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     

The first species of Sinopa (Hyaenodontida,Mammalia) from outside of North America: implications for the history of the genus in the Eocene of Asia and North America

The first Sinopa species, S. jilinia sp. nov., from outside of North America is described. It comes from the Huadian Formation, locality Gonglangtou, Jilin Province, north‐east China. The new species represents the northernmost and one of the latest and most complete Asian Prototomus‐like hyaenodontidans known. It also represents one of the youngest specimens of Sinopa, because the age of the Huadian Formation is correlated to the later Uintan and only one doubtful citation of North American Sinopa younger than the early Uintan exists. S. jilinia sp. nov. is characterized by having m3 clearly smaller than m1, very strong and extended labial molar cingulids, backward leaning protoconids in all molars and its m3 cristid obliquum joining the postvallid very labially. With S. jilina, Sinopa is the first hyaenodontidan genus known to be present on two continents during the time interval between the earliest Eocene (c. 55.0 Ma) and latest middle Eocene (40 Ma). Its occurrence in the Huadian Formation supports the idea of a faunal exchange between North America and Asia in the early middle Eocene, a hypothesis formerly based mainly on the presence of the omomyid primate Asiomomys in the Huadian Formation, on a small radiation of East Asian trogosine tillodonts and on a couple of perissodactyl genera shared between the middle Eocene of North America and the Irdinmanhan of East Asia. As with the new Sinopa species, these Asian taxa had their closest relatives in North America.     

Origin and evolution of the Pseudorhyncocyonidae,a European Paleogene family of insectivorous placental mammals

Two new species of pseudorhyncocyonid, Fordonia lawsoni sp. nov. and Leptictidium prouti sp. nov. from the UK earliest Eocene, described here, are older than any previously recorded member of the family. They are represented by teeth from numerous loci, which allow a better understanding of the sparsely known dentitions of currently known pseudorhyncocyonids. This facilitates the recognition of two further species of Leptictidium, L. listeri sp. nov. from the Middle Eocene of Germany and L. storchi sp. nov. from the Late Eocene of France. Study of occlusal relationships also helps to fill gaps in our knowledge of missing tooth loci. Cladistic analysis of pseudorhyncocyonids with their previously judged closest relatives, the Leptictidae, Pantolesta and Palaeanodonta, shows that two European species, Diaphyodectes prolatus and Palaeictops? levei, formerly thought to be leptictids, are instead primitive pseudorhyncocyonids, extending the range of the family further back in time to the Middle Paleocene. P? levei is placed in the new genus Phakodon gen. nov. The analysis also shows that the Pseudorhyncocyonidae are sister group to the other three groups combined and that family‐level differentiation in this probable clade took place as early as the earliest Paleocene.     

Diversity of Menispermaceae from the Paleocene and Eocene of South China

We present here the earliest known Asian fossil records of the Menispermaceae based on fossil fruits from Paleocene and Eocene localities in South China. A new genus and species, Paleoorbicarpum parvum sp. nov., and two new species of Stephania Loureiro, S. ornamenta sp. nov. and S. geniculata sp. nov., are recognized from Paleocene deposits of the Sanshui Basin, Guangdong, and a new occurrence of the widespread Eocene species Stephania auriformis (Hollick) Han & Manchester is recognized from the Maoming Basin, Guangdong. The Paleocene Stephania specimens described here represent the earliest fossil endocarp record of the Menispermaceae in eastern Asia. This discovery shows that the moonseed family had arrived in tropical and humid South China by at least the middle Paleocene, which provides important evidence for the origin and phytogeographic history of the family.     

