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     

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.     

An assemblage of early Oligocene lizards (Squamata) from the locality of Boutersem (Belgium), with comments on the Eocene–Oligocene transition

A diverse, new lizard assemblage from the early Oligocene of Belgium is described. The Boutersem railway local fauna is the most species‐rich lizard assemblage yet reported from the European early Oligocene. Four lizard taxa are present: Lacertidae, Anguidae, Scincoidea and Platynota. One new species is described, Folisaurus boutersemensis sp. nov . This fauna provides new insight into the profound turnover that took place during the Eocene/Oligocene boundary in Europe. The new fauna confirms a marked decrease in diversity across the Eocene/Oligocene boundary. Two groups encountered in the European late Eocene became extinct (Iguanidae*, Glyptosaurinae). Estimates of species‐level extinctions range up to 80%. These estimates include members of virtually all the families present in the late Eocene. The relative importance of climate change and biotic interactions in controlling this pattern is discussed, and negative interactions between lizards and new carnivorous mammals are favoured. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155 , 148–170.     

Intercontinental dispersal of giant thermophilic ants across the Arctic during early Eocene hyperthermals

Early Eocene land bridges allowed numerous plant and animal species to cross between Europe and North America via the Arctic. While many species suited to prevailing cool Arctic climates would have been able to cross throughout much of this period, others would have found dispersal opportunities only during limited intervals when their requirements for higher temperatures were met. Here, we present Titanomyrma lubei gen. et sp. nov. from Wyoming, USA, a new giant (greater than 5 cm long) formiciine ant from the early Eocene (approx. 49.5 Ma) Green River Formation. We show that the extinct ant subfamily Formiciinae is only known from localities with an estimated mean annual temperature of about 20°C or greater, consistent with the tropical ranges of almost all of the largest living ant species. This is, to our knowledge, the first known formiciine of gigantic size in the Western Hemisphere and the first reported cross-Arctic dispersal by a thermophilic insect group. This implies intercontinental migration during one or more brief high-temperature episodes (hyperthermals) sometime between the latest Palaeocene establishment of intercontinental land connections and the presence of giant formiciines in Europe and North America by the early middle Eocene.     

A new primitive alligatorine from the Eocene of Thailand: relevance of Asiatic members to the radiation of the group

Despite the report of fragmentary remains from China, the fossil record of primitive Alligatorinae is unbalanced with most of its members reported from North America and Europe. Here, we describe cranial, mandibular, and some postcranial elements of a new alligatorine taxon from the Late Eocene of Krabi Basin, southern Thailand. Krabisuchus siamogallicus gen. et sp. nov. was a small animal probably not surpassing 2 m in total length. Despite deformation, the remains show a dorsally elevated cranium similar to Arambourgia gaudryi and Osteolaemus tetraspis. The new alligatorine is characterized by a very short snout and a blunt dentition in the posterior region. Krabisuchus siamogallicus is the first fossil alligatorine from Asia to be incorporated into cladistic analysis. Results indicate a close relationship of the newtaxon to Procaimanoidea, Arambourgia gaudryi, and Allognathosuchus polyodon, all of them being sister groups to the Alligator clade. Alligatorine were widespread as early as the Late Eocene across the northern hemisphere. Scenarios of alligatorine dispersal during the Palaeogene are discussed in comparison with the European and North American alligatorine fossil record. Basal alligatorines were probably land‐dwelling animals contrary to modern‐day crocodylians. Such a mode of life may have allowed this group to colonize vast territories, especially during periods of global warm climates. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 158 , 608–628.     

New remains of Chambius kasserinensis from the Eocene of Tunisia and evaluation of proposed affinities for Macroscelidea (Mammalia,Afrotheria)

Abstract

Chambius kasserinensis from the late Early or early Middle Eocene Chambi locality, central Tunisia, is undoubtedly the oldest known macroscelidid and possibly the basalmost representative of the order Macroscelidea. Hence, since its discovery in 1986, Chambius has played a key role in analyses focusing on afrotherian and eutherian phylogeny; for instance, as early as 1995, Butler’s review of fossil macroscelideans highlighted the central position of Chambius in the origin of the order. Despite this, Chambius remained poorly known until recently. Here based on new mandibular fragments, well-preserved upper molars and CT scan analysis of the holotype maxilla, Chambius is revised. Its dentition is first described in detail, providing a precise characterization of the genus. Chambius is notably defined by a submolariform P₄ with a three-cusped talonid, a reduced talonid on M₂, and a prominent metaconule on M^1?2. Interestingly, the two transverse lophs of the upper molars are basically formed by preconulecristae, evoking the recently defined peculiar bilophodonty of paenungulates. Comparisons with other Paleogene and modern macroscelidids, European Louisinidae, and North American Apheliscidae are also made, allowing the various hypotheses about the origin and early evolution of macroscelidids to be reviewed.     

