Comparative embryonic development of the skeleton of the domestic turkey (Meleagris gallopavo) and other galliform birds

Ossification sequences are poorly known for birds in general, even for common domestic and experimental species. Such sequences constitute a rich source of data on character evolution, and may even provide phylogenetic information. It is not clear, however, what factors influence ossification sequences and what the relative importance of phylogeny is to the sequences. Galliformes constitute a good group to examine these variables. These birds are osteologically conservative, have precocial young, but have a broad spectrum of body sizes and incubation periods. Here, I describe the embryonic ossification of the skeleton in the domestic turkey (Meleagris gallopavo), and compare it to the domestic chicken (Gallus gallus) and the Japanese quail (Coturnix coturnix). Ossification sequences in this group are not affected by egg size or incubation period. They also appear to be independent of both the spatial location and the embryonic tissue from which the osteogenic cells originated. Accumulation of a wider sample of ossification sequences from more morphologically variable avian taxa will be necessary in order to test functional and phylogenetic effects.     

Evolution and functional significance of derived sternal ossification patterns in ornithothoracine birds

The midline pattern of sternal ossification characteristic of the Cretaceous enantiornithine birds is unique among the Ornithodira, the group containing birds, nonavian dinosaurs and pterosaurs. This has been suggested to indicate that Enantiornithes is not the sister group of Ornithuromorpha, the clade that includes living birds and their close relatives, which would imply rampant convergence in many nonsternal features between enantiornithines and ornithuromorphs. However, detailed comparisons reveal greater similarity between neornithine (i.e. crown group bird) and enantiornithine modes of sternal ossification than previously recognized. Furthermore, a new subadult enantiornithine specimen demonstrates that sternal ossification followed a more typically ornithodiran pattern in basal members of the clade. This new specimen, referable to the Pengornithidae, indicates that the unique ossification pattern observed in other juvenile enantiornithines is derived within Enantiornithes. A similar but clearly distinct pattern appears to have evolved in parallel in the ornithuromorph lineage. The atypical mode of sternal ossification in some derived enantiornithines should be regarded as an autapomorphic condition rather than an indication that enantiornithines are not close relatives of ornithuromorphs. Based on what is known about molecular mechanisms for morphogenesis and the possible selective advantages, the parallel shifts to midline ossification that took place in derived enantiornithines and living neognathous birds appear to have been related to the development of a large ventral keel, which is only present in ornithuromorphs and enantiornithines. Midline ossification can serve to medially reinforce the sternum at a relatively early ontogenetic stage, which would have been especially beneficial during the protracted development of the superprecocial Cretaceous enantiornithines.     

Development and characterization of expressed sequence tags for the turkey (Meleagris gallopavo) genome and comparative sequence analysis with other birds

Twenty-one randomly selected clones from a turkey (Meleagris gallopavo) pituitary complementary DNA (cDNA) library were sequenced to develop expressed sequence tags (ESTs) for this economically important avian species whose genome is among the least understood. Primers specific for the ESTs were used to produce amplicons from the genomic DNA of turkey, chicken (Gallus gallus), guinea fowl (Numidia meleagris), pigeon (Columba domestica), and quail (Corturnix japonica). The amplicons were sequenced and analyzed for sequence variation within- and similarity among-species and with GenBank database sequences. The proportion of shared bases between the turkey sequence and the consensus sequence from each of the other species ranged from 72% to 93% between turkey and pigeon and quail and between turkey and chicken, respectively. The total number of single nucleotide polymorphisms (SNPs) observed ranged from 3 in quail to 18 in chicken out of 4898 and 5265 bases analyzed, respectively. The most frequent nucleotide variation observed was a C-->T transition. Linkage analysis of one such SNP in the backcross progeny of the East Lansing reference DNA panel, localized TUS0005, the chicken sequence derived from primers specific for turkey TUT2E EST, to chromosome 4. The ESTs reported, as well as the SNPs may provide a useful resource for ongoing efforts to develop high utility genome maps for the turkey and chicken. The primers described can also be used as a tool in future investigations directed at further understanding the biology of the guinea fowl, pigeon and quail and their relatedness to the turkey.     