Discosorids and Oncocerids (Cephalopoda) of the Middle Ordovician Kunda and Aseri Regional Stages of Baltoscandia and the early evolution of these groups

Discosorid and oncocerid cephalopods of the Kunda and Aseri Regional Stage (Darriwilian, Middle Ordovician) from Öland Island, Sweden and North Estonia are described for the first time. We demonstrate that a high generic and specific diversity of Oncocerida was already established in the earliest Darriwilian. The new oncocerid species Neumatoceras borense nov. sp., N. breviborense nov. sp., Paramiamoceras breviventrum nov. sp., Richardsonoceras gastroscopium nov. sp., R. gerhardi nov. sp., R. goldmanni nov. sp., R. haelluddenense nov. sp., Richardsonoceroides kingi nov. sp., R. oelandense nov. sp., and R. rhytium nov. sp. are erected; Richardsonoceroides schiefferdeckeri (Dewitz) is revised. The new graciloceratid Kundoceras evansi nov. gen., nov. sp. is erected. A review of records of early oncocerids from China shows that Richardsonoceras Foerste is probably the most primitive oncocerid and supports the hypothesis that oncocerids evolved in the early Floian or latest Tremadocian from a Bassleroceras Ulrich and Foerste-like ancestor. The new discosorid genus Paldoceras nov. gen. and species Paldoceras paldiskense nov. sp. and P. neptunsakerense nov. sp. are erected. These discosorids record the morphological link between the late Darriwilian Ruedemannoceras Flower and the early Floian Apocrinoceras Teichert and Glenister, which is considered to be the earliest discosorid.     

New trogons from the early Tertiary of Germany

A new trogon (Aves, Trogoniformes), Primotrogon? pumilio sp. nov., is described from the Middle Eocene (49 Ma) of Messel in Germany. It is the first articulated skeleton of an Eocene trogon, of which only very few fragmentary remains were hitherto known. It is also the earliest complete skeleton of a trogon, exceeding previous finds (Primotrogon wintersteini Mayr 1999) by at least 15 million years. An isolated wing of a trogon is further identified from the early Oligocene (about 30–34 Ma) of the fossil site Frauenweiler in Germany. Primotrogon? pumilio and all other sufficiently well‐preserved early Tertiary trogons appear to be stem group representatives of the Trogoniformes, corresponding with molecular clock data indicating a mid‐Tertiary origin of crown group Trogoniformes.     

A New Small-Bodied Species of <Emphasis Type="Italic">Palaeonictis</Emphasis> (Creodonta,Oxyaenidae) from the Paleocene-Eocene Thermal Maximum

Oxyaenid creodonts are extinct carnivorous mammals known from the Paleogene of North America, Europe, and Asia. The genus Palaeonictis is represented by three species that together span the late Paleocene to early Eocene of North America, and at least one species from the early Eocene of Europe. Previously, only a single trigonid of Palaeonictis was known from the interval encompassing the Paleocene-Eocene Thermal Maximum (PETM) in North America. We describe Palaeonictis wingi sp. nov. from the PETM in the Cabin Fork drainage, southeastern Bighorn Basin, Wyoming, based on associated right and left dentaries with P₂-M₂. Palaeonictis wingi sp. nov. is substantially smaller than the other North American congeners, making it similar in size to P. gigantea from the earliest Eocene of Europe and the previously described PETM specimen. We suggest that a form similar to the large-bodied late Paleocene P. peloria from North America gave rise to two smaller species in the earliest Eocene of North America (P. wingi) and Europe (P. gigantea). Palaeonictis wingi may have given rise to P. occidentalis following the PETM in North America. Dispersal of Palaeonictis to Europe coincided with rapid global warming of 5–10°C and related geographic range shifts in plants and other animals during the PETM. It has been suggested that certain mammalian lineages decreased in body size during the PETM, possibly in response to elevated temperature and/or higher CO₂ levels. Results from a dietary analysis of Palaeonictis indicate that it was an omnivore that primarily consumed meat. This suggests that the decreased nutritious quality of vegetation caused by increased CO₂ levels was not the direct contributing factor that caused body size reduction of this lineage during the PETM. Other selective pressures such as temperature, aridity, and prey size may have also contributed to the smaller body size of carnivorous mammals during this interval, although the presence of smaller species could also be explained by latitudinal range shifts of mammals during the PETM.     

