Another Laurasian connection in the Early Eocene of India: Myrmecarchaea spiders (Araneae, Archaeidae).

The first fossil Archaeidae in Cambay amber from India, of Eocene age, is documented. The inclusion is a spider exuvium and is placed as Myrmecarchaea based on the presence of elongated legs, a slightly elongated pedicel with lateral spurs, and a diastema between coxae III and IV that is similar to M.antecessor from Oise amber. The previous occurrences of the genus are from Baltic and Oise amber, both of Eocene age. Because most spiders, including Archaeidae, only molt as juveniles, the exuvium does not have adult features nor have distinct species-specific features, and a new taxon is not erected. This new record further extends the distribution of the family and genus to India 50-52 million years ago. Myrmecarchaea in Indian Cambay amber provides additional evidence that India in the Early Eocene had affinities with the Palearctic mainland rather than showing Gondwanan insularity.

A diverse crinoid fauna (Echinodermata, Crinoidea) from the Lower Eocene of the Gulf of Languedoc (Corbières, Aude, southern France).

Detailed studies of the middle Ilerdian (lower Ypresian) blue marls of the Gulf of Languedoc (Corbières, Aude, France), belonging to the north Pyrenean foreland basin, have revealed a more abundant and diverse crinoid fauna than previously documented from the Lower Eocene. Here we describe five species of stalked crinoids in the family Rhizocrinidae (Cherbonniericrinus requiensis n. sp., ?Democrinus elongatus, Globulocrinus amphoraformis n. gen., n. sp., Pseudoconocrinus doncieuxi and P. lavadensis n. sp.), one barnacle-like species in the stalkless family Holopodidae (Holopus plaziati n. sp.) and a single feather star in the family Conometridae (Amphorometra atacica). Several sites have yielded brachials and rhizoids in addition to abundant aboral cups and columnals indicating in situ fossilisation of the dissociated skeletal elements. P. lavadensis n. sp. and ?D. elongatus have been collected only from outcrops located in the upper part of the middle blue marls, while P. doncieuxi predominates, with a wide range of morphological variation, in the lower blue marls. The fossil assemblage at the locality of Réqui near Montlaur differs from the others in the smaller size of most individuals and the presence of H. plaziati n. sp., C. requiensis n. sp., G. amphoraformis n. gen., n. sp., and P. doncieuxi suboblongus n. subsp. This particular association with high juvenile mortality corresponds to an unstable environment with mixed substrates (muddy and rocky). The crinoid fauna of the Corbières appears to be the most diverse of Early Eocene age known to date. With the fauna of the London Clay, a boreal formation of the same age, it shares the presence of the genera Democrinus and Amphorometra in an open-sea environment. A comparison with extant faunas allows the depth of deposition at the Ypresian sites in the Gulf of Languedoc to be estimated between from 100 and 140 meters.

The Cephalozygoptera, a new, extinct suborder of Odonata with new taxa from the early Eocene Okanagan Highlands, western North America.

We describe the Cephalozygoptera, a new, extinct suborder of Odonata, composed of the families Dysagrionidae and Sieblosiidae, previously assigned to the Zygoptera, and possibly the Whetwhetaksidae n. fam. The Cephalozygoptera is close to the Zygoptera, but differs most notably by distinctive head morphology. It includes 59 to 64 species in at least 19 genera and one genus-level parataxon. One species is known from the Early Cretaceous (Congqingia rhora Zhang), possibly three from the Paleocene, and the rest from the early Eocene through late Miocene. We describe new taxa from the Ypresian Okanagan Highlands of British Columbia, Canada and Washington, United States of America: 16 new species of Dysagrionidae of the existing genus Dysagrion (D. pruettae); the new genera Okanagrion (O. threadgillae, O. hobani, O. beardi, O. lochmum, O. angustum, O. dorrellae, O. liquetoalatum, O. worleyae, all new species); Okanopteryx (O. jeppesenorum, O. fraseri, O. macabeensis, all new species); Stenodiafanus (S. westersidei, new species); the new genus-level parataxon Dysagrionites (D. delinei new species, D. sp. A, D. sp. B, both new); and one new genus and species of the new family Whetwhetaksidae (Whetwhetaksa millerae).

