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Beta‐microseminoproteins (MSMBs) are small disulfide‐rich proteins that are conserved among vertebrates. These proteins exhibit diverse biological activities and were mainly reported to play a role in male fertility, immunity, and embryogenesis. In this work, we focused on the chicken MSMB3 protein that was previously depicted as an egg antibacterial protein. We report that MSMB3 protein is exclusively expressed in the reproductive tissues of laying hens (in contrast to chicken MSMB1 and MSMB2 paralogs), to be incorporated in the egg white during the process of egg formation. We also showed that chicken MSMB3 possesses highly conserved orthologs in bird species, including Neognathae and Palaeognathae. Chicken MSMB3 was purified from egg white using heparin affinity chromatography and was analyzed by top‐down and bottom‐up proteomics. Several proteoforms could be characterized, and a homodimer was further evidenced by NMR spectroscopy. The X‐ray structure of chicken MSMB3 was solved for the first time, revealing that this protein adopts a novel dimeric arrangement. The highly cationic MSMB3 protein exhibits a distinct electrostatic distribution compared with chicken MSMB1 and MSMB2 structural models, and with published mammalian MSMB structures. The specific incorporation of MSMB3 paralog in the egg, and its phylogenetic conservation in birds together with its peculiar homodimer arrangement and physicochemical properties, suggests that the MSMB3 protein has evolved to play a critical role during the embryonic development of avian species. These new data are likely to stimulate research to elucidate the structure/function relationships of MSMB paralogs and orthologs in the animal kingdom., Beta‐microseminoproteins (MSMBs) are small proteins conserved among vertebrates. The chicken genome contains three MSMB paralogs, which exhibit distinct tissue expression profiles, with chicken beta‐MSMB 3 (MSMB3) protein being egg‐specific. Collectively, results from protein sequence analysis, mass spectrometry analysis, and 3D structure data highlight that chicken MSMB3 possesses specific features that may be related to its presumed role in avian reproduction. ​

Although the development of the avian skeleton has attracted considerable attention, most of the studies have been concentrated on the embryonic period, while studies on the postnatal period are rare. We studied the postnatal development of the skeleton in two phylogenetically distant birds, an altricial passerine Acrocephalus scirpaceus and a semiprecocial charadriiform Chroicocephalus ridibundus. The neonates of the former, despite being altricial, have well-ossified skeleton-the degree of development approaches that of the semiprecocial gull. However, after hatching the limb bones (particularly those of the hind limb) ossify earlier in the gull which is probably related to faster acquisition of locomotor abilities. We have observed that, in contrast to previous reports from neognathous birds, in the ankle of the gull, the ascending process fuses with the astragalus rather than with the calcaneum. This type of development is present in palaeognaths and nonavian dinosaurs but has not yet been reported in neognaths. This indicates a greater diversity within Neognathae and suggests a more complex scenario for the evolution of the avian ankle. However, data from a greater number of species are needed to establish the developmental sequence ancestral for neognathous birds. Furthermore, the sequence of bone fusions in the wrist of Acrocephalus is similar to the fossil-documented evolutionary sequence observed in the phylogeny of early birds, with the semilunate carpal and major metacarpal fusing first, followed by the alular metacarpal fusing with the major metacarpal and then the major and minor metacarpal fusing proximally. These data underscore the importance of developmental studies for reconstructing the evolutionary history.

Crown birds are subdivided into two main groups, Palaeognathae and Neognathae, that can be distinguished, among other means, by the organization of the bones in their pterygoidpalatine complex (PPC). Shape variation of the vomer, which is the most anterior part of the PPC, was recently analysed with help of geometric morphometrics to discover morphological differences between palaeognath and neognath birds. Based on this study, the vomer was identified as sufficient to distinguish the two main groups (and even some inclusive neognath groups) and their cranial kinetic system. As there are notable size differences between the skulls of Palaeognathae and Neognathae, we here investigate the impact of allometry on vomeral shape and its implication for taxonomic classification by re-analysing the data of the previous study. Different types of multivariate statistical analyses reveal that taxonomic identification based on vomeral shape is strongly impaired by allometry, as the error of correct identification is high when shape data is corrected for size. This finding is evidenced by a great overlap between palaeognath and neognath subclades in morphospace. Correct taxonomic identification is further impeded by the convergent presence of a flattened vomeral morphotype in multiple neognath subclades. As the evolution of cranial kinesis has been linked to vomeral shape in the original study, the correlation between shape and size of the vomer across different bird groups found in the present study questions this conclusion. In fact, cranial kinesis in crown birds results from the loss of the jugal-postorbital bar in the temporal region and ectopterygoid in the PPC and the combination of a mobilized quadratezygomatic arch complex and a flexible PPC. Therefore, we can conclude that vomer shape itself is not a suitable proxy for exploring the evolution of cranial kinesis in crown birds and their ancestors. In contrast, the evolution of cranial kinesis needs to be viewed in context of the braincase, quadrate-zygomatic arch and the whole pterygoid-palatine complex.

Most living birds exhibit cranial kinesis—movement between the rostrum and braincase—in which force is transferred through the palatal and jugal bars. The palate alone distinguishes the Paleognathae from the Neognathae, with cranial kinesis more developed in neognaths. Most previous palatal studies were based on 2D data and rarely incorporated data from stem birds despite great interest in their kinetic abilities. Here we reconstruct the vomer of the Early Cretaceous stem bird Sapeornis and the troodontid Sinovenator , taxa spanning the dinosaur–bird transition. A 3D shape analysis including these paravians and an extensive sampling of neornithines reveals their strong similarity to paleognaths and indicates that morphological differences in the vomer between paleognaths and neognaths are intimately related to their different kinetic abilities. These results suggest the skull of Mesozoic paravians lacked the kinetic abilities observed in neognaths, a conclusion also supported by our identification of an ectopterygoid in Sapeornis here. We conclude that cranial kinesis evolved relatively late, likely an innovation of the Neognathae, and is linked to the transformation of the vomer. This transformation increased palatal mobility, enabling the evolution of a diversity of kinetic mechanisms and ultimately contributing to the extraordinary evolutionary success of this clade.