Tortoise (Reptilia,Testudinidae) radiations in Southern Africa from the Eocene to the present

Africa, inclusive of the West Indian Ocean islands, harbours 11 of the world's 16 extant testudinid genera. Fossil records indicate that testudinids originated in Asia and dispersed first to North America and Europe (Early Eocene) and later to Africa (Late Eocene). We used mitochondrial (1870 bp) and nuclear (1416 bp) DNA sequence data to assess whether molecular data support the late cladogenesis of Southern African testudinid lineages. Our results revealed strong support for the monophyly of a clade consisting of Kinixys, the two Malagasy genera and four Southern African genera (Psammobates, Stigmochelys, Homopus and Chersina). Kinixys diverged from this clade in the Late Palaeocene, suggesting that testudinids occupied Africa at an earlier date than indicated by fossil records. The Southern African tortoises consist of three, strongly supported clades: Psammobates + Stigmochelys; the five‐toed Homopus + Chersina; and the four‐toed Homopus. Due to the paraphyly of Homopus, we propose the taxonomic resurrection of Chersobius for the five‐toed Homopus species (boulengeri, signatus and solus). Cladogenesis at the genus level occurred mainly in the Eocene, with Chersina and Chersobius diverging in the Oligocene. The latter divergence coincided with species‐level radiations within Homopus (areolatus and femoralis) and Psammobates (oculifer, geometricus and tentorius). Our phylogeny could not resolve relationships within Psammobates, indicating rapid speciation between the Late Oligocene and Early Miocene. The Chersobius species were the last to diverge in the Early to Mid‐Miocene. By the Mid‐Miocene, P. tentorius started to differentiate into four lineages instead of the three recognized subspecies: P. t. tentorius, P. t. trimeni and two P. t. verroxii subclades occurring north and south of the Orange River, respectively. Terminal radiations in several taxa suggest the existence of cryptic species and a more diverse tortoise fauna than currently recognized. Factors contributing to this diversity may include the early origin of African testudinids and climatic fluctuations over a heterogeneous landscape.     

Combined phylogenetic analysis of a new North American fossil species confirms widespread Eocene distribution for stem rollers (Aves,Coracii)

We report a nearly complete skeleton of a new species of stem roller (Aves, Coracii) from the early Eocene Green River Formation of North America. The new species is most closely related to two species‐depauperate lineages, Coraciidae (rollers) and Brachypteraciidae (ground rollers), that form a monophyletic crown clade (Coracioidea) with an exclusively Old World extant distribution. Phylogenetic analysis utilizing a matrix of 133 morphological characters and sequence data from three genes (RAG‐1, c‐myc, and ND2) identifies the new species as a stem member of the Coracii more closely related to the crown clade than the only previously identified New World taxon, Primobucco mcgrewi. The phylogenetic placement of the new species and Primobucco mcgrewiindicates a widespread northern hemisphere distribution in the Eocene with subsequent restriction to Africa, Madagascar, Australia, and temperate to tropical parts of Europe and Asia. It provides evidence of further ecological diversity in early stem Coracii and convergence on crown morphologies. The new species contributes to mounting evidence that extant distributions for major avian subclades may be of comparatively recent origin. Further late Palaeogene sampling is needed to elucidate potential drivers for shifting avian distributions and disappearance of Coracii from North America. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 157 , 586–611.     

Phylogenetic revision of the Hippasterinae (Goniasteridae; Asteroidea): systematics of deep sea corallivores,including one new genus and three new species

The Hippasterinae is a subfamily within the Goniasteridae, consisting of five genera and 26 species, which occur in cold‐water settings ranging from subtidal to abyssal depths. All known genera were included in a cladistic analysis resulting in two most parsimonious trees, supporting the Hippasterinae as monophyletic. Our review supports Sthenaster emmae gen. et sp. nov. as a new genus and species from the tropical Atlantic and two new Evoplosoma species, Evoplosoma claguei sp. nov. and Evoplosoma voratus sp. nov. from seamounts in the North Pacific. Hippasteria caribaea is reassigned to the genus Gilbertaster, which previously contained a single Pacific species. Our analysis supports Evoplosoma as a derived deep water lineage relative to its continental‐shelf, shallow water sister taxa. The genus Hippasteria contains approximately 15 widely distributed, but similar‐looking species, which occur in the northern and southern hemispheres. Except for Gilbertaster, at least one species in each genus has been observed or is inferred to prey on deep‐sea corals, suggesting that this lineage is important to the conservation of deep‐sea coral habitats. The Hippasterinae shares several morphological similarities with Circeaster and Calliaster, suggesting that they may be related. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 266–301.     

