The first skull of Anthropornis grandis (Aves,Sphenisciformes) associated with postcranial elements

Associated penguin remains found in Bartonian levels of the Submeseta Formation (Seymour Island, Antarctica), including cranium and mandible, both partial tarsometatarsi, and some other fragmentary bones, are analyzed here. This specimen preserves the first cranium reliably assigned to the giant form Anthropornis grandis, and constitutes the first opportunity to taxonomically assign a cranial material to any of the Antarctic penguin species. A discussion of the diet preferences and feeding mechanisms of A. grandis is supported here by three-dimensional paleoneurological and cranial-jaw muscular reconstructions. We propose that A. grandis was a penguin with a voluminous musculature strongly attached to the neck and skull, adapted to chase and hunt fish during diving.     

Palaeeudyptes klekowskii, the best-preserved penguin skeleton from the Eocene–Oligocene of Antarctica: Taxonomic and evolutionary remarks

A new fossil penguin skeleton from the La Meseta Formation collected at the locality DPV 13/84 (Seymour Island, Antarctic Peninsula) from the crinoid horizon located 40 m above the base of the 145 m-thick Submeseta Allomember (Late Eocene–Early Oligocene?) is described. The specimen is assigned to the species Palaeeudyptes klekowskii Myrcha, Tatur and del Valle, 1990; it is the most complete penguin skeleton ever recovered from Antarctica. Discoveries like this one are significant for the study of the anatomy and evolution of penguins, in particular regarding the Antarctic species included in the genus Palaeeudyptes Huxley, 1859. P. klekowskii closely resembles its smaller congeneric species P. gunnari ( Wiman, 1905), with only the relative concavity of the margo medialis distinguishing the tarsometatarsi of both taxa. However, the results of a geometric morphometric analysis show some intra- and inter-specific variations, making possible the systematic assignment of the majority of the specimens. Size variation is congruent with the presence of two different species.     

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.     

First fossilized skin of a giant penguin from the Eocene of Antarctica

An articulated wing belonging to Palaeeudyptes gunnari containing mineralized skin was found in Lutetian (middle Eocene) sediments from Seymour Island, Antarctica. It shows the connective tissues, morphology and density of the feather follicles, and a groove pattern left by the feather calami resting on the skin. Analysis of the preserved surfaces indicates a composition of calcium phosphate. This is the most complete and only articulated wing described for this species and represents the first record of a neornithine bird preserving three-dimensional integument. Plumage density of the wing MLP 14-I-10-22, estimated by counting the follicles under a stereoscopic microscope, is lower than that of the modern Emperor penguin.     

辽宁西部下白垩统反鸟类一新科——Alethoalaorvithidae fam.nov.

记述发现于辽宁省朝阳县原家洼村中生代地层中的反鸟类一新科:真翼鸟科(Alethoalaornithidae fam.nov.)。由于该科鸟类具有肩胛骨直,第1、2指仍有爪,腓骨和胫跗骨不愈合,趾骨不太钩曲等特征,将真翼鸟科归入华夏鸟目(Cathayornithiformes)。真翼鸟科的特有特征包括:吻尖,齿少,牙齿退化的趋势明显,腕掌骨基本形成,乌喙骨远端宽,肱骨近端具明显的气窝,尾综骨长,跗蹠骨末端三趾骨等长且滑车末端基本处于同一水平面,趾骨爪长,不甚钩曲等。通过对新材料和其它早期鸟类的对比认为,反鸟类的灭绝不仅有自身构造的原因,还有它们所生活的生态环境多变的原因。     

Procellariiform remains and a new species from the latest Eocene of Antarctica

New remains from the La Meseta (Thanetian – Lutetian) and Submeseta (Lutetian – Rupelian) formations (Seymour Island, Antarctica) are tentatively assigned to Diomedeidae and Procellariidae (Procellariiformes). Based on the fossil record and several analyses that attempt to explain the evolutionary patterns of Diomedeidae, Notoleptos giglii gen. et sp. nov., based on a small tarsometatarsus, was an ancestral form that lived in Antarctica before the rise of large-sized albatrosses. Subsequent environmental cooling since the late Oligocene could have selected against small body size, to the detriment of small-sized albatrosses like Notoleptos, thus favoring large body size and setting the stage for the development of the specialized albatross flight.

ZooBank ID (LSID): urn:lsid:zoobank.org:pub:C8D9C862-1536-4EA3-8A76-70903AA75639     

Crossvallia unienwillia, a new Spheniscidae (Sphenisciformes, Aves) from the Late Paleocene of Antarctica

Modern penguins are typically small to medium-sized birds, and nearly all known modern specimens are smaller than the most archaic Paleogene relatives. Here we report an incomplete humerus, associated femur and tibiatarsus of a new spheniscid, Crossvallia unienwillia nov. gen. and sp., from the Late Paleocene (~55 million years, Myr) of Antarctica, extending the record of spheniscids by about 15 Myr. Its large size (length ~140 cm) supports the idea that large body size in penguins was acquired independently at different times (Late Paleocene and Late Eocene) under dramatically different environmental conditions. Comparison of its osteological anatomy suggests that the new taxon is closely related to the extinct Anthropornithinae rather than to other penguin lineages.     