Conselho Nacional de Desenvolvimento Cient?fico e Tecnol?gico (CNPq, Proc. PDE 204792/2018-5 ), Funda??o de Amparo a Pesquisa do Estado do Rio Grande do Sul (FAPERGS, 16/2551-000485-7 and the Biotechnology and Biological Sciences Research Council UK (BB/K008226/1. University of Kent. School of Biosciences. Canterbury, NJ, UK / Universidade Federal do Rio Grande do Sul. Departamento de Gen?tica. Porto Alegre, RS, Brazil. Universidade Federal do Pampa. Laborat?rio de Diversidade Gen?tica Animal. S?o Gabriel, RS, Brazil. Universidade Federal Rural da Amaz?nia. Laborat?rio de Reprodu??o Animal. Parauapebas, PA, Brazil. University of Kent. School of Biosciences. Canterbury, NJ, UK. Universidade Federal do Pampa. Laborat?rio de Diversidade Gen?tica Animal. S?o Gabriel, RS, Brazil. Universidade Federal do Pampa. Laborat?rio de Diversidade Gen?tica Animal. S?o Gabriel, RS, Brazil. Universidade Federal do Rio Grande do Sul. Departamento de Gen?tica. Porto Alegre, RS, Brazil. Minist?rio da Sa?de. Secretaria de Vigil?ncia em Sa?de. Instituto Evandro Chagas. Laborat?rio de Cultura de Tecidos e Citogen?tica. Ananindeua, PA, Brasil / Universidade Federal do Par?. Instituto de Ci?ncias Exatas e Naturais. Bel?m, PA, Brazil. University of Kent. School of Biosciences. Canterbury, NJ, UK. University of Kent. School of Biosciences. Canterbury, NJ, UK. Interchromosomal rearrangements involving microchromosomes are rare events in birds. To date, they have been found mostly in Psittaciformes, Falconiformes, and Cuculiformes, although only a few orders have been analyzed. Hence, cytogenomic studies focusing on microchromosomes in species belonging to different bird orders are essential to shed more light on the avian chromosome and karyotype evolution. Based on this, we performed a comparative chromosome mapping for chicken microchromosomes 10 to 28 using interspecies BAC-based FISH hybridization in five species, representing four Neoaves orders (Caprimulgiformes, Piciformes, Suliformes, and Trogoniformes). Our results suggest that the ancestral microchromosomal syntenies are conserved in Pteroglossus inscriptus (Piciformes), Ramphastos tucanus tucanus (Piciformes), and Trogon surrucura surrucura (Trogoniformes). On the other hand, chromosome reorganization in Phalacrocorax brasilianus (Suliformes) and Hydropsalis torquata (Caprimulgiformes) included fusions involving both macro- and microchromosomes. Fissions in macrochromosomes were observed in P. brasilianus and H. torquata. Relevant hypothetical Neognathae and Neoaves ancestral karyotypes were reconstructed to trace these rearrangements. We found no interchromosomal rearrangement involving microchromosomes to be shared between avian orders where rearrangements were detected. Our findings suggest that convergent evolution involving microchromosomal change is a rare event in birds and may be appropriate in cytotaxonomic inferences in orders where these rearrangements occurred.

Avian genomes typically consist of ~10 pairs of macro- and ~30 pairs of microchromosomes. While inter-chromosomally, a pattern emerges of very little change (with notable exceptions) throughout evolution, intrachromosomal changes remain relatively poorly studied. To rectify this, here we use a pan-avian universally hybridising set of 74 chicken bacterial artificial chromosome (BAC) probes on the macrochromosomes of eight bird species: common blackbird, Atlantic canary, Eurasian woodcock, helmeted guinea fowl, houbara bustard, mallard duck, and rock dove. A combination of molecular cytogenetic, bioinformatics, and mathematical analyses allowed the building of comparative cytogenetic maps, reconstruction of a putative Neognathae ancestor, and assessment of chromosome rearrangement patterns and phylogenetic relationships in the studied neognath lineages. We observe that, as with our previous studies, chicken appears to have the karyotype most similar to the ancestor, however, previous reports of an increased rate of intrachromosomal change in Passeriformes (songbirds) appear not to be the case in our dataset. The use of this universally hybridizing probe set is applicable not only for the re-tracing of avian karyotype evolution but, potentially, for reconstructing genome assemblies.

The comparative morphology of juvenile avian skulls is poorly known. Here, we survey the shape of the squamosal (os squamosum) across juvenile skulls of avian higher-level clades. In all palaeognathous birds, the rostral end of the squamosal does not surpass the parietal and does not reach the frontal. This morphology is likely to be plesiomorphic for neornithine birds. A short squamosal also occurs in some Neognathae, but in most neognathous birds the squamosal contacts the frontal, and in some taxa the bone is strongly elongated and distinctly surpasses the parietal. Some clades show a notable variation in squamosal morphology. This is, for example, true for Strigiformes, where the taxon Athene differs from the other examined owls in squamosal size, and for the Passeriformes, where Old World Suboscines are characterized by a distinctive squamosal morphology. A unique derived squamosal morphology is for the first time reported for the Apodidae and Hemiprocnidae, in which the bone forms a elongated rostral process that runs along most of the orbital rim. In non-avian theropods, the squamosal articulates with the postorbital and delimits the upper temporal opening. Extant birds lack a postorbital, but a topological correlation between the squamosal and the postorbital process is maintained in most taxa of the Neognathae. The phylogenetic significance of squamosal morphology is diminished by the fact that closely related taxa often show very disparate shapes of the bone, and squamosal morphology appears to be determined by multiple functional constraints including skull geometry, brain morphology and, possibly, nostril type.

Coordena??o de Aperfei?oamento de Pessoal de Nivel Superior Universidade Federal do Paran?. Departamento de Gen?tica. Curitiba, PR, Brazil. Universidade Federal do Pampa. S?o Gabriel, RS, Brazil. Universidade Federal do Pampa. S?o Gabriel, RS, Brazil. Universidade Federal do Par?. Bel?m, PA, Brazil / Minist?rio da Sa?de. Secretaria de Vigil?ncia em Sa?de. Instituto Evandro Chagas. Laborat?rio Cultura de Tecidos e Citogen?tica. Ananindeua, PA, Brasil. Universidade Federal do Rio Grande do Sul. Instituto de Bioci?ncias. Porto Alegre, RS, Brazil. Universidade Federal do Pampa. S?o Gabriel, RS, Brazil. Universidade Federal do Pampa. S?o Gabriel, RS, Brazil. Universidade Federal do Paran?. Departamento de Gen?tica. Curitiba, PR, Brazil. The distribution of 45S rDNA cluster in avian karyotypes varies in different aspects, such as position, number of bearer chromosomes, and bearers being macro- or microchromosomes. The present study investigated the patterns of variation in the 45S rDNA-bearer chromosomes of birds in order to understand the evolutionary dynamics of the cluster configuration and its contribution to the evolution of bird karyotypes. A total of 73 bird species were analyzed, including both published data and species for which rDNA-FISH was conducted for the first time. In most birds, the 45S rDNA clusters were located in a single pair of microchromosomes. Hence, the location of 45S rDNA in macrochromosomes, observed only in Neognathae species, seems to be a derived state, probably the result of chromosomal fusion between microchromosomes and distinct macrochromosomes. Additionally, the 45S rDNA was observed in multiple microchromosomes in different branches of the bird phylogeny, suggesting recurrence of dispersion processeses, such as duplications and translocations. Overall, this study indicated that the redistribution of the 45S rDNA sites in bird chromosomes followed different evolutionary trajectories with respect to each lineage of the class Aves.