Spore assemblages from the Lower Devonian Xujiachong Formation from Qujing,Yunnan, China

Abstract: The Lower Devonian Xujiachong Formation from the vicinity of Qujing City, Yunnan, China is interpreted as a terrestrial‐fluviatile‐lacustrine sequence. It contains important nonmarine biotas including plants, fish and invertebrates. The plants are particularly interesting as they include many endemic taxa. Dispersed spore assemblages have been recovered from the upper part of this formation. The spores are well preserved and of moderate thermal maturity. They are systematically described and four new species erected: Aneurospora xujiachongensis sp. nov., Chelinospora ouyangii sp. nov., Camptozonotriletes? luii sp. nov. and Leiozonospora xichongensis sp. nov. One new combination is proposed: Aneurospora conica (Ouyang and Lu) comb. nov. This is a rare report of a Lower Devonian dispersed spore assemblage from the South China Plate. Indeed, few dispersed spore assemblages of this age are known outside of Euramerica and Northern Gondwana. It is suggested that the Xujiachong Formation spore assemblages can all be equated to the polygonalis‐emsiensis Spore Assemblages Biozone (PE SAB) of Richardson and McGregor (1986) indicating an early (but not earliest) Pragian to ?earliest Emsian age. However, caution is urged, because biostratigraphical interpretation is difficult owing to distinct differences between dispersed spore assemblages from South China and Euramerica/Northern Gondwana. This almost certainly reflects palaeophytogeographical variation and regional endemism among early land plant floras on widely separated land masses. Palynofacies analysis supports a nonmarine origin for the deposits of the Xujiachong Formation, with the very rare marine palynomorphs that were encountered interpreted as reworked.     

New macrostomatan snake from the Paleogene of northwestern Argentina

The lower Eocene Lumbrera Formation in Salta province, northwestern Argentina, outstands for providing snake remains from a non-Patagonian Paleogene site. The material consists of articulated precloacal vertebrae that represent a new medium-sized macrostomatan snake, namely Amaru scagliai nov. gen., nov. sp. The vertebral characters of Amaru scagliai nov. gen., nov. sp., suggest affinities with advanced clades, which is consistent with the recognition of derived macrostomatans in the early Paleocene of Bolivia and early Eocene of Brazil. The new snake confirms the presence of macrostomatan snakes in South America as early as the Eocene and suggests that the southern continents may have played an unsuspected role in the origin and evolution of advanced macrostomatans during the earliest Cenozoic.     

Primates and Plesiadapiformes from Cos (Eocene; Quercy,France)

A new fauna has been collected from a fissure filling named Cos in the Quercy region, southwestern France. It includes four primate species and a plesiadapiform. The cercamoniine adapiform Protoadapis andrei Godinot and Vidalenc, nov. sp., is represented by material that adds to our knowledge of the genus Protoadapis for the upper canine, upper molars and other morphological details. It appears more primitive than P. brachyrhynchus from the Old Quercy Collections. The other cercamoniine Pronycticebus cosensis Godinot and Vidalenc, nov. sp., offers insight on intraspecific variations and other details of the genus. Both species suggest a bushy evolution within these genera. Two teeth document the presence of a third cercamoniine, Anchomomys sp. indet. The new microchoerid Quercyloris eloisae Godinot and Vidalenc, nov. gen., nov. sp., has very primitive characters and seems to document a primitive member of the Pseudoloris clade. A poorly documented paromomyid plesiadapiform is distinct enough to be named Arcius moniquae Godinot and Vidalenc, nov. sp. It represents the first discovery of a plesiadapiform in the Quercy fossil record and makes a link with paromomyids surviving until the late Lutetian European Mammal Paleogene MP 13 reference level. The primates indicate a broad age interval between MP 10 (late Ypresian) and MP 12 (mid-Lutetian). The identification of the same species of Pronycticebus and of the new genus in the Vielase fauna suggests more precisely the MP 10–MP 11 (late Ypresian–early Lutetian) time interval.     