Diversity, palaeoecology and palaeoenvironmental significance of the Eocene chondrichthyan assemblages of the Bolca Lagerstätte, Italy.

Over the last few years, the morphology, taxonomy and systematics of the cartilaginous fish taxa of the two main sites of the Bolca Lagerstätte, Italy, (Pesciara and Monte Postale sites) have been extensively discussed in a series of papers, resulting in a complete revision of this neglected component of the Eocene Tethyan ichthyofauna. Here, we provide a comprehensive overview of the diversity, palaeoecology and palaeoenvironmental significance of the two chondrichthyan assemblages of the Pesciara and Monte Postale sites. The assemblages include 14 shark species (Lamniformes and Carcharhiniformes) and batoids (Torpediniformes, Rhinopristiformes, Myliobatiformes, Platyrhinidae and Zanobatidae), as well as a single putative chimaeriform. The Pesciara and Monte Postale sites are characterized by eight chondrichthyan taxa each, but the taxonomic compositions are distinctly different reflecting the dissimilarities in the overall composition of both fish assemblages. Palaeoecological interpretations and habitat preferences of the two chondrichthyan assemblages are consistent with previously hypothesized palaeoenvironmental settings based on sedimentological, palaeontological and geochemical evidence. The chondrichthyan assemblages of the two sites appear to be constituted by ecologically vicariant taxa, with both characterized by a predominance of benthic species with durophagous/cancritrophic feeding modes. Taxonomic composition, habitat preferences and palaeobathymetric analyses support the hypothesis that both assemblages occupied tropical marine shallow waters (likely up to 50 m deep) of the inner portion of the Lessini Shelf. The taxonomic composition of both sites is considerably different from that of any other contemporaneous Tethyan and Boreal chondrichthyan assemblages.

Mosaic of plesiomorphic and derived characters in an Eocene myliobatiform batomorph (Chondrichthyes, Elasmobranchii) from Italy defines a new, basal body plan in pelagic stingrays.

BACKGROUND: End-Cretaceous niche-filling by benthic Mesozoic survivors resulted in a prominent increase of durophagous fish families, resulting in the appearance of the earliest representatives of several extant fish lineages, including the pelagic durophagous stingrays, a monophyletic clade of myliobatiform batoids that is characterized by a derived swimming mode and feeding habits. Although the earliest members appeared in the Late Cretaceous, most of the crown genera date back to the Eocene. RESULTS: In this study, we re-examine the anatomy of the Eocene eagle ray Promyliobatis gazolai (de Zigno), represented by two nearly complete and articulated specimens from the world-famous Ypresian Konservat-Lagerstätte of Bolca, in detail. This taxon exhibits a mosaic of plesiomorphic and derived characters (e.g. tail sting displaced posteriorly on the tail, at about 50-60% of tail length; pectoral fins joining in front of the head; anterior and posterior pectoral fin margins nearly straight; compagibus laminam absent; single, unfragmented mesopterygium) that clearly define a new body plan within the pelagic durophagous stingrays. CONCLUSIONS: The significant morphological differences between Promyliobatis and extant representatives of Myliobatidae, Aetobatidae, Rhinopteridae, and Mobulidae, support its placement as separate stem group member. The phylogenetic placement of Promyliobatis, based on skeletal and dental characters, strongly supports its basal position within pelagic stingrays. However, its position within the Myliobatiformes becomes unstable when stingray taxa known by fossil teeth only are included. A comparative analysis of the skeletal and tooth morphologies, as well as of the evolutionary trends of pelagic stingrays is also discussed.

Eocene sand tiger sharks (Lamniformes, Odontaspididae) from the Bolca Konservat-Lagerstätte, Italy: palaeobiology, palaeobiogeography and evolutionary significance.