The avian centrifugal visual system, which projects from the brain to the retina, has been intensively studied in several Neognathous birds that have a distinct isthmo-optic nucleus (ION). However, birds of the order Palaeognathae seem to lack a proper ION in histologically stained brain sections. We had previously reported in the palaeognathous Chilean Tinamou (Nothoprocta perdicaria) that intraocular injections of Cholera Toxin B subunit retrogradely label a considerable number of neurons, which form a diffuse isthmo-optic complex (IOC). In order to better understand how this IOC-based centrifugal visual system is organized, we have studied its major components by means of in vivo and in vitro tracing experiments. Our results show that the IOC, though structurally less organized than an ION, possesses a dense core region consisting of multipolar neurons. It receives afferents from neurons in L10a of the optic tectum, which are distributed with a wider interneuronal spacing than in Neognathae. The tecto-IOC terminals are delicate and divergent, unlike the prominent convergent tecto-ION terminals in Neognathae. The centrifugal IOC terminals in the retina are exclusively divergent, resembling the terminals from “ectopic” centrifugal neurons in Neognathae. We conclude that the Tinamou's IOC participates in a comparable general IOC-retina-TeO-IOC circuitry as the neognathous ION. However, the connections between the components are structurally different and their divergent character suggests a lower spatial resolution. Our findings call for further comparative studies in a broad range of species for advancing our understanding of the evolution, plasticity and functional roles of the avian centrifugal visual system.

We evaluated the variation in absolute size and shape of birds’ eggs and the effects of developmental mode and phylogenetic relatedness on these traits. Eggs were characterized by length, diameter, and three indices of egg shape. Indices of egg shape were calculated as the ratio of radii that described the curvature of pointed end (cloacal zone), blunt end (infundibular zone), and lateral zone to egg diameter. We found that eggs shape was less variable than the absolute size of eggs. Index of the cloacal zone was the most changeable and index of the infundibular zone was very conservative. Size and shape of eggs could be better explained with phylogenetic relatedness than developmental mode.

The first fossil remains of vertebrates, invertebrates, plants and palynomorphs of the Chorrillo Formation (Austral Basin), about 30km to the SW of the town of El Calafate (Province of Santa Cruz), are described. Fossils include the elasmarian (basal Iguanodontia) Isasicursor santacrucensis gen. et sp. nov., the large titanosaur Nullotitan glaciaris gen. et sp. nov., both large and small Megaraptoridae indet., and fragments of sauropod and theropod eggshells. The list of vertebrates is also composed by the Neognathae Kookne yeutensis gen. et sp. nov., two isolated caudal vertebrae of Mammalia indet., and isolated teeth of a large mosasaur. Remains of fishes, anurans, turtles, and snakes are represented by fragmentary material of low taxonomical value, with the exception of remains belonging to Calyptocephalellidae. On the other hand, a remarkable diversity of terrestrial and freshwater gastropods has been documented, as well as fossil woods and palinological assemblages. The Chorrillo Formation continues south, in the Las Chinas River valley, southern Chile, where it is called Dorotea Formation. Both units share in their lower two thirds abundant materials of titanosaurs, whose remains cease to appear in the upper third, registering only elasmarians (Chorrillo Formation) and hadrosaurs (Dorotea Formation). Above both units there are levels with remains of invertebrates and marine reptiles. It is striking that the dinosaurs of the lower two thirds of the Chorrillo and Dorotea formations are represented by large basal titanosaurs and Megaraptoridae coelurosaurs, being the Saltasaurinae and Aeolosaurinae sauropods and Abelisauridae theropods totally absent. In contrast, these taxa are dominant components in sedimentary units of central and northern Patagonia (e.g., Allen, Los Alamitos, La Colonia formations). Such differences could reflect, in part, a greater antiquity (i.e., late Campanian-early Maastrichtian) for the Chorrillo fossils, or, more probably, different environmental conditions. Thus, knowledge of the biota of the southern tip of Patagonia is expanded, particularly those temporarily close to the K-Pg boundary. Fil: Novas, Fernando Emilio. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentina Fil: Agnolin, Federico. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentina. Universidad Maimónides; Argentina. Fundación de Historia Natural Félix de Azara; Argentina Fil: Rozadilla, Sebastian. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentina Fil: Aranciaga Rolando, Alexis Mauro. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentina Fil: Brissón Egli, Federico. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentina Fil: Motta, Matias Javier. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentina Fil: Cerroni, Mauricio Andrés. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentina Fil: Ezcurra, Martin Daniel. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentina Fil: Martinelli, Agustín Guillermo. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentina Fil: D'angelo, Julia Soledad. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentina Fil: Alvarez Herrera, Gerardo Paulino. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentina Fil: Gentil, Adriel Roberto. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentina Fil: Bogan, Sergio. Universidad Maimónides; Argentina Fil: Chimento, Nicolás Roberto. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentina Fil: García Marsa, Jordi Alexis. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentina Fil: Lo Coco, Gastón Ernesto. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentina Fil: Miquel, Sergio Eduardo. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentina Fil: Brito, Fátima Fernanda. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentina Fil: Vera, Ezequiel Ignacio. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentina. Universidad de Buenos Aires; Argentina Fil: Perez Loinaze, Valeria Susana. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentina Fil: Fernández, Mariela. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigaciones en Biodiversidad y Medioambiente. Universidad Nacional del Comahue. Centro Regional Universidad Bariloche. Instituto de Investigaciones en Biodiversidad y Medioambiente; Argentina Fil: Salgado, Leonardo. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigación en Paleobiología y Geología; Argentina. Universidad Nacional de Río Negro; Argentina

Summary The Palaeognathae comprise the flightless ratites and the volant tinamous, and together with the Neognathae constitute the extant members of class Aves. It is commonly believed that Palaeognathae originated in Gondwana since most of the living species are found in the Southern Hemisphere [1–3]. However, this hypothesis has been questioned because the fossil paleognaths are mostly from the Northern Hemisphere in their earliest time (Paleocene) and possessed many putative ancestral characters [4]. Uncertainties regarding the origin and evolution of Palaeognathae stem from the difficulty in estimating their divergence times [1, 2] and their remarkable morphological convergence. Here, we recovered nuclear genome fragments from extinct elephant birds, which enabled us to reconstruct a reliable phylogenomic time tree for the Palaeognathae. Based on the tree, we identified homoplasies in morphological traits of paleognaths and reconstructed their morphology-based phylogeny including fossil species without molecular data. In contrast to the prevailing theories, the fossil paleognaths from the Northern Hemisphere were placed as the basal lineages. Combined with our stable divergence time estimates that enabled a valid argument regarding the correlation with geological events, we propose a new evolutionary scenario that contradicts the traditional view. The ancestral Palaeognathae were volant, as estimated from their molecular evolutionary rates, and originated during the Late Cretaceous in the Northern Hemisphere. They migrated to the Southern Hemisphere and speciated explosively around the Cretaceous-Paleogene boundary. They then extended their distribution to the Gondwana-derived landmasses, such as New Zealand and Madagascar, by overseas dispersal. Gigantism subsequently occurred independently on each landmass.

