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.     

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 genera and species of Scelionidae (Insecta: Proctotrupoidea) from Rovno amber

Two new genera and three new species of parasitic wasps from the family Scelionidae are described from the Late Eocene Rovno amber: Pseudotelea gracilis Kononova, gen. et sp. nov. (subfamily Scelioninae), Pseudidris striatus Kononova, gen. et sp. nov., and Ceratobaeoides cornutus Kononova, sp. nov. (subfamily Baeinae). The new taxa are compared with representatives of the Recent fauna.     

New trace fossils of termites (Insecta: Isoptera) from the late Eocene‐early Miocene of Egypt,and the reconstruction of ancient isopteran social behavior

Three new ichnogenera and five new ichnospecies are described for new trace fossils of termitaria (including associated gallery systems) of subterranean termites from upper Eocene through lower Miocene rocks of northern Egypt. All but two ichnospecies (Krausichnus trompitus, ichnogen, and ichnosp. nov., and A’, altus, ichnosp. nov.) show varying degrees of affinity to nests of the extant subterranean termite species Sphaerotermes sphaerothorax (Termitidae, Macro‐termitinae).

Termitichnus qatranii (Bown, 1982) is divided into two ichnospecies, the previously named T. qatranii, and a more generalized form, T. simplicidens, ichnosp. nov. Vondrichnus obovatus, ichnogen. and ichnosp. nov., is named for simple, possibly macrotermitine nests with oblate form, and Fleaglellius pagodus, ichnogen. and ichnosp. nov., records a nest form similar to V. obovatus, but one in which successive vertical growth by chamber apposition has produced a subterranean, tower‐like structure, reminiscent of that built by extant, epigeous Cubitermes. Krausichnus trompitus, ichnogen. and ichnosp. nov., and A’, altus, ichnosp. nov., record two unique nest architectures probably produced by unknown but related species of humivorous termites. The nest architecture expressed by Krausichnus is of wholly unknown affinity and is only distantly related in form to the ichnofossils of other termite nests known from the Tertiary of Egypt.

Study of the architecture of the nests of these ancient termites reveals details important in reconstructing the phylogeny of termite nests and documents two novel blueprints for chamber expansion and society budding in what were probably primitive Macrotermi‐tinae. It also indicates that at least two constructed edifices, earlier known only as epigeous manifestations of extant termite species, were almost certainly first developed by species living underground.

The extant Macrotermitinae are a subfamily of termites originally believed to have evolved in the post‐Eocene of the Ethiopian biogeo‐graphic region. We offer fossil evidence of four distinct, possibly macrotermitine structures from upper Eocene rocks. This evidence suggests that this termite subfamily (or at least their peculiar mode of nest construction) might have evolved considerably earlier, as it is already well established and exhibits several variants by the late Eocene in Egypt.     

Chulpasia and Thylacotinga, late Paleocene-earliest Eocene trans-Antarctic Gondwanan bunodont marsupials: New data from Australia

A new marsupial from the early Eocene Tingamarra Local Fauna of southeastern Queensland, Australia, is named and referred to ChulpasiaCrochet and Sigé, 1993, a genus otherwise known from early Tertiary deposits of Peru. This taxon, Chulpasia jimthorselli nov. sp., differs in upper molar morphology only in minor details from the Peruvian type species Chulpasia mattaueri and is almost identical in size. New materials referable to the Tingamarra marsupial Thylacotinga bartholomaii Archer, Godthelp and Hand are also described. Species of Chulpasia and Thylacotinga share many striking derived as well as plesiomorphic dental features that allow recognition of a new monophyletic subfamily, Chulpasiinae. Its familial relationships are in doubt, but members of the subfamily could have provided the ancestral stock for Rosendolops and other early Tertiary South American polydolopimorphian marsupials. This is the first evidence that a Gondwanan genus of therian land mammals spanned South America, Antarctica and Australia during the early Tertiary. The very close similarity between the Peruvian and Australian fossils (and suggested short time span between their stratigraphic occurrences) provides further paleontological support for a trans-Antarctic land connection between South America and Australia extending well into the early Paleogene.     

