3D atlas of tinamou (Neornithes: Tinamidae) pectoral morphology: Implications for reconstructing the ancestral neornithine flight apparatus.

Palaeognathae, the extant avian clade comprising the flightless ratites and flight-capable tinamous (Tinamidae), is the sister group to all other living birds, and recent phylogenetic studies illustrate that tinamous are phylogenetically nested within a paraphyletic assemblage of ratites. As the only extant palaeognaths that have retained the ability to fly, tinamous may provide key information on the nature of the flight apparatus of ancestral crown palaeognaths-and, in turn, crown birds-as well as insight into convergent modifications to the wing apparatus among extant ratite lineages. To reveal new information about the musculoskeletal anatomy of tinamous and facilitate development of computational biomechanical models of tinamou wing function, we generated a three-dimensional musculoskeletal model of the flight apparatus of the extant Andean tinamou (Nothoprocta pentlandii) using diffusible iodine-based contrast-enhanced computed tomography (diceCT). Origins and insertions of the pectoral flight musculature of N. pentlandii are generally consistent with those of other extant volant birds specialized for burst flight, and the entire suite of presumed ancestral neornithine flight muscles are present in N. pentlandii with the exception of the biceps slip. The pectoralis and supracoracoideus muscles are robust, similar to the condition in other extant burst-flying birds such as many extant Galliformes. Contrary to the condition in most extant Neognathae (the sister clade to Palaeognathae), the insertion of the pronator superficialis has a greater distal extent than the pronator profundus, although most other anatomical observations are broadly consistent with the conditions observed in extant neognaths. This work will help form a basis for future comparative studies of the avian musculoskeletal system, with implications for reconstructing the flight apparatus of ancestral crown birds and clarifying musculoskeletal modifications underlying the convergent origins of ratite flightlessness.

Cardiac morphology of North Island brown kiwi (Apteryx mantelli).

AIMS: To investigate the cardiac anatomy of North Island brown kiwi (Apteryx mantelli) through heart morphometric parameters measured at post-mortem examination. METHODS: Morphometric cardiac parameters were established at post-mortem examination of 20 North Island brown kiwi. Birds were classified by gender and age (chicks vs. adults). Measurements included: body mass, heart mass, sternal length, midpoint thickness of left ventricular free wall, midpoint thickness of right ventricular free wall and ratios of heart mass to body mass, left ventricular length to sternal length, right ventricular length to sternal length, length of left ventricle to right ventricle, interventricular septal thickness relative to the sternal length and interventricular septal thickness relative to the left ventricular length. Unadjusted estimates of the median difference and their 95% CI were then reported at each age and sex for all the cardiac morphometric parameters and their ratios. RESULTS: The small sample size led to wide 95% CI for the median difference between gender and age for the cardiac morphometric measurements. Nevertheless, between adult female and male kiwi, the estimated population median differences for heart mass (2.2 (95% CI = -2.9-5.6) g), length (1.2 (95% CI = -2.2-5.6) mm), width (6.1 (95% CI = -1.0-8.2) mm), left ventricular free wall length (5.5 (95% CI = -0.5-8.8) mm) and right ventricular free wall length (2.6 (95% CI = -3.7-6.9) mm) were established. In adult North Island brown kiwi, the heart mass is 0.8 (95% CI = 0.7-0.8)% of the body mass. CONCLUSIONS: The precision of the differences noted in heart measurements recorded between male and female kiwi at each age was limited by the low sample size available for this study. This led to wide CI and an inability to adjust differences observed for gender by differences in other confounders such as body size. With this caveat, there is weak evidence that adult female kiwi have a larger heart size and mass than the adult males. CLINICAL RELEVANCE: These results can be used to improve the diagnosis of cardiac disease in kiwi at post-mortem examination and aid in interpretation of the results of echocardiography in live birds for the antemortem diagnosis of cardiac disorders.

Symmetry breaking in the embryonic skin triggers directional and sequential plumage patterning.