As a result of suppressed recombination, heterogametic sex chromosomes (either Y or W) are usually assumed to gradually shorten over evolutionary time as a way to remove accumulated mutations. However, suppressed recombination removes the most obvious mechanism for excising portions of sex chromosomes. We examined ratios of W/Z chromosome size across 224 bird species from 146 genera. Much of the data were obtained from a previous study (Rutkowska et al. 2012. Biology Letters 8: 636–638), who, similar to ourselves, found no gradual decrease in W chromosome length over evolutionary time. However, we show an abrupt decrease in W chromosome length at or just after the phylogenetic split between the two extant bird superorders, Paleognathae and Neognathae, indicating that the key to understanding sex chromosome evolution may have little to do with gradual suppression of recombination.

The blood concentrations of leukocytes and the major classes of leukocytes were compared across over 200 species of birds and over 300 species of mammals based on phylogenic relations, body weight and diet. The concentration of leukocytes was markedly higher in birds than mammals reflecting elevated concentrations of all major classes but particularly those of lymphocytes, monocytes, eosinophils and basophils. Overall, there were no differences between blood leukocyte concentrations in placental and marsupial mammals. There were differences between leukocytes concentrations within the placental mammals, for instance being lower in mammalian species within the taxa Glires than in Laurasiatheria and Primatomorpha which in turn are lower than in Afrotheria and Xenarthra. There were also differences in leukocyte concentrations in birds being higher in species with the taxa Afroaves than in Australaves. There were phylogenic differences in the percentages of the leukocyte classes. For instance, in placental compared marsupial mammals, the percentage neutrophils is elevated, the percentage lymphocytes is depressed and neutrophil to lymphocyte (N:L) ratio was much higher. Moreover, the percentage heterophils is high, the percentage lymphocytes is low and heterophil to lymphocyte (H:L) ratio is greater in birds from the taxa Paleognathae than the Neognathae. There was a positive relationship between concentrations of leukocytes or neutrophils and log body weight

Even though detailed anatomical studies of the otic region exist for some avian species, there are few comparative data for most extant higher-level clades. In the present study, the external osteology of the otic region of birds is studied in a broad taxonomic context and a focus has been put on the homology of neurovascular and muscular foramina in taxa of the Neognathae. It is detailed that this part of the skull offers significant structural variation, which is of potential phylogenetic and physiological significance. The otic region of many taxa is for the first time illustrated and some osteological features are highlighted that are not or only rarely commented on in existing accounts of the otic region. These include the foramina and osseous guiding structures for the occipital ramus of arteria ophthalmica externa, as well as the skeletal correlates of ramus palatinus and ramus hyomandibularis of nervus facialis, and the foramen for musculus columellae. In addition, different conformations of the pila otica are described and some structures of unknown significance are highlighted, which require further research.

Zygodactylidae are an extinct lineage of perching birds characterized by distinct morphologies of the foot and wing elements. Although the clade has a complex taxonomic history, current hypotheses place Zygodactylidae as the sister taxon to Passeriformes (i.e., songbirds). Given the rather sparse fossil record of early passeriforms, the description of zygodactylid taxa is important for inferring potentially ancestral states in the largest radiation of living birds (i.e., the ∼6,000 species of extant passeriforms). Despite the exceptional preservation of many specimens and considerable species diversity in Zygodactylidae, the relationships among species have not been previously evaluated in a phylogenetic context. Herein, we review the fossil record of Zygodactylidae from North America and describe five new well-preserved fossils from the early Eocene Green River Formation of Wyoming. Two specimens are identified as representing a new species and the first records of the taxonZygodactylusoutside Europe. Anatomical comparisons with previously named taxa and the results of phylogenetic analysis including newly described specimens and previously named zygodactylid taxa provide the first hypothesis of the species-level relationships among zygodactylids. The monophyly of Zygodactylidae is supported in these new analyses. However, the monophyly ofPrimozygodactylusand the taxonomic distinction betweenZygodactylusandEozygodactylusremain unresolved and would likely benefit from the description of additional specimens.

Birds represent the most diverse extant tetrapod clade, with ca. 10,000 extant species, and the timing of the crown avian radiation remains hotly debated. The fossil record supports a primarily Cenozoic radiation of crown birds, whereas molecular divergence dating analyses generally imply that this radiation was well underway during the Cretaceous. Furthermore, substantial differences have been noted between published divergence estimates. These have been variously attributed to clock model, calibration regime, and gene type. One underappreciated phenomenon is that disparity between fossil ages and molecular dates tends to be proportionally greater for shallower nodes in the avian Tree of Life. Here, we explore potential drivers of disparity in avian divergence dates through a set of analyses applying various calibration strategies and coding methods to a mitochondrial genome dataset and an 18-gene nuclear dataset, both sampled across 72 taxa. Our analyses support the occurrence of two deep divergences (i.e., the Palaeognathae/Neognathae split and the Galloanserae/Neoaves split) well within the Cretaceous, followed by a rapid radiation of Neoaves near the K-Pg boundary. However, 95% highest posterior density intervals for most basal divergences in Neoaves cross the boundary, and we emphasize that, barring unreasonably strict prior distributions, distinguishing between a rapid Early Paleocene radiation and a Late Cretaceous radiation may be beyond the resolving power of currently favored divergence dating methods. In contrast to recent observations for placental mammals, constraining all divergences within Neoaves to occur in the Cenozoic does not result in unreasonably high inferred substitution rates. Comparisons of nuclear DNA (nDNA) versus mitochondrial DNA (mtDNA) datasets and NT- versus RY-coded mitochondrial data reveal patterns of disparity that are consistent with substitution model misspecifications that result in tree compression/tree extension artifacts, which may explain some discordance between previous divergence estimates based on different sequence types. Comparisons of fully calibrated and nominally calibrated trees support a correlation between body mass and apparent dating error. Overall, our results are consistent with (but do not require) a Paleogene radiation for most major clades of crown birds.

The cranial openings of nervus hypoglossus, the 12th cranial nerve, are for the first time studied across a broad range of higher avian taxa. This nerve plays an important role in the innervation of the syrinx and exits the skull through a variable number of foramina. Most previous authors described 2–3 foramina nervi hypoglossi (FNH) for neornithine birds, but the number, size, and arrangement of FNH is actually more variable than what is apparent from the literature. In the case of three foramina, there is usually a pair of caudal foramina and a rostral one, but even in closely related taxa, a great variability of the FNH pattern may exist. Many taxa of Neognathae exhibit a quadruple of symmetrically arranged FNH, in others four foramina are strung together in a line. A few taxa show more than four FNH, although in these cases the additional foramina are very small. Of particular interest is the occurrence of a very large caudal FNH in Trochilidae and many species of Passeriformes. This large foramen is suggestive of a correlation with the highly developed vocal capabilities of these birds and may transmit fibers from the tracheosyringeal portion of nucleus nervi hypoglossi, in which case it would be an osteological correlate of vocal capabilities. However, targeted neuroanatomical studies are required to determine how individual hypoglossal foramina relate to hypoglossal roots and their branches, and which of them receive fibers supplying axial, lingual, and syringeal muscles.