New ceratomorph perissodactyls (Mammalia) from the Middle and Late Eocene of Mongolia: their implications for phylogeny and dating

Eight species rank taxa arc described from Eocene sites in the Gobi Desert, Mongolia. Irdinolophus gen. now is erected for Desmalotherium mongoliense Osborn and for Irdinolophus? tuiensis sp. nov. The genus is placed at the stem of the family Depcrctellidae. Intraspecific variation in lekolophus magnus is documented for the first time; Pachylophus is synonymized with leleoloplms; and Deperetella cf.birmanica is recorded far north of its previously known range in Burma. The first known lower teeth of Colodon inceptus support referral ofDesmalotherium Jissum to Colodon. The stratigraphic range of the hyracodontid Ardynia is extended back from the late Eocene to the early middle Eocene. A range of primitive tapiroids and rhinocerotoids arc analysed cladistically to resolve the problematic affinities of the extinct families Dc-peretellidae and Rhodopagidae. The Depcrctellidae are shown to belong to a monophyletic superfamily Tapiroidca, whilst the Rhodopagidae belongs to the Rhinocerotoidea, but is distinct from the Hyracodontidae. Veragromovia, previously synonymized with the tapiroid Helaleles, is resurrected as a valid genus within the Rhodopagidae. Tsagan Khutcl and other sites in the Kholbolchi formation may be older than Irdinmanhan faunas in Asia, equaling with those from the Arshanto Formation of China of probable early Lutetian or latest Ypresian age. The Irdinmanhan Mergen fauna may be slightly older than that from Irdin Manha, because of the primitive aspect of its Lophialetes.     

Sayrevilleinae Legalov,a newly recognised subfamily of fossil weevils (Coleoptera,Curculionoidea, Attelabidae) and the use of synchrotron microtomography to examine inclusions in amber

Two genera of extinct weevils, Sayrevilleus Gratshev & Zherikhin from Cretaceous New Jersey amber and Baltocar Kuschel from Eocene Baltic amber, are recognized as close relatives based on similarities revealed by the use of synchrotron tomography and the availability of new amber inclusions. The subfamily Sayrevilleinae Legalov stat. nov. is characterized by possessing mandibles with an external cutting edge and an inner blunt edge. The subfamily is placed in the family Attelabidae (s.l.), although some characters also suggest a possible relationship with the ‘higher weevils’ comprising Caridae, Brentidae, and Curculionidae. Sayrevilleus is transferred from the tribe Auletini of Rhynchitinae to Sayrevilleinae, and Sayrevilleus grimaldii Gratshev & Zherikhin is redescribed. Baltocar Kuschel is transferred from Caridae to Sayrevilleinae and revised, its type species, Baltocar succinicus (Voss), is redescribed and three new species, Baltocar groehni Riedel sp. nov. , Baltocar hoffeinsorum Riedel sp. nov. , and Baltocar subnudus Riedel sp. nov. are described based on eight well‐preserved inclusions. The genera Orapauletes Legalov and Zherichiniletes Legalov previously assigned to Sayrevilleini are regarded as Curculionoidea incertae sedis. The Sayrevilleinae were distributed over areas of North America and Europe at least since the Late Cretaceous (c. 90 Mya) and were probably relatively diverse until the Eocene (c. 44 Mya). It is speculated that they became extinct through competition with Curculionidae, which used a similar oviposition strategy. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 165 , 773–794.     

New and little known orthopteroid insects (Polyneoptera) from fossil resins: Communication 3

New taxa of Orthoptera Ensifera are described in the families Mogoplistidae [Protomogoplistes asquamosus gen. et sp. nov. (Upper Cretaceous) in the subfamily Protomogoplistinae subfam. nov. and Archornebius balticus gen. et sp. nov. (Eocene), Pseudarachnocephalus gen. nov., P. dominicanus sp. nov., and P. latiusculus sp. nov. (all Miocene) in Mogoplistinae] and Gryllidae [Eopentacentrus borealis gen. et sp. nov. (Eocene), ?Grossoxipha feminea sp. nov. (Miocene), and Apentacentrus copalicus sp. nov. in the subfamily Pentacentrinae, ?Cyrtoxipha electrina sp. nov. and ?Cyrtoxipha illegibilis sp. nov. (both Miocene) in Trigonidiinae, and Baltonemobius fossilis gen. et sp. nov. (Eocene) in Nemobiinae]. The Miocene genera Proanaxipha Vickery et Poinar and Grossoxipha Vickery et Poinar are transferred from the subfamily Trigonidiinae to Pentacentrinae. P. latoca Vickery et Poinar and Abanaxipha longispina Vickery et Poinar are redescribed; the male of the latter species is described for the first time.     

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.     

