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Oryctos

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volume 7, 2008

Andrzej Elzanowski, The avian femur: morphology and terminology of the lateral condyle. Oryctos 7, 1-5.

In the dinosaurs, the lateral condyle of the femur is subdivided into two parts, here termed semicondyles: cranial semicondyle which articulates primarily with the fibula, and caudal semicondyle which is known in the theropods as the ectocondylar tuber and articulates with the tibia. Modern birds also have two semicondyles, fibular and tibiofibular (“tibiofibular crest”), which roughly correspond in position to the non-avian theropod semicondyles. However, the basal birds have a single rounded lateral condyle which must have undergone differentiation into two modern avian semicondyles independently of those in the non-avian theropods. Since extrapolating anatomical terms for details of the femoral lateral condyle between modern birds and theropods seems unwarranted, I propose to use two separate, consistent sets of terms, both based on the subdivision of the lateral condyle into two semicondyles, as implemented for modern birds in Table 1.



Richard Hinchliffe, Bird wing digits & their homologies: reassessment of developmental evidence for a 2,3,4 identity. Oryctos 7, 7-12.
The theory of descent of birds from theropod dinosaurs demands that their fore-limb digit identities (1,2,3) are the same as those of birds and thus the conventional embryological identification of these as 2,3,4 remains a major problem for acceptance of this theory. Are the 2,3,4 identities of bird wing digits correct? The paper analyses the developing bird wing as a specialisation of the general developmental ‘bauplan’ for the pentadactyl skeleton. Evidence from the chick embryonic skeletogenic pattern supports interpretation of the main digits as 2,3,4 on the basis of timing, position & connections, using comparative methods eg comparison with other amniote patterns of limb skeletogenesis. Fresh support for 2,3,4 identity comes from evidence of i) a temporary embryonic digit 1 in the ostrich and of ii) a condensation-specific Sox9 molecular domain in a digit 1 position in the chick wingbud. In contrast, the recent ‘frame shift’ hypothesis of Wagner proposes molecular identity transformation by which theropod identities for 1,2,3 have become shifted to avian digit condensations 2,3,4. Support for ‘frame shift’ is claimed from evidence (Vargas & Fallon 2004) that the expression domain for Hox d 13 alone characterises digit 1, but a domain for Hox d 12 and 13 characterises digits 2-5. Here it is argued that this evidence for ‘frame shift’ is speculative and insufficiently convincing to support reinterpretation of the wing digits as 1,2,3.Evidence that wing digits have been correctly identified as 2,3,4 continues to provoke doubts about the ‘dinosaur-bird’ theory. The presence of well-defined feathers in the bird-like hands of dromaeosaurs (from Liaoning, China) may be due to their possibly being secondarily flightless birds, thus suggesting that birds may derive from a lineage separate from that of theropods.

Kenneth E. Campbell, The manus of Archaeopterygians: implications for avian ancestry. Oryctos 7, 13-26.

Recent, apparent confirmation that the digits of the avian manus are II, III, and IV, and not I, II, and III as in theropods, brings into question the purported homologies thought by many to have existed between the wrist and manus of early birds and those of maniraptoran theropods. A close examination of the wrist and manus of five archaeopterygians confirms the presence of numerous previously recognized, as well as unrecognized, derived avian characters. These include, but are not limited to, Metacarpal II very short, slightly wrapping around and probably fused to Metacarpal III for much of its length; Metacarpal II with small Processus extensorius for attachment of M. extensor carpi radialis; Metacarpal IV with proximal end lying well distal to proximal ends of metacarpals II and III, wrapping under and fused to ventral surface of Metacarpal III; joint between Metacarpal II and Phalanx 1 of Digit II ginglymoid near plane of wing, allowing Digit II to function as a primitive alula; joints between metacarpals III and IV and their respective phalanges relatively immobile; and “palmar” surfaces of distal phalanges facing anteroventrad or anteriad when wing is extended. The above features are not found in theropods, but their more highly derived counterparts are found in all modern birds with wings, whether volant or not. The avian, rather than theropodian, structure of the wrist and manus of archaeopterygians indicates significant functional differences between the forelimbs of archaeopterygians and theropods, which is not surprising if different digits are involved. Some prominent “feathered dinosaurs” are recognized as having a more advanced avian manus than that found in archaeopterygians, indicating their avian, rather than dinosaurian, ancestry.



David A. Burnham, A review of the Early Cretaceous Jehol Group on northeastern China and a revision of concerning the origin of flight paradigm. Oryctos 7, 27-43.