Derived,still living cockroach genus Cariblattoides (Blattida: Blattellidae) from the Eocene sediments of Green River in Colorado,USA

Abstract Cariblattoides labandeirai sp.n. from the Eocene sediments of Green River in Colorado, USA bear only two plesiomorphies, but also several significant autapomorphies within the advanced and highly derived living cockroach genus. Thus, Cariblattoides with extant occurrence in the Caribbean and South America was historically common in the Nearctic, and represents important evidence for the occurrence of derived living genera of cockroaches ～50 Ma ago. Generally, the vast majority of living genera were absent during the Palaeocene, thus the diversification of most living cockroach lineages near the Palaeocene/Eocene boundary must have been extremely rapid. Females of living C. suave, the type species, have identical (sophisticated) coloration of pronotum, but the most related living taxa are C. piraiensis and C. fontesi from Brazil (supported by phylogenetical analysis).     

Earliest South American paucituberculatans and their significance in the understanding of ‘pseudodiprotodont’ marsupial radiations

We describe the oldest Paucituberculata marsupials, from the La Barda and Las Flores localities (Argentina; Late Palaeocene, and Early–Middle Eocene), as well as from the Itaboraí Basin (Brazil; Late Palaeocene). The new taxa are represented by very scarce, although well‐preserved, dental remains. A parsimony analysis was performed in order to evaluate the phylogenetic affinities of these taxa. Representatives of both Riolestes capricornicus gen. et sp. nov. and Bardalestes hunco gen. et sp. nov. appear to be basal paucituberculatans, and their molar features give clues on the early evolution of the representatives of this order. Within the Paucituberculata we recognize two major clades: Caenolestoidea and Palaeothentoidea. We conclude that ‘pseudodiprotodont’ marsupials of the traditional literature (i.e. Polydolopimorphia + Paucituberculata) do not form a natural group. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155 , 867–884.     

Extinct large colobines from the Plio-Pleistocene of Africa

Three genera and six species of extinct African Plio-Pleistocene Colobinae are discussed. One new genus, Rhinocolobus and three new species, R. turkanaensis, Cercopithecoides kimeui and Paracolobus mutiwa are named. These colobines show a diversity in postcranial and dental morphology not seen among extant species. Rhinocolobus was most similar to extant Colobus in postcranial morphology and had similar high-cusped shearing teeth. Cercopithecoides shows a number of postcranial skeletal features typical of terrestrial cercopithecid species and has lower cusped teeth. Paracolobus, while generally more similar to Rhinocolobus than to Cercopithecoides, is intermediate in some features of its postcranial morphology. The distribution of the various taxa among East and South African sites with different palaeoenvironments is generally consistent with the morphological interpretation. With the exception of Cercopithecoides kimeui, which persisted a little longer, these large colobines disappear from the fossil record about 1.8 million years ago. Their disappearance coincides with an interval of increasing aridity documented at Olduvai Gorge, the Omo Valley, and East Turkana.     

Phylogeny of cardiid bivalves(cockles and giant clams): revision of the Cardiinae and the importanceof fossils in explaining disjunct biogeographical distributions