Limb ossification patterns of the ichthyosaur Stenopterygius, and a discussion of the proximal tarsal row of ichthyosaurs and other neodiapsid reptiles

Limb ossification patterns for the Lower Jurassic (Toarcian) ichthyosaur, Stenopterygius , are described. It is found that limb ossification follows a continuous proximal to distal sequence from the propodial elements through to the terminal elements of 1st to 4th digit in the manus and the 1st to 3rd digit in the pes. The 5th manal and 4th pedal digit begin ossification later than more preaxial digits and also show evidence of proximal addition of elements near the distal mesopodial row in a manner consistent with delayed ossification of the 5th distal mesopodial in other diapsids. Ossification of manal elements in the Supernumerary 3–4 (S3-4) digit and the 5th digit appear interdependent; if one or the other is highly ossified, ossification of the other is retarded. The 1st pedal digit is considered to be lost in Stenopterygius and the 4th pedal digit is identified as the 5th digit. Delayed ossification of the mesopodium is not observed. The most preaxial proximal tarsal is identified as the centralc; the remaining proximal tarsals are the astragalus and calcaneum, and it is inferred that the astragalus and calcaneum ossified from within a single proximal cartilage.     

Early larval ontogeny of the Permo-Carboniferous temnospondyl Sclerocephalus

The early larval development of the temnospondyl Sclerocephalus sp. is analyzed, based on 38 specimens from the Lower Rotliegend (Permo-Carboniferous boundary) of the Saar-Nahe Basin (south-west Germany). The study focuses on the smallest larval specimens, which exemplify changes in both proportions and ossification patterns. In comparison with dissorophoid larvae, the skull ossifies more fully and at a much faster rate; the smallest specimens already have completely formed circumorbital bones that are sutured throughout. Sculpturing undergoes two marked changes, first from uniformly pitted to pits of variable size and regional differentiation, and finally to the origin of ridges. The palate of small larvae differs from that of larger specimens in patterns of dentition, having more teeth including a denticle field on the cultriform process. The mandible of small larvae is described for the first time, being narrower than in adults and having three dentigerous coronoid elements. The smallest specimens have poorly ossified neural arches, lack vertebral centra, and have faintly ossified humeri, femora, and very poorly developed distal elements. The posterior ribs, metapodia, and phalanges appeared after the dermal elements of the pectoral girdle, whereas the scapulocoracoid and ischium are absent throughout the larval period. Early growth and differentiation of the limbs and the ilium illustrates the developmental patterning of the appendages, which proceeded from proximal to distal. Dermal squamation is uniform in small stages, consisting of round or oval osteoderms with pronounced growth rings; in large larvae, they start to differentiate in certain body regions.     

Osteological development of Cope's Gray Treefrog,Hyla chrysoscelis

Hyla chrysoscelis, Cope's Gray Treefrog, is a generalized treefrog found throughout much of east‐central North America. Although it is a model for many behavioural and ecological studies, little is known of its skeletal morphology or development. Herein, we describe the postembryonic skeletal development and adult osteology of H. chrysoscelis. The adult skull is well ossified with slight dermal ornamentation, the postcranial and tadpole skeletons are fairly non‐distinct with no obvious novel morphologies, and the Gosner stage by which bony elements first appear varies. We compare the rank order sequence of ossification to that of its sibling species Hyla versicolor and use examples from this study to demonstrate current complications with conducting ossification sequence meta‐analyses.     

阿伦膦酸钠与阿法骨化醇联合治疗妇女绝经后骨质疏松症的疗效

目的:探讨妇女绝经后骨质疏松症(PMO)应用阿伦膦酸钠(ALN)与阿法骨化醇联合治疗的临床效果。方法:选取我院门诊2013年1月~2015年3月收治的98例PMO患者,按照数字随机表法均分为两组。对照组:在常规钙剂基础上,给予阿法骨化醇治疗;观察组:在对照组基础上,给予ALN治疗。记录比较两组治疗前后Ward三角区、股骨颈及腰椎L2~L4骨密度(BMD)水平,治疗前后翻身、前屈后伸及自发性腰背痛视觉模拟评分法(VAS)评分,治疗前后血清骨钙素(BGP)、骨碱性磷酸酶(BALP)、Ⅰ型胶原C端肽(CTX)及抗酒石酸酸性磷酸酶5b(TRACP-5b)水平,用药期间不良反应。结果:观察组治疗后Ward三角区、股骨颈及腰椎L2~L4 BMD水平与治疗前比较,均明显改善(P0.05),且观察组改善程度更加明显(P0.05);观察组治疗后翻身、前屈后伸及自发性腰背痛VAS评分与治疗前比较,均显著降低(P0.01),且观察组改善程度更为显著(P0.01);观察组治疗后血清BGP、BALP、CTX及TRACP-5b水平与治疗前比较,均显著下降(P0.01),且观察组改善幅度更为显著(P0.01)。结论:妇女PMO应用ALN与阿法骨化醇联合治疗更能显著提高BMD水平,缓解患者疼痛症状,改善骨代谢指标,疗效切实,安全可靠,具有较高临床参考价值。     