The unusual preservation and evolutionary significance of the avian and non-avian dinosaur fossils from western Liaoning Province in China exemplifies rare conservation deposits. Described as the Jehol Biota, the fauna includes such remarkable discoveries such as feathers and wings associated with dinosaurs as well as many new species of fossil birds preserved in abundance. Volcanic activity during the Mesozoic was crucial to the preservation of the Early Cretaceous Jehol Biota by acting as a mechanism that killed organisms en masse from volatile emissions and voluminous ash falls. One of the crucial specimens documented in the Liaoning deposits is Microraptor gui, a four-winged glider. Described as a feathered, non-avian dinosaur, the evolution of such an animal in a group closely related to birds necessitates revision of the origin of flight paradigm.



Larry D. Martin, Origins of avian flight – a new perspective. Oryctos 7, 45-54.
The discovery of a primitive bird-like dromaeosaur (Microraptor) with four functional wings vindicates Beebe’s suggestion that birds went through a tetrapteryx stage in the origin of flight. Flight originated from an arboreal gliding ancestor and Longisquama may be more central to understanding how this came about than previously supposed.

Attila Ösi, Enantiornithine bird remains from the Late Cretaceous of Hungary. Oryctos 7, 55-60.
The recently documented vertebrate fauna from the Santonian Csehbánya Formation of Iharkút, (Bakony Mts, Hungary) provides the first occurrence of the extinct avian group Enantiornithes Walker, 1981 in the Upper Cretaceous of Central Europe. A nearly complete left femur and a complete right tarsometatarsus are referred here to Enantiornithes indet., and a distal fragment of a subadult left femur and a metatarsus III described here as Aves indet., represent the earliest known avian fossil record from Hungary. The isolated two femora and the fragmentary metatarsus III indicate the presence of very small terrestrial birds in the Santonian of Europe. The well-preserved femur documents that some members of the Hungarian enantiornithines retained the small size of their Early Cretaceous ancestors. The large enantiornithine tarsometatarsus supports the existence of a larger enantiornithine bird in the area, which may have been adapted to aquatic or semi-aquatic habitats.

Larry D. Martin & Virginia L. Naples, Mandibular kinesis in Hesperornis. Oryctos 7, 61-65.

Some aspects of mandibular morphology are known for three hesperornithiform genera: Hesperornis, Parahesperornis and Baptornis. All share a distinctive intramandibular joint between the angular and the splenial. A special process of the surangular extending between the splenial and the dentary bridges the joint. The symphysis appears to have been elongate and unfused, joining anteriorly with a short intersymphyseal bone. It appears that the mandibles spread posteriorly as the jaws opened, allowing the swallowing of larger prey. The closed mandible is very slender anteriorly, resembling some cetaceans, and seems highly adapted for the capture of fish. The discovery of fish remains in a preserved stomach cast of Baptornis gives direct support for this interpretation.



Gerald Mayr, The higher-level phylogeny of birds - when morphology, molecules,and fossils coincide. Oryctos 7, 67-74.
Although the higher-level relationships of modern birds are still poorly resolved, there some clades result from cladistic analyses of both morphological and molecular data, and are in further agreement with the mosaic character distribution in Paleogene fossil taxa. Examples are sister group relationships between Galliformes (landfowl) and Anseriformes (waterfowl), Phoenicopteriformes (flamingos) and Podicipediformes (grebes), Aegothelidae (owlet-nightjars) and Apodiformes, and jacamars/puffbirds (Galbulae) and Pici (woodpeckers and allies). Recent molecular studies further support a position of Turnicidae (buttonquails) within Charadriiformes (shorebirds), which is in concordance with the mosaic distribution of turnicid and charadriiform characters in the early Oligocene taxon Turnipax. Most of the above clades have initially been suggested from studies of morphological data, and despite recent progress in molecular analyses phylogenies based on morphological characters are still needed to set fossil taxa into a phylogenetic context.

Eric Buffetaut, First evidence of the giant bird Gastornis from southern Europe: a tibiotarsus from the Lower Eocene of Saint-Papoul (Aude, southern France). Oryctos 7, 75-82.

A well preserved tibiotarsus from the Early Eocene locality of Saint-Papoul (Aude, southwestern France) is described as belonging to the giant ground bird Gastornis parisiensis Hébert, 1855, on the basis of close resemblances with the lectotype of that species. Small and variable differences in the tibiotarsi of European and North American gastornithids are considered as insufficient to justify a separation between Gastornis and Diatryma, the latter being considered as a junior synonym of the former. The discovery of Gastornis in southern France lends weight to the hypothesis according to which the large bird eggs found in the continental Lower Eocene of southern France may have been laid by gastornithids.