A new large philisid (Mammalia,Chiroptera, Vespertilionoidea) from the late Early Eocene of Chambi,Tunisia

Abstract: Among the new dental remains from the late Early Eocene of Chambi (Kasserine area, Tunisia) is a large‐sized upper molar of a new bat species, Witwatia sigei nov. sp. (Chiroptera, Vespertilionoidea, Philisidae), described herein. The locality of Chambi has revealed evidence for an early appearance of two modern microchiropteran superfamilies in Africa: Dizzya exsultans, a Philisidae, which is considered to be an archaic Vespertilionoidea, and an indeterminate Rhinolophoidea. In addition to D. exsultans, the new species, W. sigei, is the second representative of the Philisidae in this locality. W. sigei extends back to the late Early Eocene the occurrence of the genus Witwatia, which was previously only reported from the early Late Eocene of the Fayum (BQ‐2, Egypt). By analogy with the largest extant microbats, the large size of Witwatia suggests a tendency to the opportunistic diet of this taxon, thereby contrasting with the strict insectivory characterizing primitive bats found in other continents in the same epoch.     

New gall midges of the tribe Leptosynini (Diptera,Cecidomyiidae) from the Late Eocene ambers and the classification of the supertribe Heteropezidi

Three new genera and 27 new species of gall midges are described from the Late Eocene ambers: Henria baltica sp. nov., Frirenia manca sp. nov., F. musicata sp. nov., Leptosyna samlandica sp. nov., L. fastosa sp. nov. from Baltic amber and H. xystica sp. nov., H. liquida sp. nov., Stellasegna vlaskini gen. et sp. nov., S. vaporea sp. nov., S. nexa sp. nov., Rasnitsia verticosa gen. et sp. nov., F. rohdendorfi sp. nov., F. schevchenkoi sp. nov., F. melica sp. nov., F. lukashevichae sp. nov., F. leporidis sp. nov., F. marmarygma sp. nov., F. vesana sp. nov., Vincinescia alisae gen. et sp. nov., L. margarita sp. nov., L. munifera sp. nov., L. sukachevae sp. nov., L. assa sp. nov., L. larga sp. nov., L. vegeta sp. nov., L. vaticina sp. nov., and L. shcherbakovi sp. nov. from Rovno amber. Strobliella capitata Fedotova is redescribed as Henria capitata (Fedotova, 2004) (comb. nov.). Diagnoses of Henria (= Electroxylomyia Nel et Prokop, syn. nov.), Frirenia, and Leptosyna are revised. As a result, Henria comprises 3 extant and 5 extinct (Late Eocene) species, including H. eocenica (Nel et Prokop), comb. nov. (= Electroxylomyia eocenica), Frirenia comprises 1 extant and 10 Late Eocene species, and Leptosyna comprises 3 extant and 11 Late Eocene species. The tribe Heteropezini is elevated to the supertribal rank (Heteropezidi) and included in the subfamily Lasiopterinae. Leptosynini is treated as a separate tribe, and Lasiopterinae is considered as part of Cecidomyiidae s. str. (i.e., excluding Lestremiidae). Keys to the tribes and genera of Heteropezidi and to species of Henria, Stellasegna, Frirenia, and Leptosyna are provided. The gall midge faunas of the Rovno and Baltic ambers are compared. Phylogenetic relationships within the supertribe are hypothesized.     

