of the Mediterranean from Tethys Sea in the Eocene. Photosynthetic activity in seeds of Australian species of Posidonia is still unknown. This study shows oxygen production and respiration rates, and maximum PSII photochemical efficiency (Fv : Fm) in seeds of two Australian Posidonia species (P. australis and P. sinuosa), and compares these with previous results for P. oceanica. Results showed relatively high oxygen production and respiratory rates in all three species but with significant differences among them, suggesting the existence of an adaptive mechanism to compensate for the relatively high oxygen demands of the seeds. In all cases maximal photochemical efficiency of photosystem II rates reached similar values. The existence of photosynthetic activity in the seeds of all three species implicates that it was an ability probably acquired from a common ancestor during the Late Eocene, when this adaptive strategy could have helped Posidonia species to survive in nutrient-poor temperate seas. This study sheds new light on some aspects of the evolution of marine plants and represents an important contribution to global knowledge of the paleogeographic patterns of seagrass distribution.     

Le genre Leguminocythereis (Ostracodes) dans le Paléogène nord-aquitain: Espèces et populations; histoire évolutive locale

The genus Leguminocythereis, well represented inWestern Europe, is followed in North-Aquitaine (France) throughout the Eocene and at the basal Oligocene.Four species are described: L. striatopunctata, L.inflata, L. pertusa, L. aff. scrobiculata. The first two ones are polymorphic.The genus has suffered severely by the gradual setting of Eocene/Oligocene climatic crisis. Effectives grow less, specific populations structures are simplified; L. pertusa and L. aff. scrobiculata only are known at the basal Oligocene.By their locally irreversible distribution in time, themorphs appear as good stratigraphic indicators. Some of them are closely linked to special palaeoenvironments and are their markers.A comparison between the variances of some characterssuggests that, when they are isolated (out of the polymorphic structure) or/and in less favourable living conditions, the morphs develop an adaptative strategy by increasing of their genetic variability.Observed data are relevant of microevolution.     

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.