Here we report the first record of one of the most common and widespread Palaeogene selachians, the sand tiger shark Brachycarcharias, from the Ypresian Bolca Konservat-Lagerstätte. The combination of dental character of the 15 isolated teeth collected from the Pesciara and Monte Postale sites (e.g. anterior teeth up to 25 mm with fairly low triangular cusp decreasing regularly in width; one to two pairs of well-developed lateral cusplets; root with broadly separated lobes; upper teeth with a cusp bent distally) supports their assignment to the odontaspidid Brachycarcharias lerichei (Casier, 1946), a species widely spread across the North Hemisphere during the early Palaeogene. The unambiguous first report of this lamniform shark in the Eocene Bolca Konservat-Lagerstätte improves our knowledge concerning the diversity and palaeobiology of the cartilaginous fishes of this palaeontological site, and provides new insights about the biotic turnovers that involved the high trophic levels of the marine settings after the end-Cretaceous extinction.

Early African Fossils Elucidate the Origin of Embrithopod Mammals.

Modern mammals rapidly evolved in the early Cenozoic in all continental provinces, including in Africa, with one of the first placental branches, the Afrotheria [1, 2]. Afrotherian evolution is at the origin of the major radiation of African ungulate-like mammals, including extant hyrax, elephant, and sea cow orders, which all belong to the Paenungulata. The paenungulate radiation also includes the extinct order Embrithopoda of uncertain interordinal relationships, which is best known for the giant and strangely specialized Oligocene genus Arsinoitherium. The Ouled Abdoun basin, Morocco, yielded exceptional Paleocene-Eocene fossils documenting the early paenungulate evolution [3-8]. Here we report two new small Ypresian species, Stylolophus minor n.g., n.sp. and cf. Stylolophus sp., which are the earliest and most primitive embrithopods. The cladistic analysis relates the Embrithopoda to crown paenungulates as the stem-group of the Tethytheria, which makes crown tethytherians restricted to extant elephant and sea cow orders. The Embrithopoda is therefore an early tethytherian offshoot predating the elephant and sea cow divergence. The resulting phylogeny supports a strictly African early radiation of the paenungulates excluding the Phenacolophidae and Anthracobunia. It sustains an at least early Paleocene African origin of the Embrithopoda. The unique tooth pattern of the embrithopods (hyperdilambdodont and pseudolophodont molars) is resolved as evolving early and directly from the dilambdodont (W-shaped labial molar crests) ancestral paenungulate morphotype. The specialized upper molar morphology with two transverse crests is convergent and non-homologous in embrithopods and crown Tethytheria. These convergences for specialized folivorous diet were driven by free herbivorous African niches in the early Paleogene.

A synoptic review of the Eocene (Ypresian) cartilaginous fishes (Chondrichthyes: Holocephali, Elasmobranchii) of the Bolca Konservat-Lagerstätte, Italy.

Here, we review and discuss the records and taxonomy of the Ypresian (Eocene) chondrichthyans from the famous Bolca Konservat-Lagerstätte in northeastern Italy. Despite the outstanding diversity and the numerous studies focusing on the actinopterygian faunas from Pesciara and Monte Postale, the current knowledge about the systematics, taxonomy and phylogenetic relationships of the cartilaginous fishes from these Eocene sites remains elusive and largely inadequate. The celebrated Eocene Bolca Lagerstätte has yielded several exquisitely preserved articulated remains of chondrichthyan fishes in which delicate structures and soft tissues are preserved, as well as isolated teeth. The cartilaginous fish assemblage of Bolca comprises at least 17 species-level taxa belonging to 10 families in 6 orders, including selachians (Carcharhiniformes, Lamniformes), batoids (Torpediniformes, Myliobatiformes, Rajiformes) and holocephalans (Chimaeriformes). The occurrence of holocephalans represented by an isolated fin-spine of the chimeroid Ischyodus in the Bolca assemblage is reported here for the first time and represents the first record of chimeroids in the Eocene of Italy and also southern Europe. The Bolca chondrichthyan assemblage is remarkably different from those of other contemporaneous Boreal or Tethyan deposits, suggesting that its taxonomic composition is largely influenced by the palaeoenvironmental context. However, this synoptic review also highlights the importance of detailed revisions of all chondrichthyan remains from the Bolca Konservat-Lagerstätten.