Ostrich-like birds (Palaeognathae) show very little taxonomic diversity while their sister taxon (Neognathae) contains roughly 10,000 species. The main anatomical differences between the two taxa are in the crania. Palaeognaths lack an element in the bill called the lateral bar that is present in both ancestral theropods and modern neognaths, and have thin zones in the bones of the bill, and robust bony elements on the ventral surface of their crania. Here we use a combination of modeling and developmental experiments to investigate the processes that might have led to these differences. Engineering-based finite element analyses indicate that removing the lateral bars from a neognath increases mechanical stress in the upper bill and the ventral elements of the skull, regions that are either more robust or more flexible in palaeognaths. Surgically removing the lateral bar from neognath hatchlings led to similar changes. These results indicate that the lateral bar is load-bearing and suggest that this function was transferred to other bony elements when it was lost in palaeognaths. It is possible that the loss of the load-bearing lateral bar might have constrained diversification of skull morphology in palaeognaths and thus limited taxonomic diversity within the group.

International audience; To better determine the history of modern birds, we performed a genome-scale phylogenetic analysis of 48 species representing all orders of Neoaves using phylogenomic methods created to handle genome-scale data. We recovered a highly resolved tree that confirms previously controversial sister or close relationships. We identified the first divergence in Neoaves, two groups we named Passerea and Columbea, representing independent lineages of diverse and convergently evolved land and water bird species. Among Passerea, we infer the common ancestor of core landbirds to have been an apex predator and confirm independent gains of vocal learning. Among Columbea, we identify pigeons and flamingoes as belonging to sister clades. Even with whole genomes, some of the earliest branches in Neoaves proved challenging to resolve, which was best explained by massive protein-coding sequence convergence and high levels of incomplete lineage sorting that occurred during a rapid radiation after the Cretaceous-Paleogene mass extinction event about 66 million years ago.

Most systematic studies of the avian visual system have focused on Neognathous species, leaving virtually unexplored the Palaeognathae, comprised of the flightless ratites and the South American tinamous. We investigated the visual field, the retinal topography, and the pattern of retinal and centrifugal projections in the Chilean tinamou, a small Palaeognath of the family Tinamidae. The tinamou has a panoramic visual field with a small frontal binocular overlap of 20°. The retina possesses three distinct topographic specializations: a horizontal visual streak, a dorsotemporal area, and an area centralis with a shallow fovea. The maximum ganglion cell density is 61,900/ mm(2) , comparable to Falconiformes. This would provide a maximal visual acuity of 14.0 cycles/degree, in spite of relatively small eyes. The central retinal projections generally conform to the characteristic arrangement observed in Neognathae, with well-differentiated contralateral targets and very few ipsilateral fibers. The centrifugal visual system is composed of a considerable number of multipolar centrifugal neurons, resembling the "ectopic" neurons described in Neognathae. They form a diffuse nuclear structure, which may correspond to the ancestral condition shared with other sauropsids. A notable feature is the presence of terminals in deep tectal layers 11-13. These fibers may represent either a novel retinotectal pathway or collateral branches from centrifugal neurons projecting to the retina. Both types of connections have been described in chicken embryos. Our results widen the basis for comparative studies of the vertebrate visual system, stressing the conserved character of the visual projections' pattern within the avian clade.

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.

The study of avian eggshell structure, including composition, pigmentation, thickness, and strength,hasimportantecologicalandeconomicimplications.Previousinvestigatorshaveusedavarietyoftechniques to derive either direct measures or indirect estimates of eggshell thickness. Assessing the repeatability and method agreement of different techniques is necessary to permit comparison of eggshell thickness values from different studies on various genetic stocks, populations, and species. We recorded and analyzed measurements of eggshell thickness using two methods, micrometers and scanning electron microscopy (SEM), for several Palaeognathae and Neognathae taxa, including nonpasserines and passerines. Applying a tolerance-interval approach, we found that repeatability of measurements for eggs with thinner shells (

On the evolution of basicranial articulations of birds, especially of fowl and waterfowl (Galloanseres) Basicranial articulations are among the most neglected structures of the avian feeding apparatus. Based on the presented ontogenetic evidence two nonhomologous types of basicranial articulations are distinguished in birds: 1. A basipterygoid articulation developing from an early embryonic quadratopolar comissure, which is homologous to the basipterygoid articulation of other amniotes. This diarthrosis is present in all paleognathous as well as in many juvenile and some adult neognathous birds. 2. A rostropterygoid articulation developing through apposition of the pterygoid on the parasphenoid. This secondary articulation is known from galliform and ansenform birds (Galloanseres). Based on ontogenetic evidence as well as functional analysis following hypothesis explaining the evolution of rostropterygoid articulations in the Galloanseres is proposed: 1. In the groundplan of birds (Aves, sensu Gauthier 1986) a basipterygoid articulation was present continuously during ontogeny. It protected the pterygoid-quadrate articulation against propalineal distortion by controlling the movement of the caudal end of the flexible, but undivided pterygopalatinal bar. This construction is preserved in recent paleognaths. 2. In the groundplan of the Neognathae an intrapterygoid joint subdividing the pterygopalatinal bar was formed in juveniles. Consequently the basipterygoid articulation had lost its function in adults and was therefore developed only in embryonic and juvenile stages. This condition is present in many recent neognaths. 3. In the stemline of the Galloanseres the intrapterygoid joint was already formed in embryos. Since an undivided pterygopalatinal bar is absent in any ontogenetic stage of the Galloanseres, no basipterygoid articulation is being formed. The recent Galloanseres retain this developmental pattern. 4. In the groundplan of the Galloanseres a newly developed rostropterygoid articulation was present providing a support for the pterygopalatinal bar. This articulation could withstand not only pressure but also increasing tension occurring as a consequence of the development of a gliding jaw joint and a firm grip with beak tips, both consequences of increasing herbivory in the Galloanseres. Since the basipterygoid articulation was reduced, a new secondary joint had to be developed.

Deep avian evolutionary relationships have been difficult to resolve as a result of a putative explosive radiation. Our study examined ∼32 kilobases of aligned nuclear DNA sequences from 19 independent loci for 169 species, representing all major extant groups, and recovered a robust phylogeny from a genome-wide signal supported by multiple analytical methods. We documented well-supported, previously unrecognized interordinal relationships (such as a sister relationship between passerines and parrots) and corroborated previously contentious groupings (such as flamingos and grebes). Our conclusions challenge current classifications and alter our understanding of trait evolution; for example, some diurnal birds evolved from nocturnal ancestors. Our results provide a valuable resource for phylogenetic and comparative studies in birds.