The development of an organism involves the formation of patterns from initially homogeneous surfaces in a reproducible manner. Simulations of various theoretical models recapitulate final states of natural patterns, yet drawing testable hypotheses from those often remains difficult. Consequently, little is known about pattern-forming events. Here, we surveyed plumage patterns and their emergence in Galliformes, ratites, passerines, and penguins, together representing the three major taxa of the avian phylogeny, and built a unified model that not only reproduces final patterns but also intrinsically generates shared and varying directionality, sequence, and duration of patterning. We used in vivo and ex vivo experiments to test its parameter-based predictions. We showed that directional and sequential pattern progression depends on a species-specific prepattern: an initial break in surface symmetry launches a travelling front of sharply defined, oriented domains with self-organising capacity. This front propagates through the timely transfer of increased cell density mediated by cell proliferation, which controls overall patterning duration. These results show that universal mechanisms combining prepatterning and self-organisation govern the timely emergence of the plumage pattern in birds.

Genetic parameters for tonic immobility, body weight, and morphological traits of the red-winged tinamou (Rhynchotus rufescens).

This study was carried out to estimate genetic parameters for morphology, body weight, and tonic immobility traits in the red-winged tinamou (Rhynchotus rufescens). Information on 690 birds was used and genetic parameters were estimated using Bayesian methods under a multi-trait animal model. The following traits were considered in this study: tarsal length (TL), bill length (BL), wing length (WL), head width (HW), bill width (BW), mature weight (MW), weight at 90 days (W90), and tonic immobility (TI). The heritability showed estimates between 0.15 for wing length and 0.56 for bill length. Positive and negative genetic correlations were estimated, ranging from - 0.33 to 0.81. All the morphological, production, and behavioral traits studied will have moderate to high response to selection. The body weight at 90 days is a better alternative for use in breeding programs and its selection would not lead to an increase in the time of tonic immobility. Both the selection for weight gain and for reduction of tonic immobility time would lead to an increase in the size of the legs of the red-winged tinamou, which could be advantageous for thermal control of these birds in tropical systems.

The phylogenetic significance of the morphology of the syrinx, hyoid and larynx, of the southern cassowary, Casuarius casuarius (Aves, Palaeognathae).

BACKGROUND: Palaeognathae is a basal clade within Aves and include the large and flightless ratites and the smaller, volant tinamous. Although much research has been conducted on various aspects of palaeognath morphology, ecology, and evolutionary history, there are still areas which require investigation. This study aimed to fill gaps in our knowledge of the Southern Cassowary, Casuarius casuarius, for which information on the skeletal systems of the syrinx, hyoid and larynx is lacking - despite these structures having been recognised as performing key functional roles associated with vocalisation, respiration and feeding. Previous research into the syrinx and hyoid have also indicated these structures to be valuable for determining evolutionary relationships among neognath taxa, and thus suggest they would also be informative for palaeognath phylogenetic analyses, which still exhibits strong conflict between morphological and molecular trees. RESULTS: The morphology of the syrinx, hyoid and larynx of C. casuarius is described from CT scans. The syrinx is of the simple tracheo-bronchial syrinx type, lacking specialised elements such as the pessulus; the hyoid is relatively short with longer ceratobranchials compared to epibranchials; and the larynx is comprised of entirely cartilaginous, standard avian anatomical elements including a concave, basin-like cricoid and fused cricoid wings. As in the larynx, both the syrinx and hyoid lack ossification and all three structures were most similar to Dromaius. We documented substantial variation across palaeognaths in the skeletal character states of the syrinx, hyoid, and larynx, using both the literature and novel observations (e.g. of C. casuarius). Notably, new synapomorphies linking Dinornithiformes and Tinamidae are identified, consistent with the molecular evidence for this clade. These shared morphological character traits include the ossification of the cricoid and arytenoid cartilages, and an additional cranial character, the articulation between the maxillary process of the nasal and the maxilla. CONCLUSION: Syrinx, hyoid and larynx characters of palaeognaths display greater concordance with molecular trees than do other morphological traits. These structures might therefore be less prone to homoplasy related to flightlessness and gigantism, compared to typical morphological traits emphasised in previous phylogenetic studies.

