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Our understanding of the early evolution of birds has advanced over the past 2 decades, thanks to an ever-improving fossil record. Extraordinary fossils have revealed new details about the evolution of the avian brain, respiratory system, digestive tract, and reproductive system. Many of the traits most strongly associated with birds first arose in nonavian theropod dinosaurs. Theropods evolved pennaceous feathers, incipient wings, and gliding flight long before modern birds appeared. Birds likewise inherited features such as an expanded forebrain, gizzard, dorsally immobile lung, pigmented eggs, and paternal brooding system from their theropod ancestors. Yet, the earliest birds also retained primitive traits such as teeth, clawed hands, long bony tails, partially buried nests, and slower growth. The evolution of birds was profoundly influence by the Cretaceous–Paleogene mass extinction, which wiped out the previously dominant Enantiornithines (“opposite birds”). This sets the stage for modern birds to radiate into the most diverse major clade of tetrapods.

Adaptations to different diets represent a hallmark of animal diversity. The diets of birds are highly variable, making them an excellent model system for studying adaptive evolution driven by dietary changes. To test whether molecular adaptations to diet have occurred during the evolution of birds, we examined a dietary enzyme alanine-glyoxylate aminotransferase (AGT), which tends to target mitochondria in carnivorous mammals, peroxisomes in herbivorous mammals, and both mitochondria and peroxisomes in omnivorous mammals. A total of 31 bird species were examined in this study, which included representatives of most major avian lineages. Of these, 29 have an intact mitochondrial targeting sequence (MTS) of AGT. This finding is in stark contrast to mammals, which showed a number of independent losses of the MTS. Our cell-based functional assays revealed that the efficiency of AGT mitochondrial targeting was greatly reduced in unrelated lineages of granivorous birds, yet it tended to be high in insectivorous and carnivorous lineages. Furthermore, we found that proportions of animal tissue in avian diets were positively correlated with mitochondrial targeting efficiencies that were experimentally determined, but not with those that were computationally predicted. Adaptive evolution of AGT mitochondrial targeting in birds was further supported by the detection of positive selection on MTS regions. Our study contributes to the understanding of how diet drives molecular adaptations in animals, and suggests that caution must be taken when computationally predicting protein subcellular targeting.

Flight is a key feature in the evolution of birds. Wing anatomy reflects many aspects of avian biology such as flight ability. However, our knowledge of the flight musculature has many gaps still, particularly for the distal wing. Therefore, the aim of this work was to investigate the form-function relationship of the forelimb myology of birds to understand the role of individual muscles during flight. Dissections of six species of birds of prey were performed to collect numerical data of muscle architecture, which is the primary determinant of muscle function and force-generation capacity. Birds of prey are a highly diverse group that presents different flight styles throughout the taxa, making them a good model for our purposes. Wing muscle mass (MM) isometrically scaled with body mass1.035 , muscle length to MM0.343 , and fascicle length (FL) scaled allometrically to MM0.285 . The shoulder musculature scaled differently than the other regions where the FL increases more slowly than would be expected in geometrically similar animals, which affects flight mechanics. A proximal-to-distal reduction of MM occurs, which helps to minimize the wing moment of inertia during flight while allowing precise control of the distal wing. Interestingly, the distribution of MM appeared to be species-specific, suggesting a functional signal. This study provides numerical information of muscle architecture of the avian wing that helps us to understand muscle function and its implication in flight, and can be used in future studies of flight mechanics. Anat Rec, 302:1808-1823, 2019. © 2019 American Association for Anatomy.

Since the sequencing of the zebra finch genome it has become clear the avian genome, while largely stable in terms of chromosome number and gene synteny, is more dynamic at an intrachromosomal level. A multitude of intrachromosomal rearrangements and significant variation in transposable element content have been noted across the avian tree. Transposable elements (TEs) are a source of genome plasticity, because their high similarity enables chromosomal rearrangements through non-allelic homologous recombination, and they have potential for exaptation as regulatory and coding sequences. Previous studies have investigated the activity of the dominant TE in birds, CR1 retrotransposons, either focusing on their expansion within single orders, or comparing passerines to non-passerines. Here we comprehensively investigate and compare the activity of CR1 expansion across orders of birds, finding levels of CR1 activity vary significantly both between and with orders. We describe high levels of TE expansion in genera which have speciated in the last 10 million years including kiwis, geese and Amazon parrots; low levels of TE expansion in songbirds across their diversification, and near inactivity of TEs in the cassowary and emu for millions of years. CR1s have remained active over long periods of time across most orders of neognaths, with activity at any one time dominated by one or two families of CR1s. Our findings of higher TE activity in species-rich clades and dominant families of TEs within lineages mirror past findings in mammals.Author SummaryTransposable elements (TEs) are mobile, self replicating DNA sequences within a species’ genome, and are ubiquitous sources of mutation. The dominant group of TEs within birds are chicken repeat 1 (CR1) retrotransposons, making up 7-10% of the typical avian genome. Because past research has examined the recent inactivity of CR1s within model birds such as the chicken and the zebra finch, this has fostered an erroneous view that all birds have low or no TE activity on recent timescales. Our analysis of numerous high quality avian genomes across multiple orders identified both similarities and significant differences in how CR1s expanded. Our results challenge the established view that TEs in birds are largely inactive and instead suggest that their variation in recent activity may contribute to lineage-specific changes in genome structure. Many of the patterns we identify in birds have previously been seen in mammals, highlighting parallels between the evolution of birds and mammals.