Although reconstruction of the phylogeny of living birds has progressed tremendously in the last decade, the evolutionary history of Neoaves--a clade that encompasses nearly all living bird species--remains the greatest unresolved challenge in dinosaur systematics. Here we investigate avian phylogeny with an unprecedented scale of data: >390,000 bases of genomic sequence data from each of 198 species of living birds, representing all major avian lineages, and two crocodilian outgroups. Sequence data were collected using anchored hybrid enrichment, yielding 259 nuclear loci with an average length of 1,523 bases for a total data set of over 7.8 × 10(7) bases. Bayesian and maximum likelihood analyses yielded highly supported and nearly identical phylogenetic trees for all major avian lineages. Five major clades form successive sister groups to the rest of Neoaves: (1) a clade including nightjars, other caprimulgiforms, swifts, and hummingbirds; (2) a clade uniting cuckoos, bustards, and turacos with pigeons, mesites, and sandgrouse; (3) cranes and their relatives; (4) a comprehensive waterbird clade, including all diving, wading, and shorebirds; and (5) a comprehensive landbird clade with the enigmatic hoatzin (Opisthocomus hoazin) as the sister group to the rest. Neither of the two main, recently proposed Neoavian clades--Columbea and Passerea--were supported as monophyletic. The results of our divergence time analyses are congruent with the palaeontological record, supporting a major radiation of crown birds in the wake of the Cretaceous-Palaeogene (K-Pg) mass extinction.

The study of the contributions of different bones to the formation of the skeleton in birds is necessary: (1) to establish homologies in comparative anatomy; (2) to delimit each bone structure correctly, mainly in relation to the skull and mandible where the bones are fused to each other in adults; and (3) to standardize nomenclature in avian osteology. In this paper at least one young specimen belonging to each sub-family of Cuculidae was examined in order to identify each bone in terms of boundaries and contributions to skull and mandible formation. These cuckoos specimens were also compared with adults and young of turacos and hoatzin. The results show little variation of skull and jaw among the young cuckoos studied compared with the variations among adult specimens. However, it provides new suggestions for the boundaries and nomenclature of certain osseous structures in the skull and mandible of birds, specifically fissura zona flexoria craniofacialis, prominetia frontoparietalis, crista temporalis transversa, processus squamosalis, fossa laterosphenoidalis, tuberculum laterosphenoidale and processus retroangularis. This study also provides more reliable homologies for use in cladistic analysis and above all it contributes to the phylogenetic position of Cuculidae within Neognathae, specifically the skull formation suggest that turacos and hoatzin are more similar to each other than either is to the cuckoos.

Patterns of diversification and timing of evolution within Neoaves, which includes almost 95% of all bird species, are virtually unknown. On the other hand, molecular data consistently indicate a Cretaceous origin of many neoavian lineages and the fossil record seems to support an Early Tertiary diversification. Here, we present the first well-resolved molecular phylogeny for Neoaves, together with divergence time estimates calibrated with a large number of stratigraphically and phylogenetically well-documented fossils. Our study defines several well-supported clades within Neoaves. The calibration results suggest that Neoaves, after an initial split from Galloanseres in Mid-Cretaceous, diversified around or soon after the K/T boundary. Our results thus do not contradict palaeontological data and show that there is no solid molecular evidence for an extensive pre-Tertiary radiation of Neoaves.

This oological analysis supports Palaeognathae monophyly, and places Dromornithidae within Neognathae, the sister clade of Paleognathae. This Dromornithidae relationship is congruent with a recent phylogenetic hypothesis, based on cranial characters, that differs from the original assignment of this group of extinct birds to the Palaeognathae. Minor differences exist between previous palaeognath phylogenetic hypotheses and that obtained from this research. Some of the topological disparities might be influenced solely by inclusion of more fossil taxa in this analysis compared with those considering either molecular or skeletal characters. Surprisingly, oological data differentiate the Australasian palaeognath taxa from those of Africa and South America; this phylogenetic topology conforms to the paleognath geographic distribution. Regardless of the issues related to Paleognathae, results show that oological characters permit the differentiation of groups at a generic level, thus validating the use of eggs...

The tinamid bird Nothura maculosa, along with other species of the order Tinamiformes and all of the existent ratites, form the infraclass Paleognathae, the most primitive living birds. Previous work has shown that in all studied Neognathae, the ZW pair shows strictly localized recombination in a very short pseudoautosomal region, while in paleognath birds, the ZW pairs have mostly free recombination. The present observations show that the ZW pair of N. maculosa has a recombination pattern departing from both neognaths and other Paleognath birds, as there is a single crossover but occurring at random points along a significant part of the long arm of the W chromosome. This recombination pattern agrees with the presence of intercalary and terminal heterochromatin in the W chromosome, suggesting an exceptional, additional step of recombination suppression.Key words: recombination, ZW pair, avian sex chromosomes, sex chromosome heteromorphism.

SUMMARYCranial kinesis is an important feature in avian feeding behaviour and involves the transmission of quadrate movement to the upper bill by the Pterygoid–Palatinum Complex (PPC). The PPC in Palaeognathae is remarkably different from that found in Neognathae. In this study we analyse whether the special morphology of the PPC is an adaptation to the feeding behaviour of the Palaeognathae. Behavioural analyses of the rhea Rhea americana showed that the feeding behaviour of the rhea is typical `Catch and Throw' behaviour, independent of the size of the food item. Drinking is achieved by a scooping movement followed by a low-amplitude tip-up phase. During feeding rhynchokinetic movements of the upper bill were observed. However, cranial kinesis was limited and may differ from rhynchokinesis in neognathes as a clear bending zone seemed absent. Since the movement patterns are considered very similar to the basic feeding behaviour in neognathous birds it is concluded that the specific morphology of the PPC is not the result of specific functional demands from palaeognathous feeding behaviour.

The coding region of the preproinsulin gene has been cloned and partly sequenced in a variety of marine and terrestrial birds (28 species). All genes showed the “ancestral” structure with a large intron-2. The size of intron-2 changed considerably during the evolution of birds (2.4–4.2 kb). The hydrophobicity of signal peptides was conserved. Bird C-peptides were predicted to be 28 amino acids long, but circulating C-peptides would be only 26 amino acids long, with Passer as a possible exception. Bird C-peptides were found to lack the sequences identified in mammals as responsible for peptide bioactivity and the structure of the central part. In contrast, predicted insulin sequences were highly conserved. Only two types of analog were identified: the hypoactive form (GluA8), present only in Anseriformes and the hyperactive form (His A8), present in all other species. Based on 3′-nucleotide sequence analysis (extending into intron-2), birds appeared to be monophyletic. Five groups were clearly identified: Paleognathae, Galliformes, Anseriformes, Passeriformes, and Charadriiformes. Paleognathae were suggested as the basal group, supporting the traditional view of avian evolution. Subsequent branching identified a gallo-anserae group and a group containing all other Neognathae. Surprisingly, Columba livia (Columbiforme order) clustered with Galliformes. With represented species, Procellariiformes and possibly Ciconiiformes, and Pelicaniformes were suggested as paraphyletic, in agreement with conclusions from some studies based on mitochondrial DNA sequences.