How to walk carrying a huge egg? Trade-offs between locomotion and reproduction explain the special pelvis and leg anatomy in kiwi (Aves; Apteryx spp.).

Kiwi (Aves; genus Apteryx) are famous for laying an enormous egg in comparison with their relatively small body size. Considering the peculiar gait of this flightless bird, we suspected the existence of morpho-functional trade-offs between reproduction and locomotion. To understand how structural constraints, imposed by a large egg size, might influence the terrestrial locomotion of Apteryx, we analysed the anatomy of the limb osteomuscular system in two species of kiwi (Apteryx mantelli and Apteryx owenii). We performed detailed dissections and brought to light specific anatomical features of kiwi, in comparison with other ratites and neognathous birds. Our osteological study revealed a strongly curved pelvis, a rigid tail, and enlarged ribs. Our myology study showed an unusual location of the caudofemoralis muscle origin and insertion. The insertion of the pars pelvica along the entire caudal face of the femur, contrasts with the proximal insertion usually seen in other birds. Additionally, the pars caudalis originates along the entire tail, whereas it only inserts on the uropygium in the other birds. To interpret these specificities from a functional point of view, we built three-dimensional osteomuscular models based on computed tomography scans, radiographies and our dissections. We chose three postures associated with reproductive constraints: the standing position of a gravid compared with a non-gravid bird, as well as the brooding position. The 3D model of the brooding position suggested that the enlarged ribs could support the bodyweight when leaning on the huge egg in both males and females. Moreover, we found that in gravid females, the unusual shape of the pelvis and tail allowed the huge egg to sit ventrally below the pelvis, whereas it is held closer to the rachis in other birds. The specific conformation of the limb and the insertions of the two parses of the caudofemoralis help to maintain the tail flexed, and to keep the legs adducted when carrying the egg. The caudal location of the hip and its flexed position explains the long stance phase during the strange gait of kiwi, revealing the functional trade-off between reproduction and locomotion in this emblematic New Zealand bird.

Nocturnal giants: evolution of the sensory ecology in elephant birds and other palaeognaths inferred from digital brain reconstructions.

The recently extinct Malagasy elephant birds (Palaeognathae, Aepyornithiformes) included the largest birds that ever lived. Elephant bird neuroanatomy is understudied but can shed light on the lifestyle of these enigmatic birds. Palaeoneurological studies can provide clues to the ecologies and behaviours of extinct birds because avian brain shape is correlated with neurological function. We digitally reconstruct endocasts of two elephant bird species, Aepyornis maximus and A. hildebrandti, and compare them with representatives of all major extant and recently extinct palaeognath lineages. Among palaeognaths, we find large olfactory bulbs in taxa generally occupying forested environments where visual cues used in foraging are likely to be limited. We detected variation in olfactory bulb size among elephant bird species, possibly indicating interspecific variation in habitat. Elephant birds exhibited extremely reduced optic lobes, a condition also observed in the nocturnal kiwi. Kiwi, the sister taxon of elephant birds, have effectively replaced their visual systems with hyperdeveloped olfactory, somatosensory and auditory systems useful for foraging. We interpret these results as evidence for nocturnality among elephant birds. Vision was likely deemphasized in the ancestor of elephant birds and kiwi. These results show a previously unreported trend towards decreased visual capacity apparently exclusive to flightless, nocturnal taxa endemic to predator-depauperate islands.

Comparison and Evaluation of the Effectiveness of Two Approaches of Diffusible Iodine-Based Contrast-Enhanced Computed Tomography (diceCT) for Avian Cephalic Material.