A new commelinid monocot seed fossil from the early Eocene previously identified as Solanaceae.

PREMISE OF THE STUDY: Fossils provide minimum age estimates for extant lineages. Here we critically evaluate Cantisolanum daturoides Reid & Chandler and two other early putative seed fossils of Solanaceae, an economically important plant family in the Asteridae. METHODS: Three earliest seed fossil taxa of Solanaceae from the London Clay Formation (Cantisolanum daturoides) and the Poole and Branksome Sand Formations (Solanum arnense Chandler and Solanispermum reniforme Chandler) were studied using x-ray microcomputed tomography (MCT) and scanning electron microscopy (SEM). KEY RESULTS: The MCT scans of Cantisolanum daturoides revealed a high level of pyrite preservation at the cellular level. Cantisolanum daturoides can be clearly excluded from Solanaceae and has more affinities to the commelinid monocots based on a straight longitudinal axis, a prominent single layer of relatively thin-walled cells in the testa, and a clearly differentiated micropyle surrounded by radially elongated and inwardly curved testal cells. While the MCT scans show no internal preservation in Solanum arnense and Solanispermum reniforme, SEM images show the presence of several characteristics that allow the placement of these taxa at the stem node of Solanaceae. CONCLUSIONS: Cantisolanum daturoides is likely a member of commelinid monocots and not Solanaceae as previously suggested. The earliest fossil record of Solanaceae is revised to consist of fruit fossil with inflated calyces from the early Eocene of Patagonia (52 Ma) and fossilized seeds from the early to mid-Eocene of Europe (48-46 Ma). The new identity for Cantisolanum daturoides does not alter a late Cretaceous minimum age for commelinids.

Eoalosa janvieri gen. et sp. nov., a new clupeid fish (Teleostei, Clupeiformes) from the Eocene of Monte Bolca, Italy.

Fishes of the family Clupeidae are extremely abundant in the Eocene fossiliferous limestone of Monte Bolca representing the most common group from this celebrated locality. A new clupeid from the Pesciara site, Eoalosa janvieri gen. et sp. nov., is described. The new taxon exhibits a unique combination of characters supporting its recognition as a new genus and species of clupeid fish that is tentatively placed in the subfamily Alosinae. The description of this new taxon improves our knowledge of the diversity of clupeoid fishes in the Eocene of Monte Bolca.

Weddellian marine/coastal vertebrates diversity from a basal horizon (Ypresian, Eocene) of the Cucullaea I Allomember, La Meseta formation, Seymour (Marambio) Island, Antarctica

The La Meseta Formation crops out in Seymour/Marambio Island, Weddell Sea, northeast of the Antarctic Peninsula and contains one of the world's most diverse assemblages of Weddellian marine/coastal vertebrates of Early Eocene (Ypresian) age. The La Meseta Formation is composed of poorly consolidated, marine sandstones and siltstones which were deposited in a coastal, deltaic and/or estuarine environment. It includes marine invertebrates and vertebrates as well as terrestrial vertebrates and plants. The highly fossiliferous basal horizon (Cucullaea shell bed, Telm 4 of Sadler 1988) of the Cucullaea I Allomember is a laterally extensive shell bed with sandy matrix. The fish remains, including 35 species from 26 families, of the Ypresian Cucullaea bed represent one of the most abundant and diverse fossil vertebrate faunas yet recorded in southern latitudes. Stratigraphic distribution and phylogenetic relationships of the Weddellian sphenisciforms are consistent with a first radiation of this group in the Early Eocene. The first inquestionable archaeocete from Antarctica is recorded in this unit and is referred to a new taxon