Kazuhiko Sakurai, Masaichi Kimura & Takayuki Katoh, A new penguin-like bird (Pelecaniformes:Plotopteridae) from the Late Oligocene Tokoro Formation, northeastern Hokkaido, Japan. Oryctos 7, 83-94.

The skeleton of a large bird from Late Oligocene marine strata of the Tokoro Formation, exposed near Abashiri City in northeastern Hokkaido, Japan, represents a new genus and species of the extinct pelecaniform family Plotopteridae. This new specimen, Hokkaidornis abashiriensis new gen. et sp., was similar in size to Copepteryx hexeris Olson and Hasegawa, but differs from that species chiefly in characters of the anterior parts of the skeleton. The leg bones of H. abashiriensis and C. hexeris are nearly identical, and if found isolated, probably would be deemed congeneric/conspecific. The new animal from Hokkaido demonstrates that plotopterid generic diversity during Late Oligocene time was greater than previously suspected.



Ursula B. Göhlich & Marco Pavia, A new species of Palaeortyx (Aves: Galliformes: Phasianidae) from the Neogene of Gargano, Italy. Oryctos 7, 95-108.

A new species, Palaeortyx volans n. sp., of Phasianidae (Aves: Galliformes) is described from the Neogene vertebrate assemblage of the Gargano (Italy). The Gargano fossil vertebrate association has been well known since 1971, and it is considered to be an island fauna because of the high degree of endemism shown by different taxa. The original description of the fossil avifauna of Gargano referred all the phasianid specimens to Palaeortyx grivensis, a fossil species described form the Miocene (MN7+8) of La Grive-Saint-Alban (France). The present study reveals the differences between Palaeortyx volans n. sp. and the other species of Palaeortyx described until now. The new species described herein represents the smallest species of the genus known, except for P. joleaudi from La Grive-Saint-Alban. The systematic position of P. grivensis from La Grive-Saint-Alban is supported. Morphometric analysis of the fossil remains of P. volans n. sp. suggests good flying capabilities and an ecology similar to that of the recent Coturnix coturnix. These features allowed P. volans n. sp. to colonize isolated islands such as Gargano, which are normally not inhabited by short-distance flyers like the phasianids.



Carolina Acosta Hospitaleche & Claudia Tambussi,South American fossil penguins: a systematic update. Oryctos 7, 109-127.

During the last few years, we have worked on the systematics and paleobiology of the South American and Antarctic fossil penguins. As a result, we have obtained new data about their past biodiversity. Concerning South American fossil penguins, particularly the Tertiary ones, we can point out that, based on phylogenetic and morphometric analyses practiced on skulls and appendicular skeleton, we recognised three non taxonomic groups, partially in agreement partially with the systematic scheme proposed by Simpson: Paraptenodytinae, Palaeospheniscinae and Spheniscinae. These species are recorded exclusively from South America and morphologically have more resemblance with the living species than the fossil penguins from others regions of the Southern Hemisphere. This suggests that the evolutionary and biogeographical history of the penguin fauna of Argentina, Chile and Peru followed different routes from those of Antarctica, New Zealand and Australia.



Stig A. Walsh, Norman MacLeod & Mark O’Neill, Analysis of spheniscid humerus and tarsometatarsus morphological variability using DAISY automated image recognition. Oryctos 7, 129-136.

Despite a long history of research, relationships within fossil and extant Sphenisciformes remain unclear. This is largely because most fossil species were described on the basis of either the tarsometatarsus or humerus. Neither of these elements is particularly phylogenetically informative, and the extent of intraspecific morphological variation also remains unknown. Herein we investigate a new approach – the use of artificial neural-net (ANN) technology – to determine whether either of these elements can be reliably used to identify extant species. The DAISY ANN system was able to recognise most species from either tarsometatarsal or humerus morphology, but its success rate improved when the species training sets were combined into generic groups, indicating the need for larger image libraries. Our preliminary results suggest that these elements can allow reliable identifications for most taxa, but that the tarsometatarsus is on the whole a better element for this purpose. These results also demonstrate the potential for artificial neural-net technology to address problems in avian taxonomy.



Claudia Patricia Tambussi & Carolina Acosta Hospitaleche, Skull shape analysis and diet of South American fossil penguins (Sphenisciformes). Oryctos 7, 137-145.