Metaves,Mirandornithes, Strisores and other novelties – a critical review of the higher‐level phylogeny of neornithine birds

Recent hypotheses on the higher‐level phylogeny of modern birds are reviewed, and areas of agreement and major conflict are detailed, with emphasis being put on congruence among independent molecular and morphological data sets. Although molecular data significantly contributed to a better understanding of avian phylogeny, they do not seem to be free of homoplasy and caution is warranted in the interpretation of some results. The recently proposed ‘Metaves’ clade is likely to be an artefact of the β‐fibrinogen gene, and current molecular data do not yield well‐supported phylogenies for some groups whose interrelationships can be resolved with morphological evidence. There exists, however, congruent and strong molecular evidence for several novel clades that were not recognized by morphologists before, and to ease future discussions the terms Picocoraciae (non‐leptosomid ‘Coraciiformes’ and Piciformes) and Aequornithes (‘waterbird assemblage’) are introduced. Molecular studies further congruently recover some clades, which have not yet been adequately appreciated and are outlined in the present review.     

Larval and post-larval development of the branchiopod clam shrimp Cyclestheria hislopi (Baird, 1859) (Crustacea, Branchiopoda, Conchostraca, Spinicaudata)

The larval and post-larval development of Cyclestheria hislopi is examined by SEM. There are at least nine stages (excluding the adult) – six larval and three post-larval stages. The first four stages are passed within the egg-membrane. The larval and the post-larval phase are separated by a profound change in morphology that takes place between stages VI and VII. The larva shifts from a dorso-ventrally flattened 'larval' appearance up to stage VI to a laterally flattened, more 'adult' appearance from stage VII. New morphological data have been revealed by this study, including (1) a large and globular larval dorsal organ; (2) the carapace starts its development from the segments of the first and second maxillae; (3) the anterior ramus of the second antenna in adult Cyclestheria hislopi is the endopod, and the posterior ramus the exopod. Direct development of the brood in Cyclestheria hislopi – unique among conchostracans – is compared with that of the Cladocera. If Cyclestheria is the sister group to the Cladocera, as favoured in this work, the classical neoteny theory of the Cladocera must be reconsidered, as there is no particular similarity between any adults of the Cladocera and any of the larval stages of Cyclestheria . It is suggested that Cyclestheria displays the type of development present in a cladoceran ancestor. A comparison between Cyclestheria and the Upper Cambrian 'Orsten' fossil Rehbachiella kinnekullensis reveals a remarkable similarity in the endite morphology of the trunk limbs.     

Ossification and midline shape changes of the human fetal cranial base

An appreciation of ontogenetic changes to the cranial base is important for understanding the evolution of modern human skull form. Using geometric morphometric techniques, this study explores midline shape variations of the basicranium and midface during human prenatal ontogeny. In particular, the analysis sets out to explore shape variations associated with endochondral ossification and to reassess shape variations previously observed on the basis of angular measures.Fifty-four formalin-preserved human fetuses were imaged using high-resolution MRI. Coordinates for 10 landmarks defining the midline basicranium and midface were acquired and areas of ossification in the midline basioccipital, basisphenoid, and presphenoid cartilages were measured as percentages of overall cranial base area. The results show shape variations with increasing fetal size that are consistent with cranial base retroflexion, anterior facial projection and dorsal facial rotation. These growth variations are centered on the midsphenoid area and are associated with disproportionate variations of sphenoid height and length. Small but significant correlations were observed between ossification of the presphenoid cartilage and components of shape that described, among other variations, sphenoid shortening. While ossification cannot be directly linked with the shape variations observed, it seems likely that bone formation plays a role in modulating the influence of other factors on the fetal cranial base.     

Growth patterns in brooding dinosaurs reveals the timing of sexual maturity in non-avian dinosaurs and genesis of the avian condition

The timing of sexual maturation in non-avian dinosaurs is not known. In extant squamates and crocodilians it occurs in conjunction with the initial slowing of growth rates as adult size is approached. In birds (living dinosaurs) on the other hand, reproductive activity begins well after somatic maturity. Here we used growth line counts and spacing in all of the known brooding non-avian dinosaurs to determine the stages of development when they perished. It was revealed that sexual maturation occurred well before full adult size was reached-the primitive reptilian condition. In this sense, the life history and physiology of non-avian dinosaurs was not like that of modern birds. Palaeobiological ramifications of these findings include the potential to deduce reproductive lifespan, fecundity and reproductive population sizes in non-avian dinosaurs, as well as aid in the identification of secondary sexual characteristics.     