This chapter explores the evolutionary history of birds. It considers the dinosaur origins of birds and the evolution of the modern bird lineages. The development of the modern bird from its prehistoric, reptilian ancestors is analysed by discussion of important fossil specimens, particularly that of Archaeopteryx, and the development of both morphological and biomolecular phylogenies. Evolution by natural selection is explained, as are processes of evolutionary adaptation and speciation. The conservation implications of hybridization are considered and the classification and nomenclature of birds is introduced. Throughout the chapter examples of current research are presented alongside established classic studies to engage the reader and provide a route into the relevant scientific literature.

Evolution of birds from non-flying theropod dinosaurs is a classic evolutionary transition, but a deeper understanding of early flight has been frustrated by disagreement on the relationships between birds (Avialae) and their closest theropod relatives. We address this through a larger, more resolved evolutionary hypothesis produced by a novel automated analysis pipeline tailored for large morphological datasets. We corroborate the grouping of dromaeosaurids + troodontids (Deinonychosauria) as the sister taxon to birds (Paraves), as well as the recovery of Anchiornithidae as basalmost avialans. Using these phylogenetic results and available data for vaned feathered paravians, maximum and minimum estimates of wing loading and specific lift calculated using ancestral state reconstruction analysis are used as proxies for the potential for powered flight through this transition. We found a broad range of paravian ancestors with estimates approaching values that are indicative of powered flight potential. This suggests that prior to the evolution of flight there was a wider extent of experimentation with wing-assisted locomotion among paravians than previously appreciated. We recovered wing loading and specific lift estimates indicating the potential for powered flight among fossil birds as well as unenlagiine and microraptorine dromaeosaurids. In the context of our phylogeny and of Mesozoic palaeogeography, our results suggest that the potential for powered flight originated three or more times from a broad range of ancestors already nearing this potential, providing a well-supported scenario for the origin of theropod flight to further explore.

Two decades of paleontological discoveries of basal birds and non-avian theropods with preserved integumentary structures, especially in Late Jurassic to Early Cretaceous deposits from northeastern China, have greatly improved our understanding of the origin and early evolution of birds and their plumage. Here, we present a concise review of the plumage evolution within pennaraptora, the most inclusive clade containing Oviraptorosauria and Paraves. Feather or feather-like morphotypes were particularly diversified in non-avialan paravians, suggesting that they probably already fulfilled a wide array of biological roles, including thermoregulation and visual display. The feather-like structures in non-eumaniraptoran paravians were obviously not adapted for flight. However, Microraptor and maybe some of its relatives preserve large pennaceous feathers along the limbs and tail, similar in morphology and organization to those in modern birds, so that they could have functioned in active flight or passive gliding. Several aerodynamic innovations and flight-related morphological adaptations were (likely independently) experimented within the paravian clade close to the origin of birds. The origin and early evolution of complex feathers and flight abilities in paravian theropods were not linear processes, but more complex than previously thought.

The acoustic adaptation hypothesis (AAH) proposes that habitat characteristics affect the evolution of birds’ long-distance vocalizations. Although many studies have confirmed the AAH, some...

Ugolyakia kaluginae gen. et sp. nov. is described from Ugolyak (Santonian Taimyr amber), based primarily on its unbranched Rs and developed katepisternal sulcus. It is attributed to the tribe Simuliini, although it lacks calcipala and spiniform seta on the costal vein characteristic of most genera of the tribe. Possession of a claw with a large subbasal tooth and absence of significant sclerotization of the sternites suggest that U. kaluginae females were blood-sucking avian parasites. Black flies make up 3% of all insect inclusions and 5% of all identifiable dipterans in Ugolyak amber. Only two Late Cretaceous black fly specimens were previously known: a poorly preserved female from Yantardakh (Santonian Taimyr amber) and a complete one from Turonian New Jersey amber. Feathers found at nearly all Cretaceous black fly sites (and at all formations with records of Simuliini) were younger than the Hauterivian. Ugolyak black flies are thought to have inhabited the same environments as Cretaceous ornithurine birds and most likely fed on them. These insects can then be used as an indicator of this bird community, allowing a better understanding of the Late Cretaceous forest ecology of Northern Asia. The inferred presence of Ornithuromorpha at high latitudes by the Early Cretaceous implies that their high growth rate may have evolved as an adaptation to a short yearly period of productivity (probably as a compensation for the poor flight ability of their young). This further implies that advanced ornithuromorphs might have originated at higher latitudes; later, aquatic ornithuromorphs occupied niches in lower latitude regions with tropical climates such as the Chinese Jehol biota, to which they were preadapted. The inferred seasonality at higher latitudes during cold spells of the Early Cretaceous could further be viewed as a prerequisite for the evolutionary origin of the granivory.