Form and function of the skull of Recent and fossil genera of available Spheniscidae are analysed in order to infer possible dietary behaviors for extinct penguins.Skull and mandible shapes were compared using the Resistant-Fit Theta-Rho-Analysis (RFTRA) Procrustean method. Due to the availability and quality of the material, this study was based on six living species belonging to five genera (Spheniscus, Eudyptula, Eudyptes, Pygoscelis, and Aptenodytes) and two Miocene species: Paraptenodytes antarticus (Moreno and Mercerat, 1891) and Madrynornis mirandus Acosta Hospitaleche, Tambussi, Donato & Cozzuol. Seventeen landmark from the skull and five from the mandible were chosen, including homologous and geometrical points. Morphological similarities among RFTRA distances are depicted using the resulting dendrograms for UPGMA (unweighted pair-group method using arithmetic average) cluster analysis. This shape analysis allows the assessment of similarities and differences in the skulls and jaws of penguins within a more comprehensive ecomorphological and phylogenetic framework. Even though penguin diet is not well known, enough data supports the conclusion that Spheniscus + Eudyptes penguins specialize on fish and all other taxa are plankton-feeders or fish and crustacean-feeders. We compared representative species of both ecomorphological groups with the available fossil material to evaluate their feeding strategies. Penguins are the most abundant birds, indeed the most abundant aquatic tetrapods, in Cenozoic marine sediments of South America. The results arising from this study will be of singular importance in the reconstruction of those marine ecosystems.



Antoine Louchart, Yohannes Haile-Selassie, Patrick Vignaud, Andossa Likius & Michel Brunet, Fossil birds from the Late Miocene of Chad and Ethiopia and zoogeographical implications. Oryctos 7, 147-167.

Since a review of the Tertiary birds of Africa 30 years ago by Rich (1974), a great wealth of new data and new interpretations have augmented this fossil record, hitherto still relatively poor when compared with that of northern continents. A growing amount of new fossils is coming from hominid localities, now extending as early as the Late Miocene. Recently found bird fossils from the Late Miocene of Chad and Ethiopia are described here. They come respectively from several localities of Toros Menalla (Djurab Desert, Chad), dated ca 7 Ma, several localities of the Western Margin of the Middle Awash (Afar rift, Ethiopia), dated 5.6-5.8 Ma, and a locality of the Central Awash Complex of the Middle Awash, dated ca 5.2 Ma. 19 different taxa in 11 families are identified, essentially aquatic birds except for part of the fossils from one Ethiopian locality, ALA-VP-2. The bird assemblages from the three groups of localities by age and location provide some paleoenvironmental indications. Several taxa are new, or are new to Africa, and several are the earliest fossil records of a modern genus or lineage. Three extinct taxa are represented: the large darter Anhinga cf. A. pannonica, a new species of a giant saddlebill stork, genus Ephippiorhynchus, and a swan Afrocygnus chauvireae. Families with an extremely scarce record worldwide are represented here (Pandionidae, Heliornithidae). Comparisons are made which reveal some similarities between the Chadian and Ethiopian sites, as well as with Late Miocene localities in Kenya, Tunisia and Libya. At a larger scale, from the Mio-Pliocene of Africa, Leptoptilos falconeri and Anhinga cf. A. pannonica were also of Eurasian distribution, while Pavo sp. and Heliopais cf. H. personata are now typical of the Oriental Region. These represent important zoogeographical links between Africa and Eurasia at that period, compared with the relations observed today between the non-passerine birds of the Paleotropical (=Ethiopian) and Oriental regions.



Cécile Mourer-Chauviré & Denis Geraads, The Struthionidae and Pelagornithidae (Aves: Struthioniformes, Odontopterygiformes) from the late Pliocene of Ahl Al Oughlam, Morocco. Oryctos 7, 169-194.

The Pliocene locality of Ahl al Oughlam is situated at the southeastern limit of the city of Casablanca, in Morocco, on an ancient seashore of the Atlantic Ocean. It has yielded a very rich vertebrate fauna (macro- and micromammals, birds, reptiles, amphibians and fishes) including both terrestrial and marine forms. On the basis of biostratigraphy, the fauna has been dated at about 2.5 Ma, which corresponds to the latest Pliocene. The avifauna is very diverse and includes birds belonging to twelve different orders. In this paper we describe only the Struthionidae and the Pelagornithidae. Ostriches are represented by a large-sized form, referred to the extinct species Struthio asiaticus Milne-Edwards, and its eggshells, of struthioid type, are comparable to those of the recent species Struthio camelus, but thicker. The Pelagornithidae, giant marine birds with bony pseudoteeth, are represented by an extinct species of the genus Pelagornis. As far as we know, this species was probably the latest representative of the order Odontopterygiformes.