Diffusible iodine-based contrast-enhanced computed tomography presents a comparatively new tool kit for imaging fine-scale three-dimensional phenotypes that is rapidly becoming standard anatomical practice. However, relatively few studies have attempted to look at subtle differences in staining protocols or attempted to model tissue reactions to gain insight into staining mechanisms. Here, two iodine-based contrast agents, iodine-ethanol (I2 E) and iodine-potassium iodide (I2 KI) in neutral buffered formalin , were applied to avian cephalic specimens to investigate their effectiveness. We found that the two solutions had markedly different results for staining of mineralized skeletal tissues (i.e., bone). Other tissues, including muscles, epithelia, and common connective tissues (e.g., lamina propria) were assessed individually and show minor differences in the sorption of iodine. Numerical simulations suggest that different results from I2 E and I2 KI-formaldehyde staining are due to different partition coefficients and retardation factors of tissues, fixation effects, as well as distinct iodine diffusion and sorption patterns. We found a clear positive relationship between glycogen concentration and grayscale values measured within muscle, epithelia, nervous tissues, and glands. We also found the use of ethanol for tissue fixation and following I2 E staining outperforms I2 KI-formaldehyde by providing higher efficiency for acquiring greater contrast both between different soft tissues and between mineralized and nonmineralized tissues.

Is Cerebellar Architecture Shaped by Sensory Ecology in the New Zealand Kiwi (Apteryx mantelli).

Among some mammals and birds, the cerebellar architecture appears to be adapted to the animal's ecological niche, particularly their sensory ecology and behavior. This relationship is, however, not well understood. To explore this, we examined the expression of zebrin II (ZII) in the cerebellum of the kiwi (Apteryx mantelli), a fully nocturnal bird with auditory, tactile, and olfactory specializations and a reduced visual system. We predicted that the cerebellar architecture, particularly those regions receiving visual inputs and those that receive trigeminal afferents from their beak, would be modified in accordance with their unique way of life. The general stripe-and-transverse region architecture characteristic of birds is present in kiwi, with some differences. Folium IXcd was characterized by large ZII-positive stripes and all Purkinje cells in the flocculus were ZII positive, features that resemble those of small mammals and suggest a visual ecology unlike that of other birds. The central region in kiwi appeared reduced or modified, with folium IV containing ZII+/- stripes, unlike that of most birds, but similar to that of Chilean tinamous. It is possible that a reduced visual system has contributed to a small central region, although increased trigeminal input and flightlessness have undoubtedly played a role in shaping its architecture. Overall, like in mammals, the cerebellar architecture in kiwi and other birds may be substantially modified to serve a particular ecological niche, although we still require a larger comparative data set to fully understand this relationship.

Genomic support for a moa-tinamou clade and adaptive morphological convergence in flightless ratites.

One of the most startling discoveries in avian molecular phylogenetics is that the volant tinamous are embedded in the flightless ratites, but this topology remains controversial because recent morphological phylogenies place tinamous as the closest relative of a monophyletic ratite clade. Here, we integrate new phylogenomic sequences from 1,448 nuclear DNA loci totaling almost 1 million bp from the extinct little bush moa, Chilean tinamou, and emu with available sequences from ostrich, elegant crested tinamou, four neognaths, and the green anole. Phylogenetic analysis using standard homogeneous models and heterogeneous models robust to common topological artifacts recovered compelling support for ratite paraphyly with the little bush moa closest to tinamous within ratites. Ratite paraphyly was further corroborated by eight independent CR1 retroposon insertions. Analysis of morphological characters reinterpreted on a 27-gene paleognath topology indicates that many characters are convergent in the ratites, probably as the result of adaptation to a cursorial life style.

IMAGING DIAGNOSIS--USE OF RADIOGRAPHY AND COMPUTED TOMOGRAPHY IN THE DIAGNOSIS OF A MINERALIZED YOLK SAC IN A BROWN KIWI (APTERYX MANTELLI).

A 12-day-old Brown Kiwi (Apteryx mantelli) was presented with anorexia, torticollis, head-tilt, and coelomic distension. Radiographs showed an ill-defined, fat-opaque, coelomic mass displacing viscera craniodorsally. Curvilinear mineral opacities were superimposed over the ventral aspect of the mass. Computed tomography demonstrated the presence of mineral within the periphery of a fat attenuating mass consistent with a retained yolk sac. A deutectomy (yolk sac excision) was performed. Histopathology of the excised tissue confirmed the diagnosis of a retained yolk sac with multifocal mineralization.

