First record of a tarsometatarsus of Tonsala hildegardae (Plotopteridae) and other avian remains from the late Eocene/early Oligocene of Washington State (USA)

New bird fossils from the late Eocene/early Oligocene Makah Formation and the Oligocene Pysht Formation on the Olympic Peninsula (Washington State, USA) are described. A partial skeleton from the Pysht Formation includes the first reported tarsometatarsus of Tonsala hildegardae Olson, 1980, a wing-propelled diving bird of the taxon Plotopteridae. It shows that Tonsala had a tarsometatarsus that was morphologically intermediate between that of the late Eocene Phocavis and more derived plotopterids. We introduce the new taxon Tonsalinae nov. subfam. for a clade including all named plotopterids except Phocavis, Plotopterum, and the recently described Stemec. We furthermore describe a partial plotopterid pelvis and a sternum from the Makah Formation. The sternum shows a close resemblance to that of extant Phalacrocoracoidea (cormorants and darters) and may be the earliest North American record of this taxon.     

A specimen of Eocuculus Chandler, 1999 (Aves, ? Cuculidae) from the early Oligocene of France

A postcranial skeleton of a small bird from the early Oligocene locality Pichovet in Southern France is described and identified as Eocuculus cf. cherpinae Chandler, 1999. It is the second fossil record of Eocuculus which was hitherto known from a postcranial skeleton from the late Eocene of North America only. Although Eocuculus shares some derived similarities with Cuculidae (cuckoos), it distinctly differs in a number of osteological features from crown group members of this taxon. If future, more complete skeletons prove its cuculiform affinities, Eocuculus is a stem lineage representative of this taxon and not within the crown group. Recognition of Eocuculus in the early Oligocene of France provides evidence for the presence of an extinct late Eocene/early Oligocene avian taxon with an intercontinental Northern Hemisphere distribution.     

A small stem-galliform bird (Aves: Paraortygidae) from the Eocene of Uzbekistan

Landfowl (order Galliformes) are among the most characteristic birds of the modern avian faunas, but their early evolutionary history is insufficiently known. The diversity of previously described Eocene galliforms implies a great role of Eocene diversification in the early evolution of this group. However, almost nothing is known about the Eocene diversity of galliforms in Asia, even though this large continent with a variety of habitats might have played a significant role in their early evolution. Here we describe a partial coracoid from the Lutetian–Bartonian of Uzbekistan, which is the oldest diagnosable galliform bird in Asia, and is further the first landbird known from this geographical area. The specimen displays a unique morphology with few autapomorphies and an unexpected combination of similarities with the African genus Scopelortyx and Eurasian Paraortyx, and hence is described as a new taxon, Xorazmortyx turkestanensis gen. et sp. nov., within the extinct family Paraortygidae, being the first Asian representative of this clade. Similarities with the African genus Scopelortyx indicate a connection of land bird faunas between northern Africa/Arabia and Asia in the middle Eocene. Better dispersal abilities of the early galliforms Paraortygidae are inferred from the osteology of their pectoral girdle and the humerus, which do not show adaptations to the powerful burst take off (escape flight), characteristic of most modern Phasianidae.     

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.     

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.     

A new pipoid frog (Anura,Pipimorpha) from the Paleogene of Patagonia. Paleobiogeographical implications

The aim of the present contribution is to describe a new genus and species of Pipoidea from the Huitrera Formation (Eocene) from Patagonia, Argentina. The new genus shows a unique combination of characters indicating that it is a valid taxon different from other pipimorphs, including the coeval Llankibatrachus truebae. The phylogenetic analysis resulted in the nesting of the new taxon within a previously unrecognized endemic clade of South American aglossans. This new clade turns out to be the sister-group of crown-group Pipidae. This phylogenetic proposal reinforces the hypothesis sustaining the dispersal of pipids between Africa and South America through an island chain or a continental bridge across the Atlantic Ocean by Early Tertiary times.     