‘Exceptional fossils’ of dinosaurs preserving feathers have radically changed the way we view their paleobiology and the evolution of birds. Understanding how such soft tissues preserve is imperative to accurately interpreting the morphology of fossil feathers. Experimental taphonomy has been integral to such investigations. One such experiment used a printing press to mimic compaction, done subaerially and without sediment burial, and concluded that the leaking of bodily fluid could lead to the clumping of feathers by causing barbs to stick together such that they superficially resemble simpler, less derived, filamentous structures. Here we use a novel, custom-built experimental setup to more accurately mimic subaqueous burial and compaction under low-energy, fine-grain depositional environments applicable to the taphonomic settings most plumage-preserving ‘exceptional fossils’ are found in. We find that when submerged and subsequently buried and compacted, feathers do not clump together and they maintain their original arrangement. Submersion in fluid in and of itself does not lead to clumping of barbs; this would only occur upon pulling feathers out from water into air. Furthermore, sediment encases the feathers, fixing them in place during compaction. Thus, feather clumping that leads to erroneously plesiomorphic morphological interpretations may not be a taphonomic factor of concern when examining fossil feathers. Our current methodology is amenable to further improvements that will continue to more accurately mimic subaqueous burial and compaction, allowing for various hypothesis testing. Electronic supplementary material The online version of this article (10.1007/s12542-018-0411-y) contains supplementary material, which is available to authorized users.

The Jehol Biota is a world-famous Early Cretaceous terrestrial ecosystem in East Asia and has revealed exceptionally well–preserved fossils which provide significant insights into the origin and evolution of birds and feathers, and the early diversifications of mammals and angiosperms. After the long-lived controversies over the timing of the biota varying from Late Jurassic to Early Cretaceous in last century, the lifespan of the early-middle Jehol Biota has been precisely dated from Barremian to early Aptian (ca. 131–120 Ma) by a series of recent radiometric analyses. However, few well–constrained ages are available for the late Jehol Biota, hindering our complete understanding of the evolutionary history of the terrestrial ecosystem. Alternating marine and non-marine deposits of the Chengzihe Formation in eastern Heilongjiang have yielded a non-marine bivalve assemblage of the Jehol Biota, basal angiosperms, and Cretaceous marine index bivalves, which collectively offer crucial and indisputable clues against the prevailing Jurassic time of the biota since 1990s. Here we present the first discovery of tephra layers from the Chengzihe Formation, dated by zircon U-Pb geochronology as ca. 116–111 Ma, providing the first quantitative age calibration for the early angiosperms and the Jehol bivalve assemblage. Therefore, we demonstrate that the duration of the Jehol Biota extends to late Aptian–early Albian, approximately 4–9 Myr younger than the currently accepted age limits.

New data on the taxonomic and morphological diversity of Early Cretaceous Enantiornithes are reviewed. A new hypothesis concerning the phylogenetic position of Pengornithidae is proposed. These birds traditionally treated as primitive enantiornithines may in fact be more closely related to Ornithuromorpha. This phylogenetic placement implies that the fan-shaped tail and modern-type humeral joint was formed once in the early evolution of birds. Hence, the similarity between Pengornithidae and other enantiornithines may be plesiomorphic. The ecology of Early Cretaceous enantiornithines is discussed. The increased mobility of neck in Holbotia is possibly accounted for by the cranioinertial swallowing mechanism, as in modern ratites. The hypotheses of scansorial adaptations in Parapengornis and Fortunguavis are criticized. In addition, the phylogenetic position of Mystiornis is discussed.

The evolution of birds from dinosaurs is a subject that has received great attention among vertebrate paleontologists. Nevertheless, the early evolution of the paravians, the group that contains birds and their closest non-avian dinosaur relatives, remains very poorly known. Even the most basal members of one paravian lineage, the Dromaeosauridae, already show a body plan that differs substantially from their closest non-paravian relatives. Recently, the dromaeosaurid Halszkaraptor escuilliei was described from the Cretaceous of Mongolia. Halszkaraptor possesses numerous unserrated premaxillary teeth, a platyrostral rostrum with a developed neurovascular system, an elongate neck, bizarrely-proportioned forearms, and a foreword-shifted center of mass, differing markedly from other paravians. A reevaluation of the anatomy, taphonomy, environmental setting, and phylogenetic position of H. escuilliei based on additional comparisons with other maniraptorans suggests that, rather than indicating it was a semiaquatic piscivore, the body plan of this dinosaur bears features widely distributed among maniraptorans and in some cases intermediate between the conditions in dromaeosaurids and related clades. I find no evidence for a semiaquatic lifestyle in Halszkaraptor. A phylogenetic reevaluation of Halszkaraptorinae places it as the sister clade to Unenlagiinae, indicating the bizarre features of unenlagiines previously interpreted as evidence of piscivory may also represent a mosaic of plesiomorphic, derived, and intermediate features. The anatomy of Halszkaraptor reveals that dromaeosaurids still possessed many features found in more basal maniraptoran and coelurosaur clades, including some that may have been tied to herbivory. Rather than being a semiaquatic piscavore, Halszkaraptor was a basal dromaeosaurid showing transitional features.