Mathematical modeling of appendicular bone growth in glaucous‐winged gulls

Development of locomotor activity is crucial in tetrapods. In birds, this development leads to different functions for hindlimbs and forelimbs. The emergence of walking and flying as very different complex behavior patterns only weeks after hatching provides an interesting case study in animal development. We measured the diaphyseal lengths and midshaft diameters of three wing bones (humerus, ulna, and carpometacarpus) and three leg bones (femur, tibiotarsus, and tarsometatarsus) of 79 juvenile (ages 0–42 days) and 13 adult glaucous‐winged gulls (Larus glaucescens), a semiprecocial species. From a suite of nine alternative mathematical models, we used information‐theoretic criteria to determine the best model(s) for length and diameter of each bone as a function of age; that is, we determined the model(s) that obtained the best tradeoff between the minimized sum of squared residuals and the number of parameters used to fit the model. The Janoschek and Holling III models best described bone growth, with at least one of these models yielding an R² ≥ 0.94 for every dimension except tarsometatarsus diameter (R² = 0.87). We used the best growth models to construct accurate allometric comparisons of the bones. Early maximal absolute growth rates characterize the humerus, femur, and tarsometatarsus, bones that assume adult‐type support functions relatively early during juvenile development. Leg bone lengths exhibit more rapid but less sustained relative growth than wing bone lengths. Wing bone diameters are initially smaller than leg bone diameters, although this relationship is reversed by fledging. Wing bones and the femur approach adult length by fledging but continue to increase in diameter past fledging; the tibiotarsus and tarsometatarsus approach both adult length and diameter by fledging. In short, the pattern of bone growth in this semiprecocial species reflects the changing behavioral needs of the developing organism. J. Morphol., 2009. © 2008 Wiley‐Liss, Inc.     

A remarkable new ‘gruiform’ bird from the Middle Eocene of Messel (Hessen, Germany)

Salmila robusta n. gen., n. sp. (Aves: ‘Gruiformes’) is described from the Middle Eocene of Messel (Hessen, Germany). The new species is assumed to have had a predominantly terrestrial habit. It shares derived features of the carpometacarpus with the recent Psophiidae and some taxa of the Cariamae. In several features, however, it distinctly differs from other ‘gruiform’ birds (e. g. in the larger proximal end of the humerus and in the shape of the margo caudalis of the sternum). AlthoughSalmila robusta has been tentatively assigned to the Cariamae in this study, its exact systematic position deserves further investigations. The rather short tarsometatarsus of the new species resembles that of the ‘Messel-phorusrhacid’Aenigmavis sapea Peters 1987.     

The different breeding strategies of penguins: A review

The 18 penguin species are exclusively and widely distributed in the Southern hemisphere, from the Equator to the Antarctic continent, and are thus submitted to various ecological constraints in their reproductive strategy. This results in a high variability in all aspects of the breeding biology of the different species. Although penguins appear primarily adapted for a marine existence, they remain dependent on land for breeding, rearing young, and moulting. Here we describe and compare the breeding cycle of all the penguin species, highlighting the characteristics of each species in terms of breeding range, population status, threats induced by environmental changes, duration of the different phases of the breeding cycle, mate fidelity, body mass, body height, egg mass and duration of egg formation. We also focus on the breeding cycle of the genus Aptenodytes, since it largely differs from the breeding cycle of most of the other penguin species.     

The last terror birds (Aves,Phorusrhacidae): new evidence from the late Pleistocene of Uruguay

We report new fossil evidence of terror bird survival until the end of the Pleistocene in Uruguay. The new specimens comprise the distal portion of right tarsometatarsus and a left humerus; the latter is assigned to the genus Psilopterus. The sedimentary context of the remains yields a characteristic Pleistocene mammalian association along with numerical age dating giving an undoubted late Pleistocene age (OSL 96,040 ± 6300 years). We also revise and discuss the systematic placement of late Pleistocene phorusrhacid material previously published. The trophic role of terror birds and other South American carnivorous birds in late Pleistocene ecosystems should be revised based on the increasing findings of avian fossil materials.     

The Tarsometatarsus of the Ostrich Struthio camelus: Anatomy,Bone Densities,and Structural Mechanics

Background

The ostrich Struthio camelus reaches the highest speeds of any extant biped, and has been an extraordinary subject for studies of soft-tissue anatomy and dynamics of locomotion. An elongate tarsometatarsus in adult ostriches contributes to their speed. The internal osteology of the tarsometatarsus, and its mechanical response to forces of running, are potentially revealing about ostrich foot function.

