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.     

A SKULL OF THE GIANT BONY‐TOOTHED BIRD DASORNIS (AVES: PELAGORNITHIDAE) FROM THE LOWER EOCENE OF THE ISLE OF SHEPPEY

Abstract: The first substantial skull of a very large Paleogene bony‐toothed bird (Pelagornithidae) is described from the Lower Eocene London Clay of the Isle of Sheppey in England. The specimen is assigned to Dasornis emuinus (Bowerbank), based on a taxonomic revision of the large London Clay Pelagornithidae. Very large bony‐toothed birds from the London Clay were known previously from fragmentary remains of non‐comparable skeletal elements only, and Dasornis londinensis Owen, Argillornis emuinus (Bowerbank), A. longipennis Owen, and Neptuniavis miranda Harrison and Walker are considered junior synonyms of D. emuinus. The new specimen allows a definitive assignment of Dasornis to the Pelagornithidae and documents that this taxon closely resembles other bony‐toothed birds in cranial morphology. It is hypothesized that giant size (i.e. a wingspan above 4 m) evolved only once within Pelagornithidae and that Dasornis emuinus is the sister taxon of the giant Neogene bony‐toothed birds, which share a derived wing morphology.     

A stem lineage representative of buttonquails from the Lower Oligocene of Germany – fossil evidence for a charadriiform origin of the Turnicidae

A new species of the charadriiform taxon Turnipax Mayr, 2000 is described from the Lower Oligocene fossil site Frauenweiler in southern Germany. The postcranial skeleton assigned to Turnipax oechslerorum sp. nov. is very well preserved and allows the recognition of significant, previously unknown osteological details of Turnipax , especially concerning the wing and pectoral girdle bones. We provide evidence that Turnipax is a stem lineage representative of the Turnicidae (buttonquails) and synonymize Turnipacidae Mayr, 2000 with Turnicidae Gray, 1840. Turnipax is the earliest fossil representative of the Turnicidae, which otherwise have no Paleogene fossil record. Because recent molecular studies support a charadriiform origin of buttonquails, the mosaic distribution in the skeleton of Turnipax of derived features of the Turnicidae and non-turnicid charadriiform birds is of particular interest. Turnipax exhibits a more plesiomorphic morphology than extant Turnicidae, and we assume that its habitat and way of living differed from that of crown group Turnicidae, which may not have diversified before the spread of grasslands during the Oligocene and Miocene.     

Fight club: a unique weapon in the wing of the solitaire,Pezophaps solitaria (Aves: Columbidae), an extinct flightless bird from Rodrigues,Mascarene Islands

The solitaire (Columbidae; Pezophaps solitaria) of Rodrigues was an extinct giant flightless pigeon and the sister taxon to the dodo (Columbidae; Raphus cucullatus) from neighbouring Mauritius. The appearance and behaviour of the solitaire was recorded in detail by two observers before it became extinct in the mid 1700s. They described a prominent wing structure termed the ‘musket ball’ (carpal knob), which was used as a weapon and to produce an audible signal by either sex in territorial combat. Our study of subfossil solitaire bones from cave localities shows that the carpal knob formed after skeletal maturity, and reached its greatest size in adult males. We describe the morphology of the carpal knob, including its histology in thin section. It is an outgrowth of the processus extensorius of the carpometacarpus, but differs morphologically from homologous structures in other bird taxa, and thus is unique in Aves. We also compare the pectoral and wing osteology of the solitaire with that of the dodo, which had a similar morphology, but lacked any bony outgrowths on the wing. Furthermore, we suggest some biological and environmental factors leading to the evolution of this remarkable and unique carpal weapon. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 32–44.     