Though a tradeoff between growth rate and locomotor performance has been proposed, empirical data on this relationship are still limited. Here we statistically analyze the associations of growth rate and flight ability in birds by assessing how growth rate is correlated with three wing parameters of birds: flight muscle ratio, wing aspect ratio, and wing loading. We find that fast-growing birds tended to have higher flight muscle ratios and higher wing loadings than slow-growing birds, which suggests that fast-growing birds may have better takeoff performance, but lower efficiency in maneuvering flight. Accordingly, our findings suggest that the relationship between growth rate and flight ability is more complex than a simple tradeoff. Since the hindlimbs also contribute greatly to the locomotion of birds, future investigations on the relationship between growth rate and hindlimb performance will provide more insights into the evolution of birds.

The Early Cretaceous basal birds were known largely from just two-dimensionally preserved specimens from north-eastern China (Jehol Biota), which has hindered our understanding of the early evolution of birds. Here, we present a three-dimensionally-preserved skeleton (FPDM-V-9769) of a basal bird from the Early Cretaceous of Fukui, central Japan. Unique features in the pygostyle and humerus allow the assignment of FPDM-V-9769 to a new taxon, Fukuipteryx prima. FPDM-V-9769 exhibits a set of features comparable to that of other basalmost birds including Archaeopteryx. Osteohistological analyses indicate that FPDM-V-9769 is subadult. Phylogenetic analyses resolve F. prima as a non-ornithothoracine avialan basal to Jeholornis and outgroup of the Pygostylia. This phylogenetic result may imply a complex evolutionary history of basal birds. To our knowledge, FPDM-V-9769 represents the first record of the Early Cretaceous non-ornithothoracine avialan outside of the Jehol Biota and increases our understanding of their diversity and distribution during the time., A new fossil bird from the Early Cretaceous of Japan provides evidence that the evolutionary history of early birds was more complex than previously realised. This new species also has important implications for the distribution of early birds, demonstrating that they inhabited temperate, lowland regions.

The diverse array of phenotypes and courtship displays exhibited by birds-of-paradise have long fascinated scientists and nonscientists alike. Remarkably, almost nothing is known about the genomics of this iconic radiation. There are 41 species in 16 genera currently recognized within the birds-of-paradise family (Paradisaeidae), most of which are endemic to the island of New Guinea. In this study, we sequenced genomes of representatives from all five major clades within this family to characterize genomic changes that may have played a role in the evolution of the group's extensive phenotypic diversity. We found genes important for coloration, morphology, and feather and eye development to be under positive selection. In birds-of-paradise with complex lekking systems and strong sexual dimorphism, the core birds-of-paradise, we found Gene Ontology categories for “startle response” and “olfactory receptor activity” to be enriched among the gene families expanding significantly faster compared to the other birds in our study. Furthermore, we found novel families of retrovirus-like retrotransposons active in all three de novo genomes since the early diversification of the birds-of-paradise group, which might have played a role in the evolution of this fascinating group of birds.

The evolution of birds from a group of small dinosaurs between 170 million and 150 million years ago has emerged as a textbook example of a major evolutionary transformation in the fossil record ( 1 ). The attainment of powered flight—that is, active flapping that generates thrust—has been widely regarded, sometimes explicitly but often implicitly, as a long evolutionary march in which natural selection progressively refined one subgroup of dinosaurs into ever-better aerialists. However, recent fossil discoveries reveal a much more interesting story that is beginning to be corroborated by biomechanical studies. According to this story, the development of flight was chaotic, with different dinosaurs experimenting with different airborne behaviors using different airfoil and feather arrangements (see the figure), until ultimately only modern birds survived.

Most extant species are in clades with poor fossil records, and recent studies of comparative methods show they have low power to infer even highly simplified models of trait evolution without fossil data. Birds are a well-studied radiation, yet their early evolutionary patterns are still contentious. The fossil record suggests that birds underwent a rapid ecological radiation after the end-Cretaceous mass extinction, and several smaller, subsequent radiations. This hypothesized series of repeated radiations from fossil data is difficult to test using extant data alone. By uniting morphological and phylogenetic data on 604 extant genera of birds with morphological data on 58 species of extinct birds from 50 million years ago, the "halfway point" of avian evolution, I have been able to test how well extant-only methods predict the diversity of fossil forms. All extant-only methods underestimate the disparity, although the ratio of within- to between-clade disparity does suggest high early rates. The failure of standard models to predict high early disparity suggests that recent radiations are obscuring deep time patterns in the evolution of birds. Metrics from different models can be used in conjunction to provide more valuable insights than simply finding the model with the highest relative fit.