Cretaceous and Eocene fossils of the rare extant genus Synneuron Lundstrom (Diptera: Canthyloscledidae): evidence of a true Pangean clade

The first two fossil species of the canthyloscelid genus Synneuron are described based on compression wings. Synneuron eomontana sp. nov. is described from the Middle Eocene Coal Creek Member of the Kishenehn Formation, in the USA, and Synneuron jelli sp. nov. is described from the Lower Cretaceous Koonwarra Fossil Bed of the Korumburra Group, in Australia. The wings are illustrated and compared to the extant species of the genus, to species of the three other recent genera of Canthyloscelidae and to an anisopodid. A phylogenetic analysis of the relationships between the species of Synneuron was performed. The Eocene fossil S. eomontana appears as sister of the pair of recent Holarctic species of the genus, while the Australian Cretaceous species S. jelli is sister of the clade with the species of Synneuron of the northern hemisphere. The sister group of Synneuron is the canthyloscelid clade (Hyperoscelis + Canthyloscelis), for which a middle Jurassic fossil is known. At the early Cretaceous, Gondwana was already separated from Laurasia and the disjunction between the species of Synneuron in Australia and the northern hemisphere clade of the genus suggest a true pangeic origin for the genus. The biology of the canthyloscelid larvae is shaped by its trophic specialization—xylosaprophagous. This suggests that the transition from the Pangean Jurassic gymnosperm-dominated forests to the late Cretaceous angiosperm-dominated forests may be related to the low recent diversity of Synneuron or of the canthyloscelids in the world—and maybe to the extinction of the genus in the southern hemisphere. This major turnover of the vegetation type along the Cretaceous may be also somehow related to the complete extinction of other groups of flies strictly associated with gymnosperms, as may be the case of the lower brachyceran family Zhangsolvidae. This speculation needs additional corroboration from other groups, that will become available with the combination of systematics, paleontology and biogeographical information of different early Cretaceous clades.     

New species of caddisflies (Insecta: Trichoptera) from the Rovno amber,Eocene of Ukraine

A new genus and seven new species of the caddisfly families Philopotamidae, Polycentropodidae, Ecnomidae, Hydroptilidae, and Leptoceridae (Wormaldia nasticentia sp. nov., Holocentropus flexiflagrum sp. nov., Electrocyrnus perpusillus gen. et sp. nov., Archaeotinodes reveraverus sp. nov., Agraylea electroscientia sp. nov., Triplectides palaeoslavicus sp. nov., and Leptocerus solifemella sp. nov.) from the Rovno amber (Upper Eocene, Ukraine) are described     

Provannid and provannid‐like gastropods from the Late Cretaceous cold seeps of Hokkaido (Japan) and the fossil record of the Provannidae (Gastropoda: Abyssochrysoidea)

A fauna of provannid and provannid‐like shells is described from Upper Cretaceous seep carbonates in Hokkaido, Japan. We describe two new provannid species, Provanna tappuensis sp. nov. and Desbruyeresia kanajirisawensis sp. nov. , with preserved protoconchs of unquestionable provannid type with decollate apex. This material confirms the occurrence of Provannidae as early as the Middle Cenomanian. We also describe Hokkaidoconcha gen. nov. and a new family Hokkaidoconchidae fam. nov. , with two named species, H. hikidai sp. nov. and H. tanabei sp. nov . Hokkaidoconchidae are possibly related to the Provannidae, judging from a similar, but not decollate larval shell, although the juvenile teleoconch whorls differ in being of a general cerithimorph appearance and the details of the aperture are unknown. Furthermore, we review the published fossil record of Provannidae and Abyssochrysidae, and we consider that in those older than the Eocene, there is no evidence preserved that unequivocally supports a position there. The Jurassic Acanthostrophia acanthica from Italy seems to be the oldest known record of Abyssochrysidae, and the most reliable occurrence of the family, older than from the Miocene. Other fossil, pre‐Miocene species that have been classified in the Abyssochryssidae are provisionally referred to Hokkaidoconchidae. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 154 , 421–436.     