First partial face and upper dentition of the Middle Miocene hominoid Dryopithecus fontani from Abocador de Can Mata (Vallès‐Penedès Basin,Catalonia, NE Spain): Taxonomic and phylogenetic implications

A well‐preserved 11.8‐million‐years‐old lower face attributed to the seminal taxon Dryopithecus fontani (Primates, Hominidae) from the Catalan site ACM/C3‐Ae of the Hostalets de Pierola area (Vallès‐Penedès Basin, Catalonia, NE Spain) is described. The new data indicate that D. fontani is distinct at the genus level from Late Miocene European taxa previously attributed to Dryopithecus, which are here reassigned to Hispanopithecus. The new facial specimen also suggests that D. fontani and the Middle Miocene Pierolapithecus catalaunicus are not synonymous. Anatomical and morphometric analyses further indicate that the new specimen shows a combination of lower facial features—hitherto unknown in Miocene hominoids—that resembles the facial pattern of Gorilla, thus providing the first nondental evidence of gorilla‐like lower facial morphology in the fossil record. Considering the current evidence, the gorilla‐like facial pattern of D. fontani is inferred to be derived relative to previously known stem hominids, and might indicate that this taxon is either an early member of the Homininae or, alternatively, a stem hominid convergent with the lower facial pattern of Gorilla. The biogeographic implications of both alternatives are discussed. This new finding in the Hostalets de Pierola section reinforces the importance of this area for understanding the elusive question of the Middle Miocene origin and early radiation of great apes. Am J Phys Anthropol, 2009. © 2009 Wiley‐Liss, Inc.     

New specimens of the early Eocene bird Vastanavis and the interrelationships of stem group Psittaciformes

We describe new skeletal elements of Vastanavis from the early Eocene Cambay Shale Formation of western India, including a small coracoid that represents an unnamed new species, and comment on the relationships between this avian taxon and the recently described Avolatavis from the early Eocene Green River Formation in North America. Like the previously described ones, the new Vastanavis bones resemble those of the late Eocene Quercypsittidae, thus strengthening psittaciform affinities of the Indian taxon. Vastanavis differs from Avolatavis in the presence of a crista medianoplantaris on the tarsometatarsus and in claw morphology, but a fossil from the early Eocene London Clay, which was previously assigned to Vastanavidae, closely resembles Avolatavis in these features and all other osteological aspects. We show that most branches in a recent phylogeny of stem group Psittaciformes collapse after modification of a single erroneous character scoring for Vastanavis. We further describe a morphologically distinctive distal humerus of a small bird resembling the stem group nyctibiid Paraprefica, which was discovered in the most recent excavation in Vastan Lignite Mine.     

Cranial osteology of Exostinus serratus (Squamata: Anguimorpha), fossil sister taxon to the enigmatic clade Xenosaurus

Within the squamate clade Anguimorpha, Xenosaurus is an enigmatic taxon combining several apparently primitive features with a highly specialized set of autapomorphies. This combination makes the fossil record along the Xenosaurus stem particularly important for resolving the relationships of Xenosaurus with other anguimorphs, and between species of Xenosaurus.Exostinus serratus Cope, 1873 from the Oligocene of the western United States is currently hypothesized to be the immediate sister taxon to Xenosaurus. The cranial osteology of this pivotal taxon is described here for the first time from all known material, using high‐resolution X‐ray computerized tomography (CT) scanning to visualize individual elements. Exostinus serratus displays a mosaic of ancestral anguimorph features that are transformed in Xenosaurus, as well as unique synapomorphies shared with Xenosaurus. The region of the external nares is less transformed than in Xenosaurus, as are the general proportions of the bones surrounding the cartilaginous nasal capsule. However, the forms of the teeth and of the osteodermal sculpture, as well as several details of maxillary and mandibular morphology, are distinctly Xenosaurus‐like. Several autapomorphies are also present, including an abbreviated tooth row and an unusually wide palatal shelf of the maxilla. An understanding of the ways in which the non‐ossified tissues of the head influenced the development of the bones is crucial to interpreting their morphology. Furthermore, subtle anatomical features often provide important comparative information, and are emphasized herein. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 159 , 921–953.     

Middle Eocene Pelagornithidae and Gaviiformes (Aves) from the Ukrainian Paratethys