Birds are maniraptoran theropod dinosaurs. The evidence supporting the systematic position of Avialae as a derived clade within Dinosauria is voluminous and derived from multiple independent lines of evidence. In contrast, a paucity of selectively chosen data weakly support, at best, alternative proposals regarding the origin of birds and feathers. Opponents of the theory that birds are dinosaurs have frequently based their criticisms on unorthodox interpretations of paleontological data and misrepresentation of phylogenetic systematic methods. Moreover, arguments against the nested position of Avialae in Dinosauria have often conflated the logically distinct questions of avian origins, the evolution of flight, and the phylogenetic distribution of feathers. Motivated by a Perspectives article with numerous factual inaccuracies that recently appeared in The Auk, we provide a review of the full complement of facts pertaining to the avian origins debate and address the misplaced criticisms raised in...

Major transformations in brain size and proportions, such as the enlargement of the brain during the evolution of birds, are accompanied by profound modifications to the skull roof. However, the hypothesis of concerted evolution of shape between brain and skull roof over major phylogenetic transitions, and in particular of an ontogenetic relationship between specific regions of the brain and the skull roof, has never been formally tested. We performed 3D morphometric analyses to examine the deep history of brain and skull-roof morphology in Reptilia, focusing on changes during the well-documented transition from early reptiles through archosauromorphs, including nonavian dinosaurs, to birds. Non-avialan taxa cluster tightly together in morphospace, whereas Archaeopteryx and crown birds occupy a separate region. There is a one-to-one correspondence between the forebrain and frontal bone and the midbrain and parietal bone. Furthermore, the position of the forebrain–midbrain boundary correlates significantly with the position of the frontoparietal suture across the phylogenetic breadth of Reptilia and during the ontogeny of individual taxa. Conservation of position and identity in the skull roof is apparent, and there is no support for previous hypotheses that the avian parietal is a transformed postparietal. The correlation and apparent developmental link between regions of the brain and bony skull elements are likely to be ancestral to Tetrapoda and may be fundamental to all of Osteichthyes, coeval with the origin of the dermatocranium. Brain and skull development are intimately related across tetrapods. Here, the authors show a close relationship between brain and skull roof across evolutionary transitions and ontogenetic stages of reptiles.

The highly organized morphogenesis of bird feathers holds important phylo- and ontogenetic information on the evolution of birds, organogenesis, tissue regeneration, and the health status of individual animals. Altered topobiological patterns are regularly used as retrospective evidence for disturbed developmental trajectories due to the past exposure to environmental stressors. Using the most advanced high-resolution (5–70 µm) X-ray fluorescence microscopy (XFM), we describe in the feathers from three species of Procellariiformes hitherto unknown, depositions of elements (Zn, Ca, Br, Cu, Fe) that are independent of pigmentation or any underlying variation in density or polymer structure. In the case of Zn, the pattern across several species of Procellariiformes, but not other species, consisted of highly regular bands of Zn numbering 30–32, which may reflect the estimated number of days of active feather growth or the duration of the moult period. Thus, speculatively, the highly consistent Zn pattern might be the result of a so far unknown diurnal systemic regulation rather than local heterogeneity amongst the follicular stem cells.

Several flight modes are thought to have evolved during the early evolution of birds. Here, we use a combination of computational modelling and morphofunctional analyses to infer the flight properties of the raven-sized, Early Cretaceous birdSapeornis chaoyangensis—a likely candidate to have evolved soaring capabilities. Specifically, drawing information from (i) mechanical inferences of the deltopectoral crest of the humerus, (ii) wing shape (i.e. aspect ratio), (iii) estimations of power margin (i.e. difference between power required for flight and available power from muscles), (iv) gliding behaviour (i.e. forward speed and sinking speed), and (v) palaeobiological evidence, we conclude thatS. chaoyangensiswas a thermal soarer with an ecology similar to that of living South American screamers. Our results indicate that as early as 125 Ma, some birds evolved the morphological and aerodynamic requirements for soaring on continental thermals, a conclusion that highlights the degree of ecological, functional and behavioural diversity that resulted from the first major evolutionary radiation of birds.

Nest sanitation-related traits have often been explained at the intraspecific level as reducing the probability of infection or detection by predators and parasites, but its evolution within the avian phylogeny is still poorly understood. We compiled detailed information of such traits for more than 400 bird species and, by means of modern comparative methodologies, we reconstructed the evolution of adults' contribution to removing their offspring's faeces and the production of faecal sacs by nestlings. Furthermore, because the functional hypotheses used to explain nest sanitation behaviour assume potential effects of brood size, body mass, nestling period and diet, we explored the association between these traits and those related to nest sanitation in a phylogenetically controlled framework. Our results suggest that parental removal of nestling faeces has driven the evolution of faecal sacs, while the ancestral states involved birds with faecal sacs removed by parents. These results support the long-held idea that faecal sacs facilitate the removal of faeces by parents. Moreover, we found that animal diets and small body sizes have favoured the evolution of faecal sacs suggesting the existence of some chemical and physical constraints in relation to the evolution of the mucous covering. Our results highlight the importance of nest sanitation in the evolution of birds and their life history characteristics, J.D.I. was financed by a postdoctoral contract (TAHUB-104) from the program ‘Andalucía Talent Hub’ (cofunded by the European Union's Seventh Framework Program Marie Skłodowska-Curie actions (COFUND) and the regional Government of Andalucía). Funding was partially provided by the Spanish Ministerio de Economía y Competitividad (European funds (FEDER)) (CGL2013-48193-C3-1-P).