A reassessment of Eocene parrotlike fossils indicates a previously undetected radiation of zygodactyl stem group representatives of passerines (Passeriformes)

In the past years, various Eocene fossil birds were described as stem group representatives of the zygodactyl Psittaciformes (parrots). These birds show quite disparate morphologies, which cast some doubt on the correct assignment of all of them to the psittaciform stem group. A reassessment of their affinities is further needed, because it was recently proposed that among extant birds, Psittaciformes and Passeriformes (passerines) form a clade and that passerines possibly derived from a zygodactyl ancestor. Here, phylogenetic analyses are performed, which for the first time also include representatives of the Zygodactylidae, the extinct zygodactyl sister taxon of the Passeriformes. The early Eocene Psittacopes was originally described as a stem group representative of Psittaciformes. However, none of the present analyses supported psittaciform affinities for Psittacopes and instead recovered this taxon in a clade together with zygodactylids and passerines. Also part of this clade are the early Eocene taxa Pumiliornis and Morsoravis, and it is detailed that Psittacopes and the long‐beaked and presumably nectarivorous Pumiliornis, with which it has not yet been compared, are very similar in their postcranial osteology. The present analysis corroborates the hypothesis of a zygodactyl stem species of passerines. To account for these results, Psittacopes is here assigned to a new higher‐level taxon and a new name is also introduced for the clade including Zygodactylidae and Passeriformes.     

Lizards of the family Arretosauridae Gilmore, 1943 (Iguanomorpha, Iguania) from the Paleogene of Mongolia

The family Arretosauridae is an enigmatic group of Asian iguanians, which contains one species, Arretosaurus ornatus Gilmore, 1943 from the Late Eocene of Inner Mongolia (China). In this work, the fossil family is classified within the taxon Iguanomorpha (Iguanidae sensu lato) and supplemented with three new forms from the Middle Eocene and Early Oligocene of Mongolia and the previously described Hemishinisaurus latifrons from the Late Eocene of China, which was originally assigned to Anguimorpha (Xenosauridae). New data characterize Arretosauridae as an important element of the Paleogene herpetofauna of Asia.     

A New Large-Bodied Oviraptorosaurian Theropod Dinosaur from the Latest Cretaceous of Western North America

The oviraptorosaurian theropod dinosaur clade Caenagnathidae has long been enigmatic due to the incomplete nature of nearly all described fossils. Here we describe Anzu wyliei gen. et sp. nov., a new taxon of large-bodied caenagnathid based primarily on three well-preserved partial skeletons. The specimens were recovered from the uppermost Cretaceous (upper Maastrichtian) Hell Creek Formation of North and South Dakota, and are therefore among the stratigraphically youngest known oviraptorosaurian remains. Collectively, the fossils include elements from most regions of the skeleton, providing a wealth of information on the osteology and evolutionary relationships of Caenagnathidae. Phylogenetic analysis reaffirms caenagnathid monophyly, and indicates that Anzu is most closely related to Caenagnathus collinsi, a taxon that is definitively known only from a mandible from the Campanian Dinosaur Park Formation of Alberta. The problematic oviraptorosaurs Microvenator and Gigantoraptor are recovered as basal caenagnathids, as has previously been suggested. Anzu and other caenagnathids may have favored well-watered floodplain settings over channel margins, and were probably ecological generalists that fed upon vegetation, small animals, and perhaps eggs.     

A small loon and a new species of large owl from the Rupelian of Belgium (Aves: Gaviiformes,Strigiformes)

The early Oligocene Boom Formation in Belgium yielded many avian remains from the Rupelian unit-stratotype, most of which have remained unstudied so far. Here, I describe a small loon (Gaviiformes) and a new species of large owl (Strigiformes) that are represented by associated bones of a single individual each. The loon, of which wing and pectoral girdle bones are preserved, is assigned to Colymboides (?) metzleri, a species previously known from a partial skeleton from the Rupelian of Germany. The owl is based on a tarsometatarsus and distal tibiotarsus and described as a new species, Selenornis steendorpensis. It constitutes the most substantial fossil record of the taxon Selenornis, which was before known from a distal tibiotarsus from an unknown horizon of the Quercy fissure fillings in southwestern France. It is detailed that there are differences in the higher level taxonomic composition of the known early Oligocene avifaunas of northern and southern Europe, which may reflect true zoogeographic facts owing to a different climate and vegetation.     