The apparently rapid and ancient diversification of many avian orders complicates the resolution of their relationships using molecular data. Recent studies based on complete mitochondrial DNA (mtDNA) sequences or shorter lengths of nuclear sequence have helped corroborate the basic structure of the avian tree (e.g., a basal split between Paleognathae and Neognathae) but have made relatively little progress in resolving relationships among the many orders within Neoaves. We explored the potential of a moderately sized mtDNA data set ( approximately 5000 bp for each of 41 taxa), supplemented with data from a nuclear intron ( approximately 700 bp per taxon), to resolve relationships among avian orders. Our sampling of taxa addresses two issues: (1). the sister relationship and monophyly, respectively, of Anseriformes and Galliformes and (2). relationships of the enigmatic hoatzin Opisthocomus hoazin. Our analyses support a basal split between Galloanserae and Neoaves within Neognathae and monophyly of both Galliformes and Anseriformes. Within Galliformes, megapodes and then cracids branch basally. Within Anseriformes, mitochondrial data support a screamer (Anhimidae) plus magpie goose (Anseranatidae) clade. This result, however, may be an artifact of divergent base composition in one of the two anatids we sampled. With deletion of the latter taxon, Anseranas is sister to anatids as in traditional arrangements and recent morphological studies. Although our data provide limited resolution of relationships within Neoaves, we find no support for a sister relationship between either cuckoos (Cuculiformes) or turacos (Musophagiformes) and hoatzin. Both mitochondrial and nuclear data are consistent with a relationship between hoatzin and doves (Columbiformes), although this result is weakly supported. We also show that mtDNA sequences reported in another recent study included pervasive errors that biased the analysis towards finding a sister relationship between hoatzin and turacos.

Description of well-preserved fossil eggshell (oospecies Prismatoolithus levis), recently assigned to the theropod dinosaur Troodon formosus, reveals traits that are shared with the eggshell of both fossil and Recent avians. Bird-like characteristics of troodontid eggshell include: fibres associated with eisospherites that are attached to the bases of the mammillae, fine radiating crystals that form the spherulites and grade into blocky wedges of the mammillae, and prisms in the outer layer that exhibit squamatic ultrastructure. The presence of two layers (mammillary and squamatic) within the eggshell is either a theropod apomorphy or it arose within Theropoda; the external eggshell layer of crown-group avians is apomorphic with respect to other theropod eggshell. Despite disagreement on the putative sister taxon of birds, recent theropod phylogenies suggest strongly that prismatic ultrastructure evolved independently in Troodontidae and Neognathae. Our identification of squamatic ultrastructure in troodontid eggshell, which was previously thought to lack this structure, reveals problems with the previous assignment of this eggshell to the dinosauroid-prismatic morphotype and with the concept of eggshell morphotype in the current eggshell parataxonomy. We conclude that the dinosauroid-prismatic morphotype should be abandoned.

Several different types of cranial kinesis are present within modern birds, enabling them to move (part of) the upper bill relative to the braincase. This movement of the upper bill results from movement of the quadrate and the pterygoid-palatine complex (PPC). The taxon Palaeognathae is characterised by a very distinct PPC and a special type of cranial kinesis (central kinesis) that is very different from that found in the Neognathae. This has led some authors to hypothesise that there is a functional relationship between the morphology of the PPC and the type of cranial kinesis. This hypothesis is tested here by analysing the movement pattern of both the upper bill and the PPC in birds with three different types of cranial kinesis: prokinesis, distal rhynchokinesis and central rhynchokinesis. Movement patterns were determined using a Roentgen stereophotogrammetry method, which made it possible to detect very small displacements (0.5 mm) of bony elements in three dimensions, while the jaw muscles and ligaments remained intact. We found that in all types of kinesis investigated the movements of the quadrate, jugal bars and PPC are similar. Movement of the quadrate is transferred to the upper beak by the jugal bar and the PPC, which moves almost exclusively forwards and backwards, thereby elevating or depressing the upper bill. The differences between the types of kinesis lie only in the position of the point of rotation. These findings indicate that there is no correlation between the specific morphology of the PPC and the type of cranial kinesis. Several other factors, including the external forces applied during food acquisition, may influence the morphology of the PPC. Differences in PPC morphology therefore appear to be the result of different functional demands acting on the system simultaneously but with different strengths, depending on the species.

Several different types of cranial kinesis are present within modern birds, enabling them to move (part of) the upper bill relative to the braincase. This movement of the upper bill results from movement of the quadrate and the pterygoid–palatine complex (PPC). The taxon Palaeognathae is characterised by a very distinct PPC and a special type of cranial kinesis (central kinesis) that is very different from that found in the Neognathae. This has led some authors to hypothesise that there is a functional relationship between the morphology of the PPC and the type of cranial kinesis. This hypothesis is tested here by analysing the movement pattern of both the upper bill and the PPC in birds with three different types of cranial kinesis: prokinesis, distal rhynchokinesis and central rhynchokinesis. Movement patterns were determined using a Roentgen stereophotogrammetry method, which made it possible to detect very small displacements (0.5 mm) of bony elements in three dimensions, while the jaw muscles and ligaments remained intact. We found that in all types of kinesis investigated the movements of the quadrate, jugal bars and PPC are similar. Movement of the quadrate is transferred to the upper beak by the jugal bar and the PPC, which moves almost exclusively forwards and backwards, thereby elevating or depressing the upper bill. The differences between the types of kinesis lie only in the position of the point of rotation. These findings indicate that there is no correlation between the specific morphology of the PPC and the type of cranial kinesis. Several other factors, including the external forces applied during food acquisition, may influence the morphology of the PPC. Differences in PPC morphology therefore appear to be the result of different functional demands acting on the system simultaneously but with different strengths, depending on the species.

Molecular analyses show that modern birds can be divided into two major taxa, the Paleognathae and the Neognathae. This division was already proposed by Merrem in 1813, based on morphological characters. One of the most prominent discriminating characters is the morphology of the Pterygoid-Palatinum Complex (PPC), which is different in paleognathous and neognathous birds. There are very few other morphological characters that support this division and even the differences in PPC have been under dispute. A discriminant analysis based on quantitative measurements of the PPC shows that a large difference between the two morphologies exists, and that the Tinamidae posses an intermediate form. An evolutionary maximum-likelihood analysis suggests that the PPC of the Paleognathae is more primitive than that of the Neognathae. A functional interpretation of the differences in the PPC between the Paleognathae and the Neognathae indicates that the paleognathous PPC is not, as generally accepted, an adaptation related to rhynchokinesis, but probably contributes to reinforcement of the skull after the loss of both the postorbital and nasal bar.