Anatomical specializations for enhanced olfactory sensitivity in kiwi, Apteryx mantelli.

The ability to function in a nocturnal and ground-dwelling niche requires a unique set of sensory specializations. The New Zealand kiwi has shifted away from vision, instead relying on auditory and tactile stimuli to function in its environment and locate prey. Behavioral evidence suggests that kiwi also rely on their sense of smell, using olfactory cues in foraging and possibly also in communication and social interactions. Anatomical studies appear to support these observations: the olfactory bulbs and tubercles have been suggested to be large in the kiwi relative to other birds, although the extent of this enlargement is poorly understood. In this study, we examine the size of the olfactory bulbs in kiwi and compare them with 55 other bird species, including emus, ostriches, rheas, tinamous, and 2 extinct species of moa (Dinornithiformes). We also examine the cytoarchitecture of the olfactory bulbs and olfactory epithelium to determine if any neural specializations beyond size are present that would increase olfactory acuity. Kiwi were a clear outlier in our analysis, with olfactory bulbs that are proportionately larger than those of any other bird in this study. Emus, close relatives of the kiwi, also had a relative enlargement of the olfactory bulbs, possibly supporting a phylogenetic link to well-developed olfaction. The olfactory bulbs in kiwi are almost in direct contact with the olfactory epithelium, which is indeed well developed and complex, with olfactory receptor cells occupying a large percentage of the epithelium. The anatomy of the kiwi olfactory system supports an enhancement for olfactory sensitivities, which is undoubtedly associated with their unique nocturnal niche.

Paleoneurology of stem palaeognaths clarifies the plesiomorphic condition of the crown bird central nervous system.

Lithornithidae, an assemblage of volant Palaeogene fossil birds, provide our clearest insights into the early evolutionary history of Palaeognathae, the clade that today includes the flightless ratites and volant tinamous. The neotype specimen of Lithornis vulturinus, from the early Eocene (approximately 53 million years ago) of Europe, includes a partial neurocranium that has never been thoroughly investigated. Here, we describe these cranial remains including the nearly complete digital endocasts of the brain and bony labyrinth. The telencephalon of Lithornis is expanded and its optic lobes are ventrally shifted, as is typical for crown birds. The foramen magnum is positioned caudally, rather than flexed ventrally as in some crown birds, with the optic lobes, cerebellum, and foramen magnum shifted further ventrally. The overall brain shape is similar to that of tinamous, the only extant clade of flying palaeognaths, suggesting that several aspects of tinamou neuroanatomy may have been evolutionarily conserved since at least the early Cenozoic. The estimated ratio of the optic lobe's surface area relative to the total brain suggests a diurnal ecology. Lithornis may provide the clearest insights to date into the neuroanatomy of the ancestral crown bird, combining an ancestrally unflexed brain with a caudally oriented connection with the spinal cord, a moderately enlarged telencephalon, and ventrally shifted, enlarged optic lobes.

Ancient DNA reveals extreme egg morphology and nesting behavior in New Zealand's extinct moa.

New Zealand's extinct flightless moa radiated rapidly into a large number of morphologically diverse species, which produced an equally large range of egg morphologies. The exact number of moa species, as well as the characteristics of the eggs they laid, remains contentious. Moreover, like most extinct species, we understand little about their nesting and incubation habits. We used a modified ancient DNA extraction procedure to recover exogenous mitochondrial and nuclear DNA from the inside and outside surfaces of moa eggs. We used sequences from the inside of 69 eggshells to directly assign these remains to seven of the 10 currently recognized moa species. In addition we were able to assign, to the species level, six of the rare reconstructed "whole" eggs. These molecular results enabled us to identify two distinct lineages within the genus Euryapteryx. Members of these lineages differed in eggshell thickness, with one lineage being characterized by a relatively thin eggshell. Unexpectedly, several thin-shelled eggs were also shown to belong to the heaviest moa of the genera Dinornis, Euryapteryx and Emeus, making these, to our knowledge, the most fragile of all avian eggs measured to date. Moreover, sex-specific DNA recovered from the outer surfaces of eggshells belonging to species of Dinornis and Euryapteryx suggest that these very thin eggs were likely to have been incubated by the lighter males. The thin nature of the eggshells of these larger species of moa, even if incubated by the male, suggests that egg breakage in these species would have been common if the typical contact method of avian egg incubation was used.