Birds have evolved on the planet for over 150 million years and become the most speciose clade of modern vertebrates. Their biological success has been ascribed to important evolutionary novelties including feathers, powered flight, and respiratory system, some of which have a deep evolutionary history even before the origin of birds. The last two decades have witnessed a wealth of exceptionally preserved feathered non-avian dinosaurs and primitive birds, which provide the most compelling evidence supporting the hypothesis that birds are descended from theropod dinosaurs. A handful of Mesozoic bird fossils have demonstrated how birds achieved their enormous biodiversity after diverging from their theropod relatives. On basis of recent fossil discoveries, we review how these new findings add to our understanding of the early avian evolution.

A new basal Avialae, Jeholornis curvipes sp. nov., from the Yixian Formation (Lower Cretaceous) of Liaoning Province (north-eastern China) is described. A revision of long-tailed birds from China and a phylogenetic analysis of basal Avialae suggest that Jeholornithiformes were paraphyletic, with Jixiangornis orientalis being the sister-taxon of pygostylia. The phylogenetic analysis also recovered that the tail reduction is a unique event in the evolution of birds. Jeholornis species were cursorial, nonperching, and seed-eating birds.

This communication presents a concise overview of reports in the literature concerning the occurrence of extratelencephalic fibre tracts in birds and the comparability of these tracts with the mammalian pyramidal tract. Emphasis is on the intratelencephalic organization, in particular that of the intratelencephalic sensorimotor circuits processing information from all important types of sense organs. It is suggested that two descending tracts, the occipitomesencephalic tract and the basal tractus superficialis medialis in birds have the same role in guiding behaviour as the pyramidal pathway in mammals. However, the differences in origin, trajectory and targets suggest that two independent systems may have developed in birds. One of these, the basal tractus superficialis medialis, represents the homologue of the pyramidal tract. It is suggested that the occipitomesencephalic tract is a specific feature of birds that has developed during the evolution from the early dinosaurs to birds. This suggestion follows from recent observations on the evolution of birds.

Hybridization is increasingly seen as an important source of adaptive genetic variation and biotic diversity. Recent phylogenetic studies on the early evolution of birds suggest that the early diversification of neoavian orders perhaps involved a period of extensive hybridization or incomplete lineage sorting. Phylogenetic error, saturation, long-branch attraction, and convergence make it difficult to detect ancient hybridization events and differentiate them from incomplete lineage sorting using sequence data. We used recently published retroposon marker data to visualize the early radiation of Neoaves within a phylogenetic network approach, and found that the most basal neoavian taxa indeed show a complex pattern of reticulated relationships. Moreover, the reticulation levels of different parts of the network are consistent with the insertion pattern of the retroposon elements. The use of network-based analyses on homoplasy-free data shows true conflicting signals and the taxa involved that are not represented in trees.

From complex songs to simple honks, birds produce sounds using a unique vocal organ called the syrinx. Located close to the heart at the tracheobronchial junction, vocal folds or membranes attached to modified mineralized rings vibrate to produce sound. Syringeal components were not thought to commonly enter the fossil record, and the few reported fossilized parts of the syrinx are geologically young (from the Pleistocene and Holocene (approximately 2.5 million years ago to the present)). The only known older syrinx is an Eocene specimen that was not described or illustrated. Data on the relationship between soft tissue structures and syringeal three-dimensional geometry are also exceptionally limited. Here we describe the first remains, to our knowledge, of a fossil syrinx from the Mesozoic Era, which are preserved in three dimensions in a specimen from the Late Cretaceous (approximately 66 to 69 million years ago) of Antarctica. With both cranial and postcranial remains, the new Vegavis iaai specimen is the most complete to be recovered from a part of the radiation of living birds (Aves). Enhanced-contrast X-ray computed tomography (CT) of syrinx structure in twelve extant non-passerine birds, as well as CT imaging of the Vegavis and Eocene syrinxes, informs both the reconstruction of ancestral states in birds and properties of the vocal organ in the extinct species. Fused rings in Vegavis form a well-mineralized pessulus, a derived neognath bird feature, proposed to anchor enlarged vocal folds or labia. Left-right bronchial asymmetry, as seen in Vegavis, is only known in extant birds with two sets of vocal fold sound sources. The new data show the fossilization potential of the avian vocal organ and beg the question why these remains have not been found in other dinosaurs. The lack of other Mesozoic tracheobronchial remains, and the poorly mineralized condition in archosaurian taxa without a syrinx, may indicate that a complex syrinx was a late arising feature in the evolution of birds, well after the origin of flight and respiratory innovations. Fil: Clarke, Julia A.. University of Texas at Austin; Estados Unidos Fil: Chatterjee, Sankar. Texas Tech University; Estados Unidos Fil: Li, Zhiheng. University of Texas at Austin; Estados Unidos. Chinese Academy of Sciences; República de China Fil: Riede, Tobias. Midwestern University; Estados Unidos 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 Fil: Goller, Franz. University of Utah; Estados Unidos Fil: Isasi, Marcelo Pablo. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”; Argentina Fil: Martinioni, Daniel Roberto. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Austral de Investigaciones Científicas; Argentina Fil: Mussel, Francisco J.. Universidad de Buenos Aires. Facultad de Ciencias Exactas y Naturales. Departamento de Ciencias Geológicas; Argentina 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