Walter E. Boles, Systematics of the fossil Australian giant megapodes Progura (Aves: Megapodiidae). Oryctos 7, 195-215.

Australia’s largest megapode was Progura gallinacea De Vis, 1888, from Plio-Pleistocene deposits in southeastern Queensland. A second species, P. naracoortensis van Tets, 1974, from southeastern South Australia, was initially distinguished by different leg proportions and size; it was later suggested that there might be only a single, sexually dimorphic species. An examination of extensive collections of unstudied material from the Naracoorte Caves indicates that there is only a single species, with size differences related to both moderate sexual dimorphism and individual variation. Comparisons of these specimens with the three living Australian megapode taxa show that the Progura is not separable from the modern genus Leipoa. The possibility that Progura gallinacea is the megafaunal representative of the living Leipoa ocellata Gould, 1838 (Malleefowl) is discussed.



Nikita V. Zelenkov, Evgeny N. Kurochkin, Alexander A. Karhu & Peter Ballmann, Birds of the Late Pleistocene and Holocene from the Palaeolithic Djuktai Cave site of Yakutia, Eastern Siberia. Oryctos 7, 217-226.

Avian remains from the Late Palaeolithic Djuktai Cave (15000 – 740 years B.P.) are represented by more than 700 bird bones. We identified 65 species from 10 modern orders. The most abundant are remains of the Willow Grouse (Lagopus lagopus), and Ptarmigan (L. mutus). The remains of Ducks (Anatidae), Waders (Scolopacidae), and Thrushes (Turdus spp.) are less numerous. Except for a few cases, most avian species identified represent the fauna of open-land habitats, for example Otis tarda. In modern faunas these birds are inhabitants of steppe landscapes which are absent in the Aldan basin now. Thus the presence of these species indicates that there was a steppe near the Cave at the end of the Pleistocene. In addition, some of the Waterfowl bones recovered have cut-marks made by Palaeolithic man.



Joanne H. Cooper & Alan J. D. Tennyson, Wrecks and residents: the fossil gadfly petrels (Pterodroma spp.) of the Chatham Islands, New Zealand. Oryctos 7, 227-248.

During some 40 years’ research, ten species of gadfly petrel, Pterodroma spp., have been reported from Holocene fossil assemblages from the Chatham Islands, New Zealand. Statistical analysis of the assemblages’ species composition and a critical appraisal of all claimed identifications have resulted in a re-assessment of fossil Pterodroma diversity prior to human arrival. Three species are confirmed as original breeding residents; the endemic Pterodroma magentae and P. axillaris, and the undescribed extinct Pterodroma sp.1. Additionally, P. nigripennis is confirmed as a vagrant. Possible records of further vagrant species, P. inexpectata, P. macroptera or P. lessonii and P. neglecta cannot yet be confirmed.



Tommy Tyrberg, The Late Pleistocene Continental Avian extinction – an evaluation of the fossil evidence. Oryctos 7, 249-269.

This paper is a review of the characteristics, including timing, geographic distribution and severity, of the Late Pleistocene extinction among continental birds, and an evaluation of the suggested reasons for it. The analysis shows that the Late Pleistocene avian extinctions correlate well with the mammalian megafaunal extinction with respect to severity and timing, and that many of the extinct birds were probably directly or indirectly dependent on the mammalian megafauna. A large proportion of the extinct bird species had life history traits that would have made them vulnerable to human predation.



Vasily A. Ilyinsky, Locomotor Adaptations in the Hindlimbs of Owls: the Burrowing Owl (Athene cunicularia), compared to the Little Owl (Athene noctua). Oryctos 7, 271-276.

The present study provides an attempt to evaluate some morphological adaptations for terrestrial locomotion in the Burrowing Owl (Athene cunicularia). The Little Owl (Athene noctua), which is commonly considered a relatively generalised member of the genus, has also been examined for comparisons. Morphological features of the hind limb of the Burrowing Owl revealed by this analysis are interpreted as a consequence of adaptations for high-speed terrestrial locomotion and probably for digging. These adaptations are expressed, for example, in the elongation and thinning of the distal limb elements, promotion of the rotational expanse of the shank and foot, and shifting of muscle bellies in a proximal direction. However, level of the morphological modifications is greatly limited by a necessity to retain basic functional morphological specificity of the hind limb in these taxa. These adaptations, at least in owls, appear to be effective “tools” for prey capture. Some results of the current study can be partly extrapolated to other avian groups because of the fact, that some adaptations of hind limb morphology, associated with a terrestrial life-style, are universal.



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