Testing gradual and speciational models of evolution in extant taxa: the example of ratites.

Ever since Eldredge and Gould proposed their model of punctuated equilibria, evolutionary biologists have debated how often this model is the best description of nature and how important it is compared to the more gradual models of evolution expected from natural selection and the neo-Darwinian paradigm. Recently, Cubo proposed a method to test whether morphological data in extant ratites are more compatible with a gradual or with a speciational model (close to the punctuated equilibrium model). As shown by our simulations, a new method to test the mode of evolution of characters (involving regression of standardized contrasts on their expected standard deviation) is easier to implement and more powerful than the previously proposed method, but the Mesquite module comet (aimed at investigating evolutionary models using comparative data) performs better still. Uncertainties in branch length estimates are probably the largest source of potential error. Cubo hypothesized that heterochronic mechanisms may underlie morphological changes in bone shape during the evolution of ratites. He predicted that the outcome of these changes may be consistent with a speciational model of character evolution because heterochronic changes can be instantaneous in terms of geological time. Analysis of a more extensive data set confirms his prediction despite branch length uncertainties: evolution in ratites has been mostly speciational for shape-related characters. However, it has been mostly gradual for size-related ones.

Comparative intralimb coordination in avian bipedal locomotion.

Analyses of how intralimb coordination during locomotion varies within and across different taxa are necessary for understanding the morphological and neurological basis for locomotion in general. Previous findings suggest that intralimb proportions are the major source of kinematic variation that governs intralimb coordination across taxa. Also, independence of kinematics from habitat preference and phylogenetic position has been suggested for mammals. This leads to the hypothesis that among equally sized bird species exhibiting equal limb proportions, similar kinematics can be observed. To test this hypothesis, the bipedal locomotion of two distantly related ground-dwelling bird species (Eudromia elegans and Coturnix coturnix) and of a less terrestrial species (Corvus monedula) was investigated by means of a biplanar high-speed X-ray videographic analysis. Birds exhibited similar intralimb proportions and were filmed over a broad range of speed while moving on a treadmill. Joint and limb element angles, as well as pelvic rotations, were quantified. Regarding fore-aft motions of the limb joints and elements, a congruent pattern of intralimb coordination was observed among all experimental species. The sample of species suggests that this is largely independent of their habitat preference and systematic position and seems to be related to demands for coping with an irregular terrain with a minimum of necessary control. Hence, the initial hypothesis was confirmed. However, this congruence is not found when looking at medio-lateral limb motions and pelvic rotations, showing distinct differences between ground-dwellers (e.g. largely restricted to a parasagittal plane) and C. monedula (e.g. increased mobility of the hip joint).

Morphometric analysis of telencephalic structure in a variety of neognath and paleognath bird species reveals regional differences associated with specific behavioral traits.

Birds exhibit a huge array of behavior, ecology and physiology, and occupy nearly every environment on earth, ranging from the desert outback of Australia to the tropical rain forests of Panama. Some birds have adopted a fully nocturnal lifestyle, such as the barn owl and kiwi, while others, such as the albatross, spend nearly their entire life flying over the ocean. Each species has evolved unique adaptations over millions of years to function in their respective niche. In order to increase processing power or network efficiency, many of these adaptations require enlargements and/or specializations of the brain as a whole or of specific brain regions. In this study, we examine the relative size and morphology of 9 telencephalic regions in a number of Paleognath and Neognath birds and relate the findings to differences in behavior and sensory ecology. We pay particular attention to those species that have undergone a relative enlargement of the telencephalon to determine whether this relative increase in telencephalic size is homogeneous across different brain regions or whether particular regions have become differentially enlarged. The analysis indicates that changes in the relative size of telencephalic regions are not homogeneous, with every species showing hypertrophy or hypotrophy of at least one of them. The three-dimensional structure of these regions in different species was also variable, in particular that of the mesopallium in kiwi. The findings from this study provide further evidence that the changes in relative brain size in birds reflect a process of mosaic evolution.