Passerines, taxonomic group that comprises about two-thirds of the world's existing avian species, originated in the part of Gondwana that today forms Australia (Edwards and Boles, 2002). The ancient supercontinent Gondwana broke up about 180 million years ago, generating the landmasses that we recognize today as Africa, South America, Australia, Antarctica, the Indian subcontinent and the Arabian Peninsula. The book's subtitle makes reference to these major events, which are presented with great accuracy by the author. However, the aim of this book goes well beyond the topic of Australia as one of the cradles of avian evolution. More generally, and interesting not only for comparative psychologists and neuroscientists but also for the general public, this is a book on avian brains and intelligence. I suspect that, in a sense, Australian birds are an excuse for the author, an eminent animal behavior researcher who has made important contributions to the study of Australian (and non-Australian) animal cognition, to discuss some of the more general and central topics of animal intelligence in a broad perspective. In this sense, I believe the book is of interest to readers of any Hemisphere.

Assessment of competing theories for the evolution of avian flight is problematic, and tends to rest too heavily on reconstruction of the mode of life of one or a few specimens representing still fewer species. A more powerful method is to compare the sequence of character acquisition predicted by the various theories with the empirical sequence provided by cladistic phylogeny. Arboreal and cursorial theories incorrectly predict the sequence of character acquisition for several key features of avian evolution. We propose an alternative 'pouncing proavis' model for the evolution of flight. As well as being both biologically and evolutionarily plausible, the pouncing proavis model correctly predicts the evolutionary sequence of all five key features marking the evolution of birds.

This article contains a review of current scientific knowledge on the evolution of birds. Topics discussed include the origin of birds from dinosaurs, the evolution of characters associated with birds (feathers, powered flight, small body size, and beaks), the phylogenetic relationships among the main lineages of living birds and the evolutionary timing of separation of these lineages in earth history.

Coelurosauria is the most diverse clade of theropod dinosaurs. Much of this diversity is present in Paraves—the clade of dinosaurs containing dromaeosaurids, troodontids, and avialans. Paraves has over 160 million years of evolutionary history that continues to the present day. The clade represents the most diverse living tetrapod group (there are over 9000 extant species of Aves—a word used here as synonomous with “bird”), and it is at the root of the paravian radiation, when dromaeosaurids, troodontids, and avialans were diverging from one another, that we find the morphology and soft tissue changes associated with the origin of modern avian flight. Within the first 15 million years of known paravian evolutionary history members of this clade exhibited a difference of nearly four orders of magnitude in body size, a value that is similar to the extreme body size disparity present today in mammalian carnivorans, avians, and varanoid squamates. In this respect, Paraves is an important case study i...

Luis M. Chiappe, John Wiley and Sons, 2007, x+263 pp., ISBN 0-471-24723-5 For all their current popularity as objects of research interest, fossil birds remain chronically under-represented in the ...

Avian Evolution Research on the origin and evolution of birds has gathered pace in recent years, aided by a continuous stream of new fossil finds as well as molecular phylogenies. Bird origins, in particular, are now better understood than those of mammals, for which the early fossil record is relatively poor compared with that of birds. Xu et al. review progress in tracing the origins of birds from theropod dinosaurs, focusing especially on recent fossil finds of feathered dinosaurs of northeastern China. They integrate current research on developmental biology and functional anatomy with the paleontological record, to show how key features of birds—feathers, wings, and flight—originated and evolved, and radiated from their dinosaur forebears. Science , this issue [10.1126/science.1253293][1] [1]: /lookup/doi/10.1126/science.1253293

Sapeornithidae is a basal pygostylian family of Early Cretaceous primitive birds, in which only one genus and species, Sapeornis chaoyangensis, was reported before. This paper deals with a new genus and species of this family, Didactylornis jii gen. et sp. nov., which was unearthed from the Early Cretaceous Jiufotang Formation in western Liaoning. According to our phylogenetic analyses, both Didactylornis gen. nov. and Sapeornis form a sister group, which is basal to the clade formed by Confuciusornis and all the more derived birds, and more closely related to the short-tailed pygostylian birds than to the long-tailed avialian birds. The early history of pygostylian birds is poorly documented except for the studies of Confuciusornis and Sapeornis. The discovery of Didactylornis jii gen. et sp. nov. adds the new material for the study on the early evolution of birds.