Despite being the type subfamily of the Cardiidae, theCardiinae has received relatively little attention from systematists.Recent investigations of the phylogenetic relationships betweenand within various subtaxa of cockles and giant clams have incorporatedseveral cardiine representatives. These studies have implied thatCardiinae as usually construed is a paraphyletic group. The presentstudy, a cladistic analysis that includes exemplars of the 18 generallyaccepted genera and subgenera of Cardiinae, as well as exemplarsfrom two notable lineages heretofore not assigned to any genus‐leveltaxon, also indicate cardiine paraphyly. Cardiinae is restrictedto Cardium, Bucardium, Vepricardium, Dinocardium, Planicardium, Chesacardium, Acanthocardias.s., A. (Rudicardium) and Schedocardia. Theclosest relatives of Cardiinae are the Eocene Sawkinsia anda clade (the ‘CFTL lineage’) containing Plagiocardium, Maoricardium,Clinocardiinae, Fraginae, Lymnocardiinae and Tridacninae (giantclams). The remaining taxa that have usually been placed in Cardiinaeform a monophyletic group that is the sister taxon to Sawkinsia +Cardiinae + CFTL; the new subfamily Orthocardiinae is erectedfor this group. Orthocardiinae contains the taxa Orthocardium, Loxocardium, Europicardium, Hedecardiums.s., H. (Titanocardium), H. ( Iheringicardium ) subgen.nov ., Agnocardia, Afrocardium and Freneixicardia gen.nov .; only Afrocardium (nine species) and Europicardium (onespecies) are extant. Hedecardium s.l. was not restrictedto New Zealand as usually argued but was present in Myanmar andAustralia, as well as being represented in South America by H. (Iheringicardium).Renewed studies of the long‐ignored marine invertebrate Cenozoicfaunas of South America may yield additional examples of taxa longthought to be endemic to New Zealand. The only other orthocardiinewith representatives in the Americas is the extinct Agnocardia. Agnocardia’sclosest relative, Afrocardium, long misconstrued as a memberof Fraginae, is known only from Pleistocene and Recent faunas, usuallyfrom the Indian Ocean. This disjunct biogeographical distributionis found to be a function of a century of misidentification of fossiland Recent species, for species attributable to Agnocardia occurin the Palaeogene of Europe and Africa; likewise, species attributableto Afrocardium occur from the Eocene to Pliocene of Europeand Africa. A greater awareness of modern taxonomy of both fossiland Recent forms is necessary to avoid constructing erroneous biogeographicaldistributions and phylogenetic hypotheses. Discrepancies betweenthe results of (1) rigorous character analysis and phylogenetichypotheses and (2) published stratigraphic and palaeobiogeographicaldistributions of fossils, are not necessarily the results of aninadequate fossil record. Instead, stratigraphic and palaeobiogeographicalanomalies may be the result of inadequate attention paid to theactual fossils by taxonomists. © 2002 The LinneanSociety of London. Zoological Journal of the Linnean Society,2002, 136 , 321?369     

Late Cretaceous to early tertiary Dinoflagellate cysts from Narasapur Well-1, Godavari-Krishna basin,South India

Dinoflagellate cyst assemblages are described fromsome of the richest horizons in the Narasapur Well-1, Godavari-Krishna basin, Andhra-Pradesh, India. Twenty-five taxa belonging to nineteen genera are recorded. The majority exhibit close morphological similarity with species described earlier from Europe, North America and Australia. One new genus, Godavariella, and three new species, Godavariella venkatachalae, Fibrocysta variabilis and Cyclonephelium indicum, are proposed; several more new forms are described but, since meagrely represented, are not named. The dinoflagellate assemblages, along with spore and pollen studies carried out earlier by Venkatachala & Sharma (1984), suggest a Late Cretaceous (Maastrichtian) to Early Tertiary (Palaeocene) age for these sediments.     

FRUITS AND SEEDS OF TRIBE GORDONIEAE (THEACEAE) FROM THE EOCENE OF NORTH AMERICA

A new genus and species Gordoniopsis polysperma and two new species of Gordonia (Gordonieae, Camellioideae, Theaceae) are described based on fossil fruit and seed remains. These specimens are part of a large flora consisting of various plant organs from the middle Eocene Claiborne Formation in western Kentucky and Tennessee. Gordoniopsis is a five-valved loculicidally dehiscent capsule similar to capsules of Gordonia but differing in having unwinged seeds and a greater number of seeds per locule. The two Gordonia species are among the earliest unequivocal records of the genus and two of only four fossil Gordonia species known with in situ seeds. Two extinct genera, Gordoniopsis and Andrewsiocarpon, and the extant genus Gordonia in the tribe Gordonieae are known from the middle Eocene Claiborne flora, suggesting an early radiation within the tribe. Based on a survey of Recent fruits and seeds we concur with Keng's proposal to merge Laplacea with Gordonia.     

The origins of modern nonmarine ostracod faunas: evidence from the Late Cretaceous and Early Palaeogene of Mongolia

The Cretaceous and Tertiary development of Mongolian non-marine ostracod faunas is reviewed. During the Late Cretaceous and Early Palaeogene, representatives of the Cypridoidea were widespread and common, Cytheroidea less so and the Darwinuloidea comparatively rare. The evolutionary history of the subfamily Talicyprideinae is considered, with reference to the genera Talicypridea, Altanicypris, Khandiaand Bogdocypris. It is suggested that the extinct Talicyprideinae were related to the mid-Cretaceous to Recent subfamily Cypridinae (e.g. the genus Cypris), both belonging to the family Cyprididae. It is shown that early representatives of the Cyprididae, one of the most diverse non-marine cypridoidean families today, were present from Early Cretaceous onwards (e.g. Lycopterocypris, Mongolocypris), alongside the dominant Cretaceous cypridoideans, the Cyprideidae (e.g. Cypridea), which became extinct in the Palaeogene.     

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.