Cortical growth marks reveal extended juvenile development in New Zealand moa.

Cyclical growth marks in cortical bone, deposited before attainment of adult body size, reflect osteogenetic changes caused by annual rhythms and are a general phenomenon in non-avian ectothermic and endothermic tetrapods. However, the growth periods of ornithurines (the theropod group including all modern birds) are usually apomorphically shortened to less than a year, so annual growth marks are almost unknown in this group. Here we show that cortical growth marks are frequent in long bones of New Zealand's moa (Aves: Dinornithiformes), a recently extinct ratite order. Moa showed the exaggerated K-selected life-history strategy formerly common in the New Zealand avifauna, and in some instances took almost a decade to attain skeletal maturity. This indicates that reproductive maturity in moa was extremely delayed relative to all extant birds. The two presently recognized moa families (Dinornithidae and Emeidae) also showed different postnatal growth rates, which were associated with their relative differences in body size. Both species of giant Dinornis moa attained their massive stature (up to 240 kg live mass) by accelerating their juvenile growth rate compared to the smaller emeid moa species, rather than by extending the skeletal growth period.

Definitive fossil evidence for the extant avian radiation in the Cretaceous.

Long-standing controversy surrounds the question of whether living bird lineages emerged after non-avian dinosaur extinction at the Cretaceous/Tertiary (K/T) boundary or whether these lineages coexisted with other dinosaurs and passed through this mass extinction event. Inferences from biogeography and molecular sequence data (but see ref. 10) project major avian lineages deep into the Cretaceous period, implying their 'mass survival' at the K/T boundary. By contrast, it has been argued that the fossil record refutes this hypothesis, placing a 'big bang' of avian radiation only after the end of the Cretaceous. However, other fossil data--fragmentary bones referred to extant bird lineages--have been considered inconclusive. These data have never been subjected to phylogenetic analysis. Here we identify a rare, partial skeleton from the Maastrichtian of Antarctica as the first Cretaceous fossil definitively placed within the extant bird radiation. Several phylogenetic analyses supported by independent histological data indicate that a new species, Vegavis iaai, is a part of Anseriformes (waterfowl) and is most closely related to Anatidae, which includes true ducks. A minimum of five divergences within Aves before the K/T boundary are inferred from the placement of Vegavis; at least duck, chicken and ratite bird relatives were coextant with non-avian dinosaurs.

Osteological description of casque ontogeny in the southern cassowary (Casuarius casuarius) using micro-CT imaging.

Extant cassowaries (Casuarius) are unique flightless birds found in the tropics of Indo-Australia. They have garnered substantial attention from anatomists with focus centered on the bony makeup and function of their conspicuous cranial casques, located dorsally above the orbits and neurocranium. The osteological patterning of the casque has been formally described previously; however, there are differing interpretations between authors. These variable descriptions suggest that an anatomical understanding of casque anatomy and its constituent elements may be enhanced by developmental studies aimed at further elucidating this bizarre structure. In the present study, we clarify casque osteology of the southern cassowary (C. casuarius) by detailing casque anatomy across an extensive growth series for the first time. We used micro-computed tomography (µCT) imaging to visualize embryonic development and post-hatching ontogeny through adulthood. We also sampled closely related emus (Dromaius novaehollandiae) and ostriches (Struthio camelus) to provide valuable comparative context. We found that southern cassowary casques are comprised of three paired (i.e., nasals, lacrimals, frontals) and two unpaired elements (i.e., mesethmoid, median casque element). Although lacrimals have rarely been considered as casque elements, the contribution to the casque structure was evident in µCT images. The median casque element has often been cited as a portion of the mesethmoid. However, through comparisons between immature C. casuarius and D. novaehollandiae, we document the median casque element as a distinct unit from the mesethmoid.