La Formación La Meseta aflora en la Isla Seymour/Marambio, Mar de Weddell, noreste de la Península Antártica y contiene una de las asociaciones de vertebrados costeros/marinos de edad Eoceno temprano (Ypresiano) más diversa que se conoce a nivel mundial. Esta unidad está compuesta por areniscas marinas pobremente consolidadas las cuales fueron depositadas en ambientes costeros, deltaicos y/o estuarinos. Esta incluye invertebrados y vertebrados marinos así como plantas y vertebrados terrestres. El horizonte basal (el banco de Cucullaea, Telm 4) del Alomiembro Cucullaea I es lateralmente extensor y altamente fosilífero. Los restos de peces del banco de Cucullaea (Ypresiano) incluyen 35 especies con 26 familias y representa una de las más abundantes y diversas fauna de vertebrados fósiles registradas en latitudes altas. La distribución estratigráfica y las relaciones filogenéticas de los pingüinos fósiles (Sphenisciformes) son consistentes con la primera radiación de este grupo en el Eoceno temprano. El primer incuestionable Archaeoceti de Antártida es registrado en esta unidad y es referido un nuevo taxón

A new genus and species of Heroini (Perciformes: Cichlidae) from the early Eocene of southern South America

The Lumbrera Formation is the uppermost unit of the Salta Group, which crops out in northwestern Argentina. The paleoenvironment of the Lumbrera Formation is interpreted as a perennial lake deposited under temperate climatic conditions during the early to middle Eocene. Its fossil content is made up of palynomorphs, insects, crocodiles, turtles, lizards, and mammals, besides an ichthyofauna formed by cichlids, poeciliids and dipnoans. åPlesioheros chauliodus is described based on a single individual from this formation, which was fossilized as a lateral view impression (missing anal and caudal fins). It can be distinguished from other cichlids by a moderately deep body, enlarged anterior dentary teeth bearing subapical cusp, a low abdominal vertebral count (10), five canal openings in the dentary, and XI + 12 dorsal-fin rays. A phylogenetic analysis, using the matrix by Kullander (1998), recovered åPlesioheros within Heroini. This species was recovered most closely related to Australoheros and to the deep-bodied South American heroins. The occurrence of an Eocene Heroini, as well as of other cichlid lineages in the same stratigraphical level, is evidence of an ancient diversification in this family. This ancient age supports the hypothesis that the Cichlidae originated on Gondwana.

A Formação Lumbrera é a unidade do topo do Grupo Salta, aflorante na região noroeste da Argentina. O paleoambiente da Formação Lumbrera tem sido interpretado como um lago perene depositado sob um clima temperado durante o início do Eoceno. Seu conteúdo fóssil é formado por palinomorfos, insetos, crocodilos, tartarugas, lagartos, mamíferos, além de uma ictiofauna que inclui ciclídeos, poeciliídeos e dipnóicos. åPlesioheros chauliodus é descrito com base em um único indivíduo coletado nesta formação, preservado como impressão em vista lateral (faltando as nadadeiras anal e caudal). Ele pode ser distinguido de outros ciclídeos por um corpo moderadamente alto, dentes anteriores do dentário aumentados e portando uma cúspide lingual subapical, uma baixa contagem de vértebras abdominais, cinco aberturas do canal sensorial no dentário, e XI + 12 raios na dorsal. Na análise filogenética, usando a matriz apresentada por Kullander (1998), åPlesioheros resultou dentro da Tribo Heroini mais relacionado a Australoheros e aos heroinis de corpo alto sul-americanos. A ocorrência de um heroini fóssil desta idade, bem como das outras linhagens de ciclídeos no mesmo nível estratigráfico, é uma evidência da diversificação inicial da família, corroborando fortemente a hipótese de que os Cichlidae se originaram no Gondwana ainda durante o Cretáceo.