The complete mitochondrial DNA, mtDNA, molecule of the greater rhea, Rhea americana, was sequenced. The size of the molecule is 16,710 nucleotides. The organization of the molecule conforms with that described for the chicken and the ostrich. It has been shown previously that relative to other vertebrates the NADH3 gene of the ostrich has an insertion of one nucleotide in position 174 of the gene. The same observation was made in the rhea and in the newly sequenced NADH3 gene of the emu, Dromaius novaehollandiae. Comparison with the NADH3 gene of the chicken and the rook suggests that the inserted nucleotide may be deleted by RNA editing. The divergence between the two struthioniform species, the ostrich and the rhea, was molecularly dated at approximately 51 million years before present, MYBP. This dating is more recent than commonly acknowledged. Phylogenetic analysis of the complete cytochrome b genes of seven avian orders placed the Passeriformes basal in the avian tree with the Struthioniformes among the remaining Neognathae. These findings challenge the commonly accepted notion that the most basal avian divergence is that between the Palaeognathae and Neognathae.

Background Secondary edentulism (toothlessness) has evolved on multiple occasions in amniotes including several mammalian lineages (pangolins, anteaters, baleen whales), birds, and turtles. All edentulous amniote clades have evolved from ancestors with enamel-capped teeth. Previous studies have documented the molecular decay of tooth-specific genes in edentulous mammals, all of which lost their teeth in the Cenozoic, and birds, which lost their teeth in the Cretaceous. By contrast with mammals and birds, tooth loss in turtles occurred in the Jurassic (201.6-145.5 Ma), providing an extended time window for tooth gene degradation in this clade. The release of the painted turtle and Chinese softshell turtle genomes provides an opportunity to recover the decayed remains of tooth-specific genes in Testudines. Results We queried available genomes of Testudines (Chrysemys picta [painted turtle], Pelodiscus sinensis [Chinese softshell turtle]), Aves (Anas platyrhynchos [duck], Gallus gallus [chicken], Meleagris gallopavo [turkey], Melopsittacus undulatus [budgerigar], Taeniopygia guttata [zebra finch]), and enamelless mammals (Orycteropus afer [aardvark], Choloepus hoffmanni [Hoffmann’s two-toed sloth], Dasypus novemcinctus [nine-banded armadillo]) for remnants of three enamel matrix protein (EMP) genes with putative enamel-specific functions. Remnants of the AMBN and ENAM genes were recovered in Chrysemys and retain their original synteny. Remnants of AMEL were recovered in both testudines, although there are no shared frameshifts. We also show that there are inactivated copies of AMBN, AMEL and ENAM in representatives of divergent avian lineages including Galloanserae, Passeriformes, and Psittaciformes, and that there are shared frameshift mutations in all three genes that predate the basal split in Neognathae. Among enamelless mammals, all three EMP genes exhibit inactivating mutations in Orycteropus and Choloepus. Conclusions Our results highlight the power of combining fossil and genomic evidence to decipher macroevolutionary transitions and characterize the functional range of different loci involved in tooth development. The fossil record and phylogenetics combine to predict the occurrence of molecular fossils of tooth-specific genes in the genomes of edentulous amniotes, and in every case these molecular fossils have been discovered. The widespread occurrence of EMP pseudogenes in turtles, birds, and edentulous/enamelless mammals also provides compelling evidence that in amniotes, the only unique, non-redundant function of these genes is in enamel formation.

In birds, there is a retinofugal projection from the brain to the retina originating from the isthmo optic nucleus (ION) in the midbrain. Despite a large number of anatomical, physiological and histochemical studies, the function of this retinofugal system remains unclear. Several functions have been proposed including: gaze stabilization, pecking behavior, dark adaptation, shifting attention, and detection of aerial predators. This nucleus varies in size and organization among some species, but the relative size and morphology of the ION has not been systematically studied. Here, we present a comparison of the relative size and morphology of the ION in 81 species of birds, representing 17 different orders. Our results show that several orders of birds, besides those previously reported, have a large, well-organized ION, including: hummingbirds, woodpeckers, coots and allies, and kingfishers. At the other end of the spectrum, parrots, herons, waterfowl, owls and diurnal raptors have relatively small ION volumes. ION also appears to be absent or unrecognizable is several taxa, including one of the basal avian groups, the tinamous, which suggests that the ION may have evolved only in the more modern group of birds, Neognathae. Finally, we demonstrate that evolutionary changes in the relative size and the cytoarchitectonic organization of ION have occurred largely independent of phylogeny. The large relative size of the ION in orders with very different lifestyles and feeding behaviors suggest there is no clear association with pecking behavior or predator detection. Instead, our results suggest that the ION is more complex and enlarged in birds that have eyes that are emmetropic in some parts of the visual field and myopic in others. We therefore posit that the ION is involved in switching attention between two parts of the retina i.e. from an emmetropic to a myopic part of the retina.

Tinamiphilopsis ariconte n. sp., a syringophilid mite, is described from three tinamou species (Tinamidae), Rhynchotus rufescens (Temminck) and Nothura boraquira (Spix) in Paraguay and N. minor (Spix) in Brazil. These data indirectly support the hypothesis that the cheyletoid-like predatory ancestor of the Syringophilidae switched to parasitism before the divergence of the avian hosts into the two major clades Palaeognathae and Neognathae.

The vast majority of extant birds possess highly differentiated Z and W sex chromosomes. Nucleotide sequence data from gametologs (homologs on opposite sex chromosomes) suggest that this divergence occurred throughout early bird evolution via stepwise cessation of recombination between identical sex chromosomal regions. Here, we investigated avian sex chromosome differentiation from a novel perspective, using retroposon insertions and random insertions/deletions for the reconstruction of gametologous gene trees. Our data confirm that the CHD1Z/CHD1W genes differentiated in the ancestor of the neognaths, whereas the NIPBLZ/NIPBLW genes diverged in the neoavian ancestor and independently within Galloanserae. The divergence of the ATP5A1Z/ATP5A1W genes in galloanserans occurred independently in the chicken, the screamer, and the ancestor of duck-related birds. In Neoaves, this gene pair differentiated in each of the six sampled representatives, respectively. Additionally, three of our investigated loci can be utilized as universal, easy-to-use independent tools for molecular sexing of Neoaves or Neognathae.

Publisher Summary This chapter examines the skull and mandible of the pigeon, Columba livia that belongs to Aves, or birds. Aves are characterized by complete loss of teeth and a very large sternal keel. They are subdivided into Palaeognathae and Neognathae, which are differen­tiated primarily on the basis of palatal features. The paleognaths include two lineages—the flightless ratites and the tinamous. The structure of birds has been highly modified to meet the demands of flight. The skel­eton has a number of such modifications, mostly in the forelimbs, which are adapted to form a bony frame for the wings. Also, however, the skeleton has undergone changes to reduce weight, clearly it is an advantage in a flying animal, and mobility, which gives the wings a rigid support. Several bones of many birds are pneumatic—hollow and con­taining extensions of the respiratory system's air sacs. In females only the left-side reproduc­tive organs are present, those of the right side having degenerated soon after their initial formation. The ovary lies on the anterior end of the left kidney. Its morphology and size varies in accordance with the breeding season.