Abstract: We describe avian remains from Novopskov, a new middle Eocene marine locality in Ukraine. The fossils constitute the most substantial collection of Palaeogene bird bones from Eastern Europe and contribute to a better knowledge of the Paratethyan seabird fauna. Most of the specimens belong to Pelagornithidae (bony‐toothed birds), and two species of very different size can be distinguished. The larger of these is tentatively referred to Dasornis sp., the smaller to Odontopteryx toliapica. The specimens include skeletal elements that were not described for Palaeogene bony‐toothed birds and document previously unknown morphological differences between Palaeogene and Neogene Pelagornithidae. It is argued that the purported crane Eobalearica tugarinovi, from the middle Eocene of Kyrgyzstan, is probably also a bony‐toothed bird. A new genus and species of small Gaviiformes, Colymbiculus udovinchenkoi, is described, which is the earliest fossil record of a loon from Europe, preceding the next oldest specimens by more than 10 myr. The Ukrainian fossils document profound differences between middle Eocene and extant marine avifaunas of Europe, and whereas the middle Eocene Paratethyan avifauna appears to have been similar to that of the North Sea with regard to pelagornithid diversity, the absence of prophaethontids and relative abundance of Gaviiformes may indicate faunistic differences concerning the remaining seabirds.     

Skeletal morphology and the phylogeny of skuas (Aves: Charadriiformes,Stercorariidae)

The skuas (Aves: Charadriiformes, Stercorariidae) consist of two assemblages. On the basis of size, plumage, and distributional similarities, each of the two assemblages has long been considered monophyletic, and this traditional hypothesis has commonly been manifested in the recognition of two genera, Stercorarius and Catharacta; conversely, more recently collected molecular and ectoparasite evidence yields an alternative hypothesis, in which one member of Stercorarius, Stercorarius pomarinus, is more closely related to the forms in Catharacta than to the other Stercorarius sp. In this study we used skeletal morphology to test the competing hypotheses of skua phylogeny. Cladistic analysis of 141 osteological characters provided strong support for the molecular/ectoparasite hypothesis. However, those skeletal data did not support a sister‐taxon relationship between S. pomarinus and Catharacta skua, as inferred from mitochondrial DNA sequence data; instead, they resolved pomarinus as the sister of a monophyletic Catharacta. Additionally, our skeletal evidence did not support a sister‐group relationship between skuas and auks, as constraining skua/auk monophyly increased the tree length by nearly 5%. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 157 , 612–621.     

The sculpture and morphology of postcranial dermal armor plates and associated bones in gasterosteiforms and syngnathiforms inhabiting Estonian coastal waters

Lees, J., Märss, T., Wilson, M. V. H., Saat, T. and ?pilev, H. 2011. The sculpture and morphology of postcranial dermal armor plates and associated bones in gasterosteiforms and syngnathiforms inhabiting Estonian coastal waters. ―Acta Zoologica (Stockholm) 93 : 422–435. Five fish species inhabiting Estonian coastal waters (Gasterosteus aculeatus L., Pungitius pungitius (L.), and Spinachia spinachia (L.) of the order Gasterosteiformes and Syngnathus typhle L. and Nerophis ophidion (L.) of the order Syngnathiformes) are described on the basis of the sculpture and morphology of their postcranial dermal armor plates, as revealed and illustrated by SEM images. This study shows that the shapes of these superficial skeletal elements vary by species as well as by their position on the body, whereas the sculpture on the bones is taxon specific. The detailed features allow the identification of isolated fossil and subfossil remains and show promise for future systematics studies.     

Diversification in an Early Cretaceous avian genus: evidence from a new species of Confuciusornis from China

A new species of Confuciusornis, the oldest known beaked bird, is erected based on a nearly complete fossil from the Early Cretaceous Yixian Formation of western Liaoning, northeast China. C. feducciai is the largest and shows the highest ratio of the forelimb to the hindlimb among all known species of Confuciusornis. The skeletal qualitative autapomorphies, including a V-shaped furcula, a rectangular deltopectoral crest, the absence of an oval foramen at the proximal end of the humerus, the very slender alular digit, a relatively much longer ischium which is two-thirds the length of the pubis, and the morphology of sternum, strongly suggest the new specimen is a valid distinctive taxon. Detailed comparison with other described species provides sound evidence for diversification in the Early Cretaceous avian genus Confuciusornis. Anatomical features suggest an arboreal habit of the new bird.     

