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     

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.     

New specimens of the early Eocene stem group galliform Paraortygoides (Gallinuloididae), with comments on the evolution of a crop in the stem lineage of Galliformes

Two new specimens of the fossil stem group galliform Paraortygoides messelensis Mayr 2000 (Gallinuloididae) are described from the Middle Eocene of Messel in Germany, including a complete skeleton in which the hitherto unknown skull of this species is preserved. The shorter and more protruding crista deltopectoralis of the humerus, also for the first time visible in one of the new specimens, shows gallinuloidids to be the sister taxon of all other, extinct and extant, galliform birds. Gallinuloidids distinctly differ from modern Galliformes in several other plesiomorphic osteological features, mainly of the pectoral girdle, of which the absence of a spina interna on the sternum is here reported for the first time. It is assumed that major evolutionary transformations in the stem lineage of Galliformes are related to the evolution of a large crop, which appears to have been absent in gallinuloidids. The vegetarian food component of gallinuloidids thus probably mainly consisted of soft plant matter rather than coarse material such as seeds.     

REPTILIAN PHYSIOLOGY AND THE FLIGHT CAPACITY OF ARCHAEOPTERYX

Current scenarios frequently interpret the Late Jurassic bird Archaeopteryx as having had an avian-type physiology and as having been capable of flapping flight, but only from “the trees downward.” It putatively lacked capacity for takeoff and powered flight from the ground upward. Data from extant reptiles indicate that if Archaeopteryx were physiologically reptilian, it would have been capable of ground upward takeoff from a standstill, as well as “trees downward” powered flight. This conclusion is based largely on a previously unrecognized attribute of locomotory (skeletal) muscle in a variety of extant reptiles: During “burst-level” activity, major locomotory muscles of a number of active terrestrial taxa generate at least twice the power (watts kg^?1 muscle tissue) as those of birds and mammals. Reptilian physiological status also helps resolve the apparently uneven development of various flight support structures in Archaeopteryx (e.g., well-developed flight features but relatively unspecialized pectoral girdle, supracoracoideus muscles, etc.). Endothermy and capacity for longer-distance powered flight probably evolved only in Early Cretaceous birds, which were the first birds to have a keeled sternum, strap-like coracoid, and hypocleidium-bearing furcula.     

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.     

An assessment of the diversity of early Miocene Scolopaci (Aves,Charadriiformes) from Saint‐Gérand‐le‐Puy (Allier,France)

Abstract: ‘Totanus’lartetianus, Elorius paludicola and ‘Tringa’gracilis are the three scolopacid birds from the early Miocene of Saint‐Gérand‐le‐Puy described by the French palaeontologist Milne‐Edwards in the 19th century. Since then, no revision of these taxa has been performed. Our re‐examination revealed that not much of the material originally assigned to ‘Totanus’lartetianus can be retained within the species. Presumably plesiomorphic features of the humerus – and potentially the coracoid we attributed to this taxon – indicate that it is misplaced in the extant scolopacid genus and may not even belong to the Scolopacidae (sandpipers and allies), and we therefore place it in the new genus Scolopacimilis. Comparisons of the material assigned to Elorius paludicola and ‘Tringa’gracilis show that they are morphologically similar, both exhibiting distinct scolopacid anatomical features. The latter, however, cannot be referred to the extant taxon Tringa and is classified into the new genus Parvelorius. We further introduce three new species, ?Elorius limosoides sp. nov., and ?Parvelorius calidris sp. nov., which we have tentatively assigned to the extinct scolopacid genera Elorius and Parvelorius, respectively, and Becassius charadriioides gen. et. sp. nov., which, together with Scolopacimilis, display a morphology uncharacteristic for extant Scolopaci. For the first time we have assigned skulls to some of the postcranial elements described in this study. The presence of at least six species of Scolopaci from the early Miocene considerably increases the number of members of the group known from this time.     

Cenozoic mystery birds – on the phylogenetic affinities of bony‐toothed birds (Pelagornithidae)

Mayr, G. (2011) Cenozoic mystery birds – on the phylogenetic affinities of bony‐toothed birds (Pelagornithidae). —Zoologica Scripta, 40, 448–467. The extinct Cenozoic bony‐toothed birds (Pelagornithidae) are characterized by the occurrence of unique spiky projections of the osseous jaws and are among the most distinctive neornithine taxa. Earlier authors considered these marine birds to be most closely related to ‘Pelecaniformes’ or Procellariiformes, but recent phylogenetic analyses resulted in a sister group relationship to Anseriformes. This latter hypothesis was, however, coupled with a non‐monophyly of galloanserine or even neognathous birds, which is not supported by all other current analyses. The character evidence for anseriform affinities of pelagornithids is thus reassessed, and it is detailed that the alleged apomorphies cannot be upheld. Pelagornithids lack some key apomorphies of galloanserine birds, and analysis of 107 anatomical characters did not support anseriform affinities, but resulted in a sister group relationship between Pelagornithidae and Galloanseres. By retaining a monophyletic Galloanseres, this result is in better accordance with widely acknowledged hypotheses on the higher‐level phylogeny of birds. The (Pelagornithidae + Galloanseres) clade received, however, only weak bootstrap support, and some characters, such as the presence of an open frontoparietal suture, may even support a position of Pelagornithidae outside crown‐group Neognathae.     