A new species of Limnofregata (Pelecaniformes: Fregatidae) from the Early Eocene Wasatch Formation of Wyoming: implications for palaeoecology and palaeobiology

A humerus and a coracoid from the Early Eocene Wasatch Formation in the Washakie Basin of south‐western Wyoming are the oldest materials (by ~2 million years) of the pelecaniform Limnofregata (Aves) and represent a new large species, Limnofregata hutchisoni sp. nov. This fossil is the oldest known member of the frigatebird lineage. Other than its large size relative to Limnofregata azygosternon and L. hasegawai, the new material is very similar morphologically to other known Limnofregata specimens. The size of this new species is comparable to the largest living species (e.g. Fregata minor and Fregata magnifiscens) and much larger than the two described species of Limnofregata. This fossil indicates that the hard minimum date previously advocated for molecular calibration of the split between Fregatidae and Suloidea is an underestimate by approximately two million years. The presence of early pelecaniform bird lineages (represented by Limnofregata and Masillastega) in limnic ecosystems prior to their known occurrences in marine deposits/habitats appears to indicate that some clades of pelecaniform birds may have undergone an evolutionary transition from freshwater to marine habitats in a pattern reminiscent of what has been suggested during the evolution of pinnipeds or that their palaeoecology included broader niches ranging across a variety of aquatic habitats. That transition in habitat occupation and the origin of many of the characteristic biological aspects present in the crown frigatebird clade likely occurred during a significant temporal gap (> 45 million years) in the fossil record of the frigatebird lineage after these earliest occurrences in the Early Eocene and before the oldest records of the extant Fregata species in the Pleistocene.     

New adapiform primate of Old World affinities from the Devil's Graveyard Formation of Texas

Most adapiform primates from North America are members of an endemic radiation of notharctines. North American notharctines flourished during the Early and early Middle Eocene, with only two genera persisting into the late Middle Eocene. Here we describe a new genus of adapiform primate from the Devil’s Graveyard Formation of Texas. Mescalerolemur horneri, gen. et sp. nov., is known only from the late Middle Eocene (Uintan) Purple Bench locality. Phylogenetic analyses reveal that Mescalerolemur is more closely related to Eurasian and African adapiforms than to North American notharctines. In this respect, M. horneri is similar to its sister taxon Mahgarita stevensi from the late Duchesnean of the Devil’s Graveyard Formation. The presence of both genera in the Big Bend region of Texas after notharctines had become locally extinct provides further evidence of faunal interchange between North America and East Asia during the middle Eocene. The fact that Mescalerolemur and Mahgarita are both unknown outside of Texas also supports prior hypotheses that low-latitude faunal assemblages in North America demonstrate increased endemism by the late middle Eocene.     

Etheliaceae fam. nov. (Gigartinales,Rhodophyta), with a clarification of the generitype of Ethelia and the addition of six novel species from warm waters

Based upon COI‐5P, LSU rDNA, and rbcL sequence data and morphological characteristics, six new members of the noncalcified crustose genus of red algae Ethelia are described in a new family, Etheliaceae (Gigartinales), sister to the recently described Ptilocladiopsidaceae. The novel species are described from subtropical to tropical Atlantic and Indo‐Pacific Ocean basins; E. mucronata sp. nov. and E. denizotii sp. nov. from southern and northern Western Australia respectively, E. wilcei sp. nov. from the Cocos (Keeling) Islands of Australia, E. suluensis sp. nov. from the Philippines, E. umbricola sp. nov. from Bermuda and E. kraftii sp. nov. from Lord Howe Island, Australia. The generitype, Ethelia biradiata, originally reported from the Seychelles, Indian Ocean, is added to the Western Australian flora.     

New species of sap beetles (Coleoptera: Nitidulidae: Epuraeinae,Cybocephalinae) from the Baltic amber

A new genus, Baltoraea gen. nov., two new species of this genus, B. insignis sp. nov. and B. simillima sp. nov., and three other new species, Cybocephalus (Cybocephalus) balticus sp. nov., C. (C.) electricus sp. nov. and C. (C.) kerneggeri sp. nov. are described from the Late Eocene Baltic amber. The systematic position of the genus Baltoraea and presumable bionomics of the new sap beetles are discussed.