Methods/Principal Findings

Computed tomography (CT) reveals anatomy and bone densities in tarsometatarsi of an adult and a young juvenile ostrich. A finite element (FE) model for the adult was constructed with properties of compact and cancellous bone where these respective tissues predominate in the original specimen. The model was subjected to a quasi-static analysis under the midstance ground reaction and muscular forces of a fast run. Anatomy–Metatarsals are divided proximally and distally and unify around a single internal cavity in most adult tarsometatarsus shafts, but the juvenile retains an internal three-part division of metatarsals throughout the element. The juvenile has a sparsely ossified hypotarsus for insertion of the m. fibularis longus, as part of a proximally separate third metatarsal. Bone is denser in all regions of the adult tarsometatarsus, with cancellous bone concentrated at proximal and distal articulations, and highly dense compact bone throughout the shaft. Biomechanics–FE simulations show stress and strain are much greater at midshaft than at force applications, suggesting that shaft bending is the most important stressor of the tarsometatarsus. Contraction of digital flexors, inducing a posterior force at the TMT distal condyles, likely reduces buildup of tensile stresses in the bone by inducing compression at these locations, and counteracts bending loads. Safety factors are high for von Mises stress, consistent with faster running speeds known for ostriches.

Conclusions/Significance

High safety factors suggest that bone densities and anatomy of the ostrich tarsometatarsus confer strength for selectively critical activities, such as fleeing and kicking predators. Anatomical results and FE modeling of the ostrich tarsometatarsus are a useful baseline for testing the structure’s capabilities and constraints for locomotion, through ontogeny and the full step cycle. With this foundation, future analyses can incorporate behaviorally realistic strain rates and distal joint forces, experimental validation, and proximal elements of the ostrich hind limb.     

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.     

Fossil penguin (Aves: Sphenisciformes) cranial material from the Eocene of Seymour Island (Antarctica)

Our knowledge of the cranial morphology of early penguins remains poor, particularly for Paleogene taxa. This paper describes a partial penguin skull and additional isolated cranial elements from the Eocene La Meseta Formation of Seymour Island, Antarctica. These specimens cannot be assigned to named taxa at present, but there is a strong possibility they belong to La Meseta penguins known only from postcranial elements. The skull shares extensive dorsal development of the temporal fossae with extant and fossil Spheniscus and the fossil penguins Paraptenodytes and Marplesornis, indicating the adductor complex was powerful in early penguins. Partial mandibles belonging to a much larger penguin are similar to Paraptenodytes and differ from all living penguins in the lack of a hooked medial process of the articular and the presence of a foramen anterior to the mandibular cotyles. Given the rarity of penguin cranial remains, these specimens provide important new insight into early penguin evolution.     

The complete sequence of the mitochondrial genome of the African Penguin (Spheniscus demersus)

The complete mitochondrial genome of the African Penguin (Spheniscus demersus) was sequenced. The molecule was sequenced via next generation sequencing and primer walking. The size of the genome is 17,346 bp in length. Comparison with the mitochondrial DNA of two other penguin genomes that have so far been reported was conducted namely; Little blue penguin (Eudyptula minor) and the Rockhopper penguin (Eudyptes chrysocome). This analysis made it possible to identify common penguin mitochondrial DNA characteristics. The S. demersus mtDNA genome is very similar, both in composition and length to both the E. chrysocome and E. minor genomes. The gene content of the African penguin mitochondrial genome is typical of vertebrates and all three penguin species have the standard gene order originally identified in the chicken. The control region for S. demersus is located between tRNA-Glu and tRNA-Phe and all three species of penguins contain two sets of similar repeats with varying copy numbers towards the 3′ end of the control region, accounting for the size variance. This is the first report of the complete nucleotide sequence for the mitochondrial genome of the African penguin, S. demersus. These results can be subsequently used to provide information for penguin phylogenetic studies and insights into the evolution of genomes.     

Morphological aspects of the heart of the northern rockhopper penguin (Eudyptes chrysocome moseleyi ): possible implication in diving behavior and ecology?