Anatomy and early development of the pectoral girdle,fin, and fin spine of sturgeons (Actinopterygii: Acipenseridae)

Acipenseriformes hold an important place in the evolutionary history of bony fishes. Given their phylogenetic position as extant basal Actinopterygii, it is generally held that a thorough understanding of their morphology will greatly contribute to the knowledge of the evolutionary history and the origin of diversity for the major osteichthyan clades. To this end, we examined comparative developmental series from the pectoral girdle in Acipenser fulvescens, A. medirostris, A. transmontanus, and Scaphirhynchus albus to document, describe, and compare ontogenetic and allometric differences in the pectoral girdle. We find, not surprisingly, broad congruence between taxa in the basic pattern of development of the dermal and chondral elements of the pectoral girdle. However, we also find clear differences in the details of structure and development among the species examined in the dermal elements, including the clavicle, cleithrum, supracleithrum, posttemporal, and pectoral‐fin spine. We also find differences in the internal fin elements such as the distal radials as well as in the number of fin rays and their association with the propterygium. Further, there are clear ontogenetic differences during development of the dermal and chondral elements in these species and allometric variation in the pectoral‐fin spine. The characters highlighted provide a suite of elements for further examination in studies of the phylogeny of sturgeons. Determining the distribution of these characters in other sturgeons may aid in further resolution of phylogenetic relationships, and these data highlight the role that ontogenetic and comparative developmental studies provide in systematics. J. Morphol. 276:241–260, 2015. © 2014 Wiley Periodicals, Inc.     

The bizarre wing of the Jamaican flightless ibis Xenicibis xympithecus: a unique vertebrate adaptation

Birds have frequently evolved to exploit insular environments by becoming adapted to a terrestrial lifestyle and losing the ability to fly, usually via reducing the wings and pectoral girdle. The enigmatic flightless ibis Xenicibis xympithecus (Threskiornithidae) from the Quaternary of Jamaica provides a rare example of flight loss in ibises. We report on previously undescribed fossils of Xenicibis, and show that the wing differed radically from that of all other birds, flightless or volant. The metacarpus is elongate, grotesquely inflated and has extremely thick walls; phalanges are short and block-like; the radius is distally expanded; and the humerus is elongate. The furcula, coracoid and sternum are all well developed. We propose that the elongate forelimb and massive hand functioned in combat as a jointed club or flail. This hypothesis is supported by the morphology of the carpometacarpus, by features permitting rapid extension of the wing and by the presence of fractures in wing bones. Although other birds use the wings as weapons, none resemble Xenicibis, which represents a unique and extraordinary morphological solution to this functional problem. Xenicibis strikingly illustrates how similar selective pressures, acting on a similar starting point, can result in novel outcomes.     

Pectoral girdle and forelimb variation in extant Crocodylia: the coracoid–humerus pair as an evolutionary module

To date, all statements about evolutionary morphological transformation in Crocodylia have essentially been based on qualitative observations. In the present study, we assessed the morphological variation and covariation (integration) between the scapula, coracoid, humerus, radius, and ulna of 15 species of Crocodylidae, Alligatoridae, and Gavialis + Tomistoma using three‐dimensional geometric morphometrics. The results obtained reveal that the variation of elements within species (intraspecific) is large. However, despite this variability, variation across species (interspecific) is mainly concentrated in two dimensions where the disparity is constrained: ‘robusticity’ and ‘twist’ (forelimbs) and ‘robusticity’ and ‘flexion’ (pectoral girdle). Robusticity (first dimension of variation) embodies a set of correlated geometrical features such as the broadening of the girdle heads and blades, or the enlargement of proximal and distal bone ends. The twist is related to the proximal and/or distal epiphyses in the forelimb elements, and flexion of the scapula and coracoid blades comprises the second dimension of variation. In all crocodylians, forelimb integration is characterized by the strong correlations of a humerus–ulna–radius triad and by a radius–ulna pair, thus forming a tight forelimb module. Unexpectedly, we found that the humerus and coracoid form the most integrated pair, whereas the scapula is a more variable and relatively independent element. The integration pattern of the humerus–coracoid pair distinguishes a relatively robust configuration in alligatorids from that of the remainder groups. The patterns of variation and integration shared by all the analyzed species have been interpreted as an inherited factor, suggesting that developmental and functional requirements would have interacted in the acquisition of a semi‐aquatic and versatile locomotion at the Crocodylia node at least 65 Mya. Our findings highlight the need to incorporate the humerus–coracoid pair in biodynamic and biomechanical studies. © 2012 The Linnean Society of London     