It has been suggested that relative egg size in living birds is strongly correlated with the developmental mode of the young; "altricial" (helpless) or "precocial" (independent). Using a data set of extant taxa we show that altricial birds lay relatively larger eggs than their precocial counterparts but that this may be due to the small size of most altricial species. Smaller birds tend to lay relatively small eggs compared to large species. Nonetheless, a predictive egg mass-body mass relationship extends into the avian fossil record. Such a relationship is important to our understanding of avian evolution because relative egg size (and thus available developmental mode) was constrained in many early birds—oviduct diameter was limited by the presence of pubic fusion. Therefore we document the evolution of avian developmental strategies using morphology-based phylogenies for Mesozoic and extant avians and corroborate correlations between developmental strategies, egg weight and female body mass. The sequential loss of precocial features in hatchlings characterises the evolution of birds while altriciality is derived within Neoaves. A set of precocial strategies is seen in earlier lineages, including basal Neornithes (modern birds) and are implied in their Mesozoic counterparts—skeletal constraints on egg size, present in many Jurassic and Early Cretaceous birds (Archaeopteryx, Confuciusornis, Enantiornithes) were lost in later diverging lineages. Attributes of precociality were already present in a number of lineages of non-avian maniraptoran theropods. We propose that the evolution of "unrestricted egg size" may have precipitated subsequent development of the diverse reproductive strategies seen in living birds.

The skeletons of birds are universally described as lightweight as a result of selection for minimizing the energy required for flight. From a functional perspective, the weight (mass) of an animal relative to its lift-generating surfaces is a key determinant of the metabolic cost of flight. The evolution of birds has been characterized by many weight-saving adaptations that are reflected in bone shape, many of which strengthen and stiffen the skeleton. Although largely unstudied in birds, the material properties of bone tissue can also contribute to bone strength and stiffness. In this study, I calculated the density of the cranium, humerus and femur in passerine birds, rodents and bats by measuring bone mass and volume using helium displacement. I found that, on average, these bones are densest in birds, followed closely by bats. As bone density increases, so do bone stiffness and strength. Both of these optimization criteria are used in the design of strong and stiff, but lightweight, manmade airframes. By analogy, increased bone density in birds and bats may reflect adaptations for maximizing bone strength and stiffness while minimizing bone mass and volume. These data suggest that both bone shape and the material properties of bone tissue have played important roles in the evolution of flight. They also reconcile the conundrum of how bird skeletons can appear to be thin and delicate, yet contribute just as much to total body mass as do the skeletons of terrestrial mammals.

In the years between 1912 and 1916, the Danish artist and graphic designer Gerhard Heilmann published a series of articles in the journal of the Danish Ornithological Society. From the outset, Heilmann's work aroused international interest, and in 1926 it was published in English as The origin of birds, setting the international agenda for research in bird evolution for the next 40 years. In Denmark, however, Heilmann's highly original work was generally ignored or even ridiculed by zoologists. This article demonstrates how Heilmann's artistic abilities played an important role in securing him international renown as a palaeontologist, while at the same time his lack of scientific credentials led to his complete isolation from the Danish zoological establishment. And it suggests that Heilmann's unyielding efforts to solve the riddle of bird evolution in the borderland between art and science, reflected a deeply felt emotional and spiritual need to counteract the religious dogmatism that had permea...

The stone curlew, also known as thick-knee (Burhinus oedicnemus, BOE), represents a phylogenetically young species of the shorebirds (Charadriiformes) that exhibits one of the most atypical genome organizations known within the class of Aves, due to an extremely low diploid number (2n = 42) and only 6 pairs of microchromosomes in its complement. This distinct deviation from the ‘typical’ avian karyotype is attributed to repeated fusions of ancestral microchromosomes. In order to compare different species with this atypical avian karyotype and to investigate the chromosome rearrangement patterns, chromosome-specific painting probes representing the whole genome of the stone curlew were used to delineate chromosome homology between BOE and 5 species belonging to 5 different avian orders: herring gull (Charadriiformes), cockatiel (Psittaciformes), rock pigeon (Columbiformes), great gray owl (Strigiformes) and Eurasian coot (Gruiformes). Paints derived from the 20 BOE autosomes delimited 28 to 33 evolutionarily conserved segments in the karyotypes of the 5 species, similar to the number recognized by BOE paints in such a basal lineage as the chicken (28 conserved segments). This suggests a high degree of conservation in genome organization in birds. BOE paints also revealed some species-specific rearrangements. In particular, chromosomes BOE1–4 and 14, as well as to a large extent BOE5 and 6, showed conserved synteny with macrochromosomes, whereas homologous regions for BOE7–13 are found to be largely distributed on microchromosomes in the species investigated. Interestingly, the 6 pairs of BOE microchromosomes 15–20 appear to have undergone very few rearrangements in the 5 lineages investigated. Although the arrangements of BOE homologous segments on some chromosomes can be explained by complex fusions and inversions, the occurrence of homologous regions at multiple sites may point to fission of ancestral chromosomes in the karyotypes of the species investigated. However, the present results demonstrate that the ancestral microchromosomes most likely experienced fusion in the stone curlew lineage forming the medium-sized BOE chromosomes, while they have been conserved as microchromosomes in the other neoavian lineages.