Uniquely derived upper molar morphology of Eocene Amphipithecidae (Primates: Anthropoidea): Homology and phylogeny

The extinct Southeast Asian primate family Amphipithecidae is regularly cited in discussions of anthropoid origins, but its phylogenetic position remains controversial. In part, the lack of consensus regarding amphipithecid relationships can be attributed to uncertainty regarding the homology of upper molar structures in this group. Here, we describe a virtually pristine upper molar of Pondaungia cotteri from the late middle Eocene Pondaung Formation of Myanmar, which is the first example of a relatively unworn and well-preserved amphipithecid upper molar ever recovered. The distolingual upper molar cusp in this new specimen of Pondaungia appears to be a lingually displaced and enlarged metaconule, rather than a hypocone or pseudohypocone as previous workers have thought. Reassessment of the upper molar morphology of other amphipithecids and putative amphipithecids reveals a very similar pattern in Siamopithecus, Myanmarpithecus and Ganlea, all of which are interpreted as having upper molars showing many of the same derived features apparent in Pondaungia. In contrast, the upper molar morphology of Bugtipithecus diverges radically from that of undoubted amphipithecids, and the latter taxon is excluded from Amphipithecidae on this basis. Phylogenetic analyses of several character–taxon matrices culled from the recent literature and updated to reflect the new information on amphipithecid upper molar morphology yield similar results. Consensus tree topologies derived from these analyses support amphipithecid monophyly and stable relationships within Amphipithecidae. Amphipithecids appear to be stem members of the anthropoid clade.     

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

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

Craniodental morphology and systematics of a new family of hystricognathous rodents (Gaudeamuridae) from the late eocene and early oligocene of Egypt

Background

Gaudeamus is an enigmatic hystricognathous rodent that was, until recently, known solely from fragmentary material from early Oligocene sites in Egypt, Oman, and Libya. Gaudeamus'' molars are similar to those of the extant cane rat Thryonomys, and multiple authorities have aligned Gaudeamus with Thryonomys to the exclusion of other living and extinct African hystricognaths; recent phylogenetic analyses have, however, also suggested affinities with South American caviomorphs or Old World porcupines (Hystricidae).

Methodology/Principal Findings

Here we describe the oldest known remains of Gaudeamus, including largely complete but crushed crania and complete upper and lower dentitions. Unlike younger Gaudeamus species, the primitive species described here have relatively complex occlusal patterns, and retain a number of plesiomorphic features. Unconstrained parsimony analysis nests Gaudeamus and Hystrix within the South American caviomorph radiation, implying what we consider to be an implausible back-dispersal across the Atlantic Ocean to account for Gaudeamus'' presence in the late Eocene of Africa. An analysis that was constrained to recover the biogeographically more plausible hypothesis of caviomorph monophyly does not place Gaudeamus as a stem caviomorph, but rather as a sister taxon of hystricids.

Conclusions/Significance

We place Gaudeamus species in a new family, Gaudeamuridae, and consider it likely that the group originated, diversified, and then went extinct over a geologically brief period of time during the latest Eocene and early Oligocene in Afro-Arabia. Gaudeamurids are the only known crown hystricognaths from Afro-Arabia that are likely to be aligned with non-phiomorph members of that clade, and as such provide additional support for an Afro-Arabian origin of advanced stem and basal crown members of Hystricognathi.     

A new tarkadectine primate from the Eocene of Inner Mongolia,China: phylogenetic and biogeographic implications

Tarka and Tarkadectes are Middle Eocene mammals known only from the Rocky Mountains region of North America. Previous work has suggested that they are members of the Plagiomenidae, an extinct family often included in the order Dermoptera. Here we describe a new primate, Tarkops mckennai gen. et sp. nov., from the early Middle Eocene Irdinmanha Formation of Inner Mongolia, China. The new taxon is particularly similar to Tarka and Tarkadectes, but it also displays many features observed in omomyids. A phylogenetic analysis based on a data matrix including 59 taxa and 444 dental characters suggests that Tarkops, Tarka and Tarkadectes form a monophyletic group—the Tarkadectinae—that is nested within the omomyid clade. Within Omomyidae, tarkadectines appear to be closely related to Macrotarsius. Dermoptera, including extant and extinct flying lemurs and plagiomenids, is recognized as a clade nesting within the polyphyletic group of plesiadapiforms, therefore supporting the previous suggestion that the relationship between dermopterans and primates is as close as that between plesiadapiforms and primates. The distribution of tarkadectine primates on both sides of the Pacific Ocean basin suggests that palaeoenvironmental conditions appropriate to sustain primates occurred across a vast expanse of Asia and North America during the Middle Eocene.     

The first New World occurrence of the Eocene birdPlesiocathartes (Aves: ?Leptosomidae)

Two new species of the Eocene avian taxonPlesiocathartes (Aves: ?Leptosomidae) are described. They are similar in size and overall appearance to other members of the genus, as well as to the extant Madagascan cuckoo-rollerLeptosomus discolor. So far, putative fossil Leptosomidae have been found only in the Eocene European de-posits of Messel, Geiseltal, Quercy, and London Clay. Both new species are from the Lower Eocene Green River Formation (USA) and thus the first New World representatives of this taxon.     