Growth of the pectoral girdle of the Leopard frog, Rana pipiens (Anura: Ranidae)

This article describes the growth of the anuran pectoral girdle of Rana pipiens and compares skeletal development of the shoulder to that of long bones. The pectoral girdle chondrifies as two halves, each adjacent to a developing humerus. In each, the scapula and coracoid form as single foci of condensed chondrocytes that fuse, creating a cartilaginous glenoid bridge articulating with the humerus. Based on histological sections, both the dermal clavicle and cleithrum begin to ossify at approximately the same time as the periosteum forms around the endochondral bones. The dermal and endochondral bones of the girdle form immobile joints with neighboring girdle elements; however, the cellular organization and growth pattern of the scapula and coracoid closely resemble those of a long bone. Similar to a long bone epiphysis, distal margins of both endochondral elements have zones of hyaline, stratified, and hypertrophic cartilages. As a result, fused elements of the girdle can grow without altering the glenoid articulation with the humerus. Comparisons of anuran long bone and pectoral girdle growth suggest that different bones can have similar histology and development regardless of adult morphology.     

The pectoral anatomy of selected Ostariophysi. II. The Cypriniformes and siluriformes

The pectoral myology and osteology of the cyprinoids Notemigonus crysoleucas, the golden shiner, and Catostomus commersonnii, the common white sucker, resemble those of generalized, lower teleosts in structure and function, except in features related to the manipulation of the massive fifth ceratobranchial of cyprinoids by muscles attaching on the girdle. Catostomus is more specialized in having unique intercostal muscles to the girdle, complex subclavian arteries and lack of a superficial trapezius muscle. The bony pectoral anatomy of the siluriform, Ictalurus nebulosus, the brown bullhead, is highly specialized in relation to the presence and locking of the massive pectoral spine which is formed of fused dorsal and ventral propterygial rays; there is consolidation of the girdle through fusion of bones, presence of unique stabilizing bony structures, firm symphyseal union of bilateral girdles and the presence of friction-surfaces of girdle and spine for locking. The movements of the spine are specialized in the greater guidance offered by the girdle. Myological specializations are related mainly to ventral appendicular muscles which lock the spine. The nervous and arterial systems are generalized.     

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.     

A chiton without a foot

Abstract: The palaeoloricate ‘polyplacophorans’ are an extinct paraphyletic group of basal chiton‐like organisms known primarily from their fossilized valves. Their phylogenetic placement remains contentious, but they are likely to include both stem‐group Polyplacophora and stem‐group Aplacophora. Candidates for the latter position include ‘Helminthochiton’thraivensis from the Ordovician of Scotland, which we redescribe here through a combined optical and micro‐CT (XMT) restudy of the type material. The 11 specimens in the type series are all articulated, presenting partial or complete valve series as well as mouldic preservation of the girdle armature; they demonstrate a vermiform body plan. The valves are typically palaeoloricate in aspect, but differ in detail from all existing palaeoloricate genera; we hence erect Phthipodochiton gen. nov. to contain the species. The most notable feature of the fossils is the spicular girdle; this is impersistently preserved, but demonstrably wraps entirely around the ventral surface of the animal, implying that a ‘true’ (i.e. polyplacophoran like) foot was absent, although we do not exclude the possibility of a narrow solenogastre‐like median pedal groove having been present. Phthipodochiton thraivensis presents an apparent mosaic of aplacophoran and polyplacophoran features and as such will inform our understanding of the relationship between these groups of extant molluscs. An inference may also be drawn that at least some other palaeoloricates possessed an ‘armoured aplacophoran’ body plan, in contrast to the ‘limpet‐like’ body plan of extant Polyplacophora.     

How can ontogeny help us to understand the morphology of anuran pectoral girdle?

The ontogenetic development of the pectoral girdle in seven anuran species (Xenopus laevis, Discoglossus pictus, Bombina bombina, Bombina variegata, Pelobates fuscus, Bufo bufo and Rana dalmatina) was studied using cleared and stained specimens. The epicoracoid cartilage was found to develop in two different ways resulting in an arciferal or firmisternal type of the pectoral girdle. In the arciferal one, the epicoracoid originates from a medial prolongation of the procoracoid cartilage and broadly overlaps its counter part during further development. In the firmisternal pectoral girdle, the epicoracoids are formed by the widened cartilaginous medial edges of the coracoids that fuse together along the midline. Polarization of ontogenetic characters shows, that omosternum evolves inside Anura, and the type of sternum occurring in basal Anura seems to be an apomorphy of all Batrachia. The sternal elements have a single or paired rudiment, their development is connected with M. rectus abdominis or a zonal area and they remain cartilaginous or ossify during postmetamorphic development. An occurrence of omosternum in Barbourula busuangensis was described for the first time.     