Exceptional avian herbivores: multiple transitions toward herbivory in the bird order Anseriformes and its correlation with body mass

Herbivory is rare among birds and is usually thought to have evolved predominately among large, flightless birds due to energetic constraints or an association with increased body mass. Nearly all members of the bird order Anseriformes, which includes ducks, geese, and swans, are flighted and many are predominately herbivorous. However, it is unknown whether herbivory represents a derived state for the order and how many times a predominately herbivorous diet may have evolved. Compiling data from over 200 published diet studies to create a continuous character for herbivory, models of trait evolution support at least five independent transitions toward a predominately herbivorous diet in Anseriformes. Although a nonphylogenetic correlation test recovers a significant positive correlation between herbivory and body mass, this correlation is not significant when accounting for phylogeny. These results indicate a lack of support for the hypothesis that a larger body mass confers an advantage in the digestion of low‐quality diets but does not exclude the possibility that shifts to a more abundant food source have driven shifts toward herbivory in other bird lineages. The exceptional number of transitions toward a more herbivorous diet in Anseriformes and lack of correlation with body mass prompts a reinterpretation of the relatively infrequent origination of herbivory among flighted birds.     

Higher-order phylogeny of modern birds (Theropoda, Aves: Neornithes) based on comparative anatomy. II. Analysis and discussion

In recent years, avian systematics has been characterized by a diminished reliance on morphological cladistics of modern taxa, intensive palaeornithogical research stimulated by new discoveries and an inundation by analyses based on DNA sequences. Unfortunately, in contrast to significant insights into basal origins, the broad picture of neornithine phylogeny remains largely unresolved. Morphological studies have emphasized characters of use in palaeontological contexts. Molecular studies, following disillusionment with the pioneering, but non-cladistic, work of Sibley and Ahlquist, have differed markedly from each other and from morphological works in both methods and findings. Consequently, at the turn of the millennium, points of robust agreement among schools concerning higher-order neornithine phylogeny have been limited to the two basalmost and several mid-level, primary groups. This paper describes a phylogenetic (cladistic) analysis of 150 taxa of Neornithes, including exemplars from all non-passeriform families, and subordinal representatives of Passeriformes. Thirty-five outgroup taxa encompassing Crocodylia, predominately theropod Dinosauria, and selected Mesozoic birds were used to root the trees. Based on study of specimens and the literature, 2954 morphological characters were defined; these characters have been described in a companion work, approximately one-third of which were multistate (i.e. comprised at least three states), and states within more than one-half of these multistate characters were ordered for analysis. Complete heuristic searches using 10 000 random-addition replicates recovered a total solution set of 97 well-resolved, most-parsimonious trees (MPTs). The set of MPTs was confirmed by an expanded heuristic search based on 10 000 random-addition replicates and a full ratchet-augmented exploration to ascertain global optima. A strict consensus tree of MPTs included only six trichotomies, i.e. nodes differing topologically among MPTs. Bootstrapping (based on 10 000 replicates) percentages and ratchet-minimized support (Bremer) indices indicated most nodes to be robust. Several fossil Neornithes (e.g. Dinornithiformes, Aepyornithiformes) were placed within the ingroup a posteriori either through unconstrained, heursitic searches based on the complete matrix augmented by these taxa separately or using backbone-constraints. Analysis confirmed the topology among outgroup Theropoda and achieved robust resolution at virtually all levels of the Neornithes. Findings included monophyly of the palaeognathous birds, comprising the sister taxa Tinamiformes and ratites, respectively, and the Anseriformes and Galliformes as monophyletic sister-groups, together forming the sister-group to other Neornithes exclusive of the Palaeognathae (Neoaves). Noteworthy inferences include: (i) the sister-group to remaining Neoaves comprises a diversity of marine and wading birds; (ii) Podicipedidae are the sister-group of Gaviidae, and not closely related to the Phoenicopteridae, as recently suggested; (iii) the traditional Pelecaniformes, including the shoebill (Balaeniceps rex) as sister-taxon to other members, are monophyletic; (iv) traditional Ciconiiformes are monophyletic; (v) Strigiformes and Falconiformes are sister-groups; (vi) Cathartidae is the sister-group of the remaining Falconiformes; (vii) Ralliformes (Rallidae and Heliornithidae) are the sister-group to the monophyletic Charadriiformes, with the traditionally composed Gruiformes and Turniciformes (Turnicidae and Mesitornithidae) sequentially paraphyletic to the entire foregoing clade; (viii) Opisthocomus hoazin is the sister-taxon to the Cuculiformes (including the Musophagidae); (ix) traditional Caprimulgiformes are monophyletic and the sister-group of the Apodiformes; (x) Trogoniformes are the sister-group of Coliiformes; (xi) Coraciiformes, Piciformes and Passeriformes are mutually monophyletic and closely related; and (xii) the Galbulae are retained within the Piciformes. Unresolved portions of the Neornithes (nodes having more than one most-parsimonious solution) comprised three parts of the tree: (a) several interfamilial nodes within the Charadriiformes; (b) a trichotomy comprising the (i) Psittaciformes, (ii) Columbiformes and (iii) Trogonomorphae (Trogoniformes, Coliiformes) + Passerimorphae (Coraciiformes, Piciformes, Passeriformes); and (c) a trichotomy comprising the Coraciiformes, Piciformes and Passeriformes. The remaining polytomies were among outgroups, although several of the highest-order nodes were only marginally supported; however, the majority of nodes were resolved and met or surpassed conventional standards of support. Quantitative comparisons with alternative hypotheses, examination of highly supportive and diagnostic characters for higher taxa, correspondences with prior studies, complementarity and philosophical differences with palaeontological phylogenetics, promises and challenges of palaeogeography and calibration of evolutionary rates of birds, and classes of promising evidence and future directions of study are reviewed. Homology, as applied to avian examples of apparent homologues, is considered in terms of recent theory, and a revised annotated classification of higher-order taxa of Neornithes and other closely related Theropoda is proposed. (c) 2007 The Linnean Society of London, Zoological Journal of the Linnean Society, 2007, 149, 1-95.     