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

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

New larger foraminifers from the Eocene of the Crimea (Ukraine) and Mangyshlak (Western Kazakhstan)

Two new species of Nummulites and one subspecies of Orbitoclypeus from the Eocene of the Crimean Mountains and Mangyshlak Peninsula are described in detail. Nummulites alexisi sp. nov. and Orbitoclypeus munieri major subsp. nov. come from the Upper Ypresian, while Nummulites ninikae sp. nov. is from the Lower-Middle Lutetian. The phylogenetic position of new taxa is discussed.     

New and little-known orthopteroid insects (Polyneoptera) from fossil resins: Communication 4

New taxa of Ensifera and Caelifera orthopterans (Insecta, Orthoptera), from the families Gryllotalpidae [Marchandiinae, subfam. nov. (Lower Cretaceous)], Haglotettigoniidae [?Haglotettigonia aenigmatosa, sp. nov. (Lower Cretaceous)], Tettigoniidae [Meconematinae: Archixizicus occidentalis, gen. et sp. nov. (Eocene), Eogrigoriora gracilis, gen. et sp. nov. (Eocene), Miophlugis rostratus, gen. et sp. nov. (Miocene)], Stenopelmatidae [Siinae: Electrosia baltica, gen. et sp. nov. (Eocene); Gryllacridinae: Plesiolarnaca prior, gen. et sp. nov. (Eocene)] and Tridactylidae [Mongoloxyinae: Birmitoxya intermedia, gen. et sp. nov. (Upper Cretaceous). The Eocene species Lipotactes martynovi Zeun. and L. bispinatus Weidn. are transferred to the genus Eomortoniellus Zeun. (Tettigoniidae: Tympanophorinae); Prorhaphidophora zeuneri Chop. and P. tachycinoides Chop. are transferred to the genus Protroglophilus Gor. (Rhaphidophoridae: Protroglophilinae). The Eocene species E. handlirschi Zeun., species of the genus Protroglophilus, and a possible member of the genus Succinotettix Piton (Tetrigidae: Tetriginae), as well as a Miocene representative of the genus Archaeoellipes Heads (Tridactylidae: Tridactylinae) are also discussed.     

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.     

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.     

Eocene and Oligocene chitons (Polyplacophora) from the Paris and Hampshire Basins

Eocene and Oligocene chitons (Polyplacophora) from the Paris Basin of N France are described along with comparative material from the Hampshire Basin of the UK. The assemblages include eight species, five of which are new: Ischnochiton fehsei sp. nov., Stenoplax monila sp. nov., Chaetopleura gaasi sp. nov., C. abbessi sp. nov. and Tonicella lira sp. nov. Other taxa in the assemblages are Leptochiton cf. algesirensis, I. vectensis and S. anglica.     

The Beothukis/Culmofrons problem and its bearing on Ediacaran macrofossil taxonomy: evidence from an exceptional new fossil locality

The late Ediacaran siliciclastic successions of eastern Newfoundland, Canada, are renowned for their fossils of soft‐bodied macro‐organisms, which may include some of the earliest animals. Despite the potential importance of such fossils for evolutionary understanding, the taxonomic framework within which Ediacaran macrofossils are described is not clearly defined. Rangeomorphs from a newly discovered fossil surface on the Bonavista Peninsula, Newfoundland, require us to reconsider contemporary use of morphological characters to distinguish between genera and species within Ediacaran taxa. The new surface exhibits remarkable preservational fidelity, resolving features smaller than 0.1 mm in dimension in both frondose and non‐frondose taxa. Such preservation permits the recognition of rarely observed fourth‐ and fifth‐order rangeomorph branching, offering unparalleled opportunities to investigate the fine‐scale construction of rangeomorph taxa including Culmofrons plumosa Laflamme et al., 2012. Our observations enable resolution of taxonomic issues relating to rangeomorphs, specifically overlap between the diagnoses of the frondose genera Beothukis Brasier and Antcliffe, 2009 and Culmofrons. We propose a taxonomic framework for all Ediacaran macrofossils whereby gross architecture, the presence/absence of discrete morphological characters and consideration of growth programme are used to distinguish genera, whereas morphometric or continuous characters define taxa at the species level. On the basis of its morphological characters, Culmofrons plumosa is herein synonymized to a species (Beothukis plumosa comb. nov.) within the genus Beothukis. This discussion emphasizes the need to standardize the taxonomic approach used to describe Ediacaran macrofossil taxa at both the genus and species levels, and raises important considerations for future formulation of higher‐level taxonomic groups.