A New Species of Pengornithidae (Aves: Enantiornithes) from the Lower Cretaceous of China Suggests a Specialized Scansorial Habitat Previously Unknown in Early Birds

We describe a new enantiornithine bird, Parapengornis eurycaudatus gen. et sp. nov. from the Lower Cretaceous Jiufotang Formation of Liaoning, China. Although morphologically similar to previously described pengornithids Pengornis houi, Pengornis IVPP V18632, and Eopengornis martini, morphological differences indicate it represents a new taxon of the Pengornithidae. Based on new information from this specimen we reassign IVPP V18632 to Parapengornis sp. The well preserved pygostyle of the new specimen elucidates the morphology of this element for the clade, which is unique in pengornithids among Mesozoic birds. Similarities with modern scansores such as woodpeckers may indicate a specialized vertical climbing and clinging behavior that has not previously been inferred for early birds. The new specimen preserves a pair of fully pennaceous rachis-dominated feathers like those in the holotype of Eopengornis martini; together with the unique morphology of the pygostyle, this discovery lends evidence to early hypotheses that rachis-dominated feathers may have had a functional significance. This discovery adds to the diversity of ecological niches occupied by enantiornithines and if correct reveals are remarkable amount of locomotive differentiation among Enantiornithes.     

A tiny stem group representative of Pici (Aves, Piciformes) from the early Oligocene of the Czech Republic

We describe a new stem group representative of Pici (woodpeckers, honeyguides, barbets, and allies) from the early Oligocene (Rupelian) of the Czech Republic. The holotype of Picavus litencicensis, gen. et sp. nov. is a postcranial skeleton with well-preserved feather remains. The new species is distinguished from crown group Pici in several plesiomorphic features and is assigned to the new taxon Picavidae, fam. nov. The absence of an elongated accessory trochlea for the reversed fourth toe shows P. litencicensis to be the most basal representative of Pici, and concerning the morphology of the distal tarsometatarsus the species represents a morphological link between Pici and their sister taxon, the Galbulae. Like all other early Oligocene piciform birds, P. litencicensis is a tiny bird, the size of the smallest extant Pici. Because all Palaeogene Pici were found in Europe and some of these are outside the crown group, an Old World origin of Pici is likely. With definitive crown group representatives of Pici being unknown before the late Oligocene/early Miocene, the fossil record does not support earlier molecular calibrations, which resulted in a late Cretaceous divergence of crown group Pici.     

A small, “wader-like” bird from the Early Eocene of Messel (Germany)

A small bird with a shorebird-like beak and a very long hind toe is described from the Early Eocene of Messel in Germany. Vanolimicola longihallucis, gen. et sp. nov. is one of the few candidate species for a “wading bird” from Messel, that is, a bird that may have foraged along the shoreline of the ancient lake. A few features indicate a relationship to charadriiform birds and Jacanidae (jacanas) in particular, but charadriiform affinities are only weakly supported. In the preserved skeletal elements, V. longihallucis also resembles the taxon Songzia from the Early Eocene of China, which is considered to be a representative of the Ralloidea (rails and allies). A potential biogeographic significance of the new Messel fossil is strengthened by similar fossils from the North American Green River Formation.     

A new Late Eocene primate from the Krabi Basin (Thailand) and the diversity of Palaeogene anthropoids in southeast Asia

According to the most recent discoveries from the Middle Eocene of Myanmar and China, anthropoid primates originated in Asia rather than in Africa, as was previously considered. But the Asian Palaeogene anthropoid community remains poorly known and inadequately sampled, being represented only from China, Myanmar, Pakistan and Thailand. Asian Eocene anthropoids can be divided into two distinct groups, the stem group eosimiiforms and the possible crown group amphipithecids, but the phylogenetic relationships between these two groups are not well understood. Therefore, it is critical to understand their evolutionary history and relationships by finding additional fossil taxa. Here, we describe a new small-sized fossil anthropoid primate from the Late Eocene Krabi locality in Thailand, Krabia minuta, which shares several derived characters with the amphipithecids. It displays several unique dental characters, such as extreme bunodonty and reduced trigon surface area, that have never been observed in other Eocene Asian anthropoids. These features indicate that morphological adaptations were more diversified among amphipithecids than was previously expected, and raises the problem of the phylogenetic relations between the crown anthropoids and their stem group eosimiiforms, on one side, and the modern anthropoids, on the other side.