The morphology and systematics of Mammalodon colliveri (Cetacea: Mysticeti), a toothed mysticete from the Oligocene of Australia

Mammalodon colliveri is an unusual toothed archaic mysticete (Cetacea) from the Upper Oligocene Jan Juc Formation of south‐east Australia. The morphology of the holotype skull and postcrania are described in detail. Superimposed on the generally plesiomorphic archaeocete‐like morphology of Mammalodon are subtle mysticete synapomorphies. Derived features of Mammalodon include a short and bluntly rounded rostrum, reduced premaxillae, and anterodorsally directed orbits. Within Mysticeti, this suite of features is unique. The aberrant rostral morphology of Mammalodon suggests specialization for suction feeding. Janjucetus hunderi is placed in an expanded family Mammalodontidae. Phylogenetic analysis corroborates the monophyly of Basilosauridae, Neoceti, Odontoceti, and Mysticeti, and yields a novel hypothesis of toothed mysticete relationships: a basal clade of undescribed toothed mysticetes from South Carolina, a Llanocetidae + Mammalodontidae clade, and monophyletic Aetiocetidae are posited as successive sister taxa to edentulous baleen whales (Chaeomysticeti). Toothed archaic mysticetes clearly employed diverse prey capture strategies, with exaptations for filter feeding evolving sequentially in stem group Mysticeti. A stratigraphically calibrated phylogeny implies that the initial diversification of Mysticeti occurred during the Late Eocene. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 158 , 367–476.     

The suction mechanism of the pipid frog,Pipa pipa (Linnaeus, 1758)

Most suction‐feeding, aquatic vertebrates create suction by rapidly enlarging the oral cavity and pharynx. Forceful enlargement of the pharynx is powered by longitudinal muscles that retract skeletal elements of the hyoid, more caudal branchial arches, and, in many fish, the pectoral girdle. This arrangement was thought to characterize all suction‐feeding vertebrates. However, it does not exist in the permanently aquatic, tongueless Pipa pipa , an Amazonian frog that can catch fish. Correlating high‐speed (250 and 500 fps) video records with anatomical analysis and functional tests shows that fundamental features of tetrapod body design are altered to allow P. pipa to suction‐feed. In P. pipa , the hyoid apparatus is not connected to the skull and is enclosed by the pectoral girdle. The major retractor of the hyoid apparatus arises not from the pectoral girdle but from the femur, which lies largely within the soft tissue boundaries of the trunk. Retraction of the hyoid is coupled with expansion of the anterior trunk, which occurs when the hypertrophied ventral pectoral elements are depressed and the urostyle and sacral vertebra are protracted and slide forward on the pelvic girdle, thereby elongating the entire trunk. We suggest that a single, robust pair of muscles adduct the cleithra to depress the ventral pectoral elements with force, while modified tail muscles slide the axial skeleton cranially on the pelvic girdle. Combined hyoid retraction, axial protraction, and pectoral depression expand the buccopharyngeal cavity to a volume potentially equal to that of the entire resting body of the frog. Pipa may be the only tetrapod vertebrate clade that enlarges its entire trunk during suction‐feeding.     

GYRACANTHIDES HAWKINSI SP. NOV. (ACANTHODII, GYRACANTHIDAE) FROM THE LOWER CARBONIFEROUS OF QUEENSLAND, AUSTRALIA, WITH A REVIEW OF GYRACANTHID TAXA

Abstract:  A new species of Gyracanthides from the mid-Viséan Ducabrook Formation of Middle Paddock site, near Springsure in the Drummond Basin, central Queensland, is based on isolated three-dimensionally preserved elements. The specimens comprise paired and unpaired spines and pectoral girdle elements, procoracoids and scapulocoracoids, and include growth series. The morphology, especially of the shoulder girdle bones and the form and tubercular ornamentation of the paired fin spines, is used to distinguish the new taxon. These characters also help differentiate the numerous described gyracanthid species. Aspects of palaeobiology including possible sexual dimorphism are explored. A hypothetical reconstruction of the fish is based on our interpretation of the articulation of isolated elements combined with examination of wear patterns on fin spines. Gyracanthides hawkinsi sp. nov. is compared with other Australian taxa as well as with gyracanthids from North America, Europe, Russia, Iran, Africa and Antarctica, some of which are tentatively reassigned here to the Gondwanan genus Gyracanthides .