PHYLOGENY AND FORELIMB DISPARITY IN WATERBIRDS

Previous work has shown that the relative proportions of wing components (i.e., humerus, ulna, carpometacarpus) in birds are related to function and ecology, but these have rarely been investigated in a phylogenetic context. Waterbirds including “Pelecaniformes,” Ciconiiformes, Procellariiformes, Sphenisciformes, and Gaviiformes form a highly supported clade and developed a great diversity of wing forms and foraging ecologies. In this study, forelimb disparity in the waterbird clade was assessed in a phylogenetic context. Phylogenetic signal was assessed via Pagel's lambda, Blomberg's K, and permutation tests. We find that different waterbird clades are clearly separated based on forelimb component proportions, which are significantly correlated with phylogeny but not with flight style. Most of the traditional contents of “Pelecaniformes” (e.g., pelicans, cormorants, and boobies) cluster with Ciconiiformes (herons and storks) and occupy a reduced morphospace. These taxa are closely related phylogenetically but exhibit a wide range of ecologies and flight styles. Procellariiformes (e.g., petrels, albatross, and shearwaters) occupy a wide range of morphospace, characterized primarily by variation in the relative length of carpometacarpus and ulna. Gaviiformes (loons) surprisingly occupy a wing morphospace closest to diving petrels and penguins. Whether this result may reflect wing proportions plesiomorphic for the waterbird clade or a functional signal is unclear. A Bayesian approach detecting significant rate shifts across phylogeny recovered two such shifts. At the base of the two sister clades Sphenisciformes + Procellariiformes, a shift to an increase evolutionary rate of change is inferred for the ulna and carpometacarpus. Thus, changes in wing shape begin prior to the loss of flight in the wing‐propelled diving clade. Several shifts to slower rate of change are recovered within stem penguins.     

Thoracic morphology of a flightless mexican grasshopper,Barytettix psolus: Comparison with the locust,Schistocerca gregaria

The thoracic morphology of a flightless grasshopper, Barytettix psolus, is described and compared to that of locusts, Schistocerca gregaria, to evaluate modifications to skeleton, muscles, and the nervous system which have accompanied secondary loss of flight. Barytettix lacks hindwings, has immobile vestiges of forewings and is devoid of skeletal specializations for wing movement and flight. Its pterothoracic musculature resembles that of Schistocerca except for the absence of those muscles which, in locusts, have the primary function of moving the wings, the dorsal longitudinal, tergosternal, first basalar, pleuroalar, and dorsal accessory muscles. Pterothoracic ganglia of Barytettix resemble those of Schistocerca in their gross features, number, and primary branching pattern of nerves, with differences in detail relating to reduction of the flight muscles. The combination of features exhibited in Barytettix represents an extreme reduction in the specialization for wing movements and flight displayed by most acridids, at a level which exceeds that of many brachypterous and some apterous species. While skeletal fusion and loss of muscles indicate loss of flight, the accompanying thoracic stiffening and increase in overall body density may promote more efficient jumping as a means of locomotion.     

Guyiling jianboni gen. et sp.n., an antlion‐like lacewing,illuminating homologies and transformations in Neuroptera wing venation

A new genus and species of antlion‐like fossil lacewing, Guyiling jianboni gen. et sp.n. (Insecta: Neuroptera) are described based on a single specimen from the Yixian Formation (Liaoning Province, China; Early Cretaceous). The new taxon exhibits derived traits such as distally dilated antennae and well‐developed anterior Banksian line (known in Myrmeleontidae), but also genuine plesiomorphies (at the level of Myrmeleontiformia), such as the divergence of a distinct CuA1 stem from MP2 + CuA1 (forewing), and a basal origin of MA (diverging from RP + MA; both wing pairs). This combination is unique among the ‘Palaeoleontidae’, a group better considered as a paraphyletic assemblage of various stem‐Myrmeleontiformia. The wing morphology of the new species is considered in the light of a survey of wing venation topological homologies (and implied transformations) among several Neuroptera families. The survey includes a revision of the holotype of Leptolingia jurassica Ren (Grammolingiidae; Jiulongshan Formation, Daohugou locality, Inner Mongolia Autonomous Region, China; middle Jurassic). The forewing morphology of Guyiling jianboni gen. et sp.n. demonstrates that the fusion of MP2 with CuA, and the differentiation of CuA into two distinct main stems (namely CuA1 and CuA2) are traits shared with Myrmeleontidae and Ascalaphidae (and possibly Nemopteridae). However, the survey also demonstrates that a fusion of MP2 with CuA occurred repeatedly among Neuropterida, although by means of various modalities (namely translocation vs regular fusion). The ‘pectinate fusion’ of CuA(1) with MP2 [i.e. CuA1 has no distinct stem and emits successive branches from MP2 + CuA(1) partim] is a further step in this fusion, and occurred repeatedly as well (at least in the hind wings of Osmylidae and Nymphidae, and both fore‐ and hindwings of a sub‐group of Myrmeleontiformia including Myrmeleontidae and Ascalaphidae, and possibly Nemopteridae). It is anticipated that the current contribution will constitute useful background information for further studies, focusing on particular transformation cases, ideally including a documentation of intraspecific variation.     