We compared the heart morphology of the small, deep-diving northern rockhopper penguin to the hearts of small, shallow-diving and large, deep-diving penguin species. The rockhopper penguin had a heart larger than expected for its body mass, and its heart weight/body weight was significantly greater than in the larger Adélie penguin. We found the rockhopper's right ventricle weight/heart weight to be significantly greater than this relationship in both the larger chinstrap and Adélie penguins. The relationship of the right to left ventricular weights in the rockhopper heart is not different to that of the large, deepest-diving emperor penguin. A larger heart in the rockhopper penguin might be related to its diving behavior and ecology if it contributes to diving efficiency during foraging by increasing lung perfusion during surface recovery. This would lead to decreased surface time. Accepted: 20 May 2000     

云南禄丰晚中新世鸟类

云南禄丰晚中新世古猿产地的鸟类化石,共计3目、3科、6属、6种,包括2新属4新种。另外尚有科、目未定之一特殊鸟类标本。     

A new Lower Cretaceous bird from China and tooth reduction in early avian evolution

A new avian genus and species, Zhongjianornis yangi gen. et sp. nov., is reported from the Lower Cretaceous lacustrine deposits of the Jiufotang Formation in Liaoning, northeast China. The new taxon is characterized by possessing the following combination of features: upper and lower jaws toothless, snout pointed, humerus with large and robust deltopectoral crest, second phalanx of the major manual digit longer than the first phalanx, unguals of the alular and major digits of similar length and significantly shorter than the corresponding penultimate phalanges, tibiotarsus slender and more than twice the length of the tarsometatarsus, and metatarsal IV longer than the other metatarsals. Phylogenetic analysis indicates that Zhongjianornis is phylogenetically basal to Confuciusornis and the dominant Mesozoic avian groups, Enantiornithes and Ornithurae, and therefore provides significant new information regarding the diversification of birds in the Early Cretaceous. It also represents the most basal bird that completely lacks teeth, suggesting that tooth loss was more common than expected in early avian evolution and that the avian beak appeared independently in several avian lineages, most probably as a response to selective pressure for weight reduction. Finally, the presence of a significantly enlarged humeral deltopectoral crest suggests that Zhongjianornis shares with other basal birds such as Jeholornis, Sapeornis and Confuciusornis a distinctive mode of adaptation for flight contrasting with that seen in more advanced birds, which instead possess an elongated sternum and a prominent keel.     

Assessing penguin colony size and distribution using digital mapping and satellite remote sensing

Changes in penguin abundance and distribution can be used to understand the response of species to climate change and fisheries pressures, and as a gauge of ecosystem health. Traditionally, population estimates have involved direct counts, but remote sensing and digital mapping methodologies can provide us with alternative techniques for assessing the size and distribution of penguin populations. Here, we demonstrate the use of a field-based digital mapping system (DMS), combining a handheld geographic information system with integrated geographical positioning system as a method for: (a) assessing penguin colony area and (b) ground-truthing colony area as derived from satellite imagery. Work took place at Signy Island, South Orkneys, where colonies of the three congeneric pygoscelid penguins: Adélie Pygoscelis adeliae, chinstrap P. antarctica and gentoo P. papua were surveyed. Colony areas were derived by mapping colony boundaries using the DMS with visual counts of the number of nesting birds made concurrently. Area was found to be a good predictor for number of nests for all three species of penguin. Using a maximum likelihood multivariate classification of remotely sensed satellite imagery (QuickBird2, 18 January 2010; Digital Globe ID: 01001000B90AD00), we were able to identify penguin colonies from the spectral signature of guano and differentiate between colonies of Adélie and chinstrap penguins. The area classified (all species combined) from satellite imagery versus area from DMS data was closely related (R ² = 0.88). Combining these techniques gives a simple and transferrable methodology for examining penguin distribution and abundance at local and regional scales.     

Low MHC variation in the endangered Galápagos penguin (<Emphasis Type="Italic">Spheniscus mendiculus</Emphasis>)

The major histocompatibility complex (MHC) is one of the most polymorphic regions of the genome, likely due to balancing selection acting to maintain alleles over time. Lack of MHC variability has been attributed to factors such as genetic drift in small populations and relaxed selection pressure. The Galápagos penguin (Spheniscus mendiculus), endemic to the Galápagos Islands, is the only penguin that occurs on the equator. It relies upon cold, nutrient-rich upwellings and experiences severe population declines when ocean temperatures rise during El Niño events. These bottlenecks, occurring in an already small population, have likely resulted in reduced genetic diversity in this species. In this study, we used MHC class II exon 2 sequence data from a DRB1-like gene to characterize the amount of genetic variation at the MHC in 30 Galápagos penguins, as well as one Magellanic penguin (S. magellanicus) and two king penguins (Aptenodytes patagonicus), and compared it to that in five other penguin species for which published data exist. We found that the Galápagos penguin had the lowest MHC diversity (as measured by number of polymorphic sites and average divergence among alleles) of the eight penguin species studied. A phylogenetic analysis showed that Galápagos penguin MHC sequences are most closely related to Humboldt penguin (Spheniscus humboldti) sequences, its putative sister species based on other loci. An excess of non-synonymous mutations and a pattern of trans-specific evolution in the neighbor-joining tree suggest that selection is acting on the penguin MHC.