The fossil record and limb disparity of enantiornithines, the dominant flying birds of the Cretaceous

Morphometric and stratigraphic analyses that encompass the known fossil record of enantiornithine birds (Enantiornithes) are presented. These predominantly flighted taxa were the dominant birds of the second half of the Mesozoic; the enantiornithine lineage is known to have lasted for at least 60 million years (Ma), up until the end of the Cretaceous. Analyses of fossil record dynamics show that enantiornithine 'collectorship' since the 1980s approaches an exponential distribution, indicating that an asymptote in proportion of specimens has yet to be achieved. Data demonstrate that the fossil record of enantiornithines is complete enough for the extraction of biological patterns. Comparison of the available fossil specimens with a large data set of modern bird (Neornithes) limb proportions also illustrates that the known forelimb proportions of enantiornithines fall within the range of extant taxa; thus these birds likely encompassed the range of flight styles of extant birds. In contrast, most enantiornithines had hindlimb proportions that differ from any extant taxa. To explore this, ternary diagrams are used to graph enantiornithine limb variation and to identify some morphological oddities ( Otogornis , Gobipteryx ); taxa not directly comparable to modern birds. These exceptions are interesting – although anatomically uniform, and similar to extant avians in their wing proportions, some fossil enantiornithines likely had flight styles not seen among their living counterparts.     

Development of the osteocranium in the suckermouth armored catfish Ancistrus cf. triradiatus (Loricariidae, siluriformes)

The development of the osteocranium of the suckermouth armored catfish Ancistrus cf. triradiatus is described based on specimens ranging from prehatching stages to juvenile stages where the osteocranium is more or less fully formed. The first bony elements that arise are the opercle, jaws, and lateralmost branchiostegal rays, as well as the basioccipital and parasphenoid in the skull floor. The supracleithrum and the membranous and perichondral pterotic components form one large, double-layered skull bone during ontogeny, without clear evidence of the involvement of a supratemporal. The Baudelot's ligament ossifies from two sides, i.e., from the basioccipital medially and the supracleithrum laterally. The lower jaw consists of a dentary, mentomeckelian, and angulo-articular, which all soon fuse. The parurohyal, formed by the fusion of a ventral sesamoid bone and a dorsal cartilage element associated with the first basibranchial, is pierced by a vein, unlike in some other siluriforms. The interhyal cartilage disappears during ontogeny; medially of it, a small sesamoid bone appears in a ligament. The largest, canal-bearing cheek plate is not homologous to the interopercle. The results of the present research, with emphasis on bone formations and homologies, are compared with studies on related catfishes.