Alchemilla austriaca,spec. nova,(Rosaceae) — eine neue Art aus den Ostalpen

Alchemilla austriaca is a new species which belongs to the group ofA. demissa, A. frigens, A. longana, A. longiuscula, A. semisecta, andA. sinuata. The holotype specimen as well as leaf and flower details are illustrated (Figs. 1–3). A complete character analysis is given, differences and similarities of allied species are presented in two tables, and the position of the group within the genus is discussed.A. austriaca so far is known only from the Austrian Alps and mainly from the central ranges (distribution map: Fig. 4). Its wet subalpine and alpine habitats are characterized by species lists.

     

A partial skeleton of a new fossil loon (Aves, Gaviiformes) from the early Oligocene of Germany with preserved stomach content

A partial skeleton of a new fossil loon (Aves, Gaviiformes), ?Colymboides metzleri n.sp., is described from the early Oligocene (Rupelian) of Frauenweiler in Germany. The new species resembles the early Miocene species Colymboides minutus in size and overall morphology, but differs in several osteological details. The specimen represents the first associated remains of an early Tertiary loon. Preserved stomach content further provides the first direct evidence that early Tertiary loons were already specialized towards a piscivorous diet, hunting fishes in a marine environment.Communicated by F. Bairlein     

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.     

A new basal ornithuromorph bird (Aves: Ornithothoraces) from the Early Cretaceous of China with implication for morphology of early Ornithuromorpha

Ornithuromorpha is the most derived avian group in the Early Cretaceous, advanced members of which encompass all living birds (Neornithes). Here we report on a new basal ornithuromorph bird, Bellulia rectusunguis gen. et sp. nov., represented by a nearly complete skeleton from the Early Cretaceous Jehol Biota in northeastern China. A comprehensive phylogenetic analysis resolved the new taxon in a basal position that is only more derived than Archaeorhynchus and Jianchangornis among ornithuromorphs, increasing the morphological diversity of basal ornithuromorphs. The new specimen has a V‐shaped furcula with a short hypocleidium, a feature otherwise known only in Schizooura among Cretaceous ornithuromorphs. We discuss the implications of the new taxon on the evolution of morphology of primitive ornithuromorphs, particularly of pectoral girdle, sternum and limb proportion pertaining to powered flight. The preserved gastroliths and pedal morphology indicate herbivory and lakeshore adaption for this new species. © 2015 The Linnean Society of London     

New Specimens of Yanornis Indicate a Piscivorous Diet and Modern Alimentary Canal

A crop adapted for an herbivorous diet of seeds has previously been documented in the Early Cretaceous birds Sapeornis and Hongshanornis. Here we report on several specimens of Yanornis that preserve a crop containing fish. One specimen preserves two whole fish in the oesophagus, indicating that Early Cretaceous birds shared trophic specializations with Neornithes for the increased energetic demands of flight – namely the storing of food for later consumption when the stomach is full. Whole fish also indicate that despite their presence, teeth were not used to orally process food, suggesting the hypertrophied dentition in this taxon were utilized in prey capture. The presence of macerated fish bones in the crop of other specimens indicates the highly efficient advanced muscular system of peristalsis responsible for moving ingested items between different segments of the alimentary canal was also in place. Despite the fact many features of the modern avian alimentary canal are inferred to compensate for the absence of teeth in birds (expandable oesophagus, grinding gizzard), the derived alimentary canal was apparently present in toothed Cretaceous birds. Although Yanornis was considered to have switched their diet from piscivorous to herbivorous, based on position and morphology we reinterpret the gastroliths reported in one specimen as sand impacted in the intestines, and reconstruct the taxon as primarily piscivorous. This is a novel interpretation for fossilized gastroliths, and the first documentation of this condition in the fossil record.