What's in a Wing? Aptian Cretaceous Enantiornithine Bird Shows Perfect Flight Adaptation

Birds have an enormously long evolutionary history: The earliest of them, the famed Archaeopteryx, lived 150 million years ago in what is today southern Germany. However, whether these early birds were capable of flying -- and if so, how well -- has remained shrouded in scientific controversy. A new discovery published in the journal Scientific Reports documents the intricate arrangement of the muscles and ligaments that controlled the main feathers of the wing of an ancient bird, supporting the notion that at least some of the most ancient birds performed aerodynamic feats in a fashion similar to those of many living birds.

An international team of Spanish paleontologists and NHM's Director of the Dinosaur Institute, Dr. Luis M. Chiappe, studied the exceptionally preserved wing of a 125-million-year-old bird from central Spain. Beyond the bones preserved in the fossil, the tiny wing of this ancient bird reveals details of a complex network of muscles that in modern birds controls the fine adjustments of the wing's main feathers, allowing birds to master the sky.

"The anatomical match between the muscle network preserved in the fossil and those that characterize the wings of living birds strongly indicates that some of the earliest birds were capable of aerodynamic prowess like many present-day birds," said Chiappe, the investigation's senior scientist.

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open access paper link.

Evolution of Assymmetical Feathers in Basal Cretaceous Avians

Barb geometry of asymmetrical feathers reveals a transitional morphology in the evolution of avian flight

Authors:

Feo et al

Abstract:

The geometry of feather barbs (barb length and barb angle) determines feather vane asymmetry and vane rigidity, which are both critical to a feather's aerodynamic performance. Here, we describe the relationship between barb geometry and aerodynamic function across the evolutionary history of asymmetrical flight feathers, from Mesozoic taxa outside of modern avian diversity (Microraptor, Archaeopteryx, Sapeornis, Confuciusornis and the enantiornithine Eopengornis) to an extensive sample of modern birds. Contrary to previous assumptions, we find that barb angle is not related to vane-width asymmetry; instead barb angle varies with vane function, whereas barb length variation determines vane asymmetry. We demonstrate that barb geometry significantly differs among functionally distinct portions of flight feather vanes, and that cutting-edge leading vanes occupy a distinct region of morphospace characterized by small barb angles. This cutting-edge vane morphology is ubiquitous across a phylogenetically and functionally diverse sample of modern birds and Mesozoic stem birds, revealing a fundamental aerodynamic adaptation that has persisted from the Late Jurassic. However, in Mesozoic taxa stemward of Ornithurae and Enantiornithes, trailing vane barb geometry is distinctly different from that of modern birds. In both modern birds and enantiornithines, trailing vanes have larger barb angles than in comparatively stemward taxa like Archaeopteryx, which exhibit small trailing vane barb angles. This discovery reveals a previously unrecognized evolutionary transition in flight feather morphology, which has important implications for the flight capacity of early feathered theropods such as Archaeopteryx and Microraptor. Our findings suggest that the fully modern avian flight feather, and possibly a modern capacity for powered flight, evolved crownward of Confuciusornis, long after the origin of asymmetrical flight feathers, and much later than previously recognized.

Early Birds Were not Very Ecologically Diverse

Birds come in astounding variety—from hummingbirds to emus—and behave in myriad ways: they soar the skies, swim the waters, and forage the forests. But this wasn't always the case, according to research by scientists at the University of Chicago and the Field Museum.

The researchers found a striking lack of diversity in the earliest known fossil bird fauna (a set of species that lived at about the same time and in the same habitat). "There were no swans, no swallows, no herons, nothing like that. They were pretty much all between a sparrow and a crow," said Jonathan Mitchell, PhD student in the Committee on Evolutionary Biology, and lead author of the new study, published May 28, 2014, in Proceedings of the Royal Society B.

The scientists examined a group of bird fossils dating back to the Cretaceous period, around 125 million years ago, relatively soon after the emergence of birds. The fossils were collected from an area in China where there was once violent volcanic activity, leading to a plethora of well-preserved fossils as intermittent eruptions periodically killed many birds. The researchers examined the diversity of species in this sample. However, because fossils indicate only the physical characteristics of the birds, understanding the diversity in how the birds behaved required significant scientific legwork.

To tease out the ecological roles played by the prehistoric birds, the researchers used modern-day birds to build a statistical technique that could relate the physical characteristics of a bird to its diet, behavior and habitat. Long legs might be associated with birds that wade through water, for instance, and the shape of the beak might hint at what the bird ate. For this purpose, the scientists painstakingly measured 1,400 modern birds—mostly from the Field Museum's collections—and extracted the correlations between these measurements and the birds' behavior.

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Fortunguavis xiaotaizicus: A New Scansorial Cretaceous Enantiornithine Bird

A new robust enantiornithine bird from the Lower Cretaceous of China with scansorial adaptations

Authors:

Wang et al

Abstract:

We describe a new enantiornithine bird, Fortunguavis xiaotaizicus, gen. et sp. nov, from the Lower Cretaceous lacustrine deposits of the Jiufotang Formation in northeastern China. The new taxon has a strongly dorsoventrally bowed furcula indicating that enantiornithines evolved furcular morphologies in parallel with ornithuromorphs. The new specimen has very robust limbs compared with other enantiornithines and has an unique foot morphology with metatarsal II much shorter than metatarsal IV, robust pedal digits, and strongly recurved pedal unguals. Although recurved unguals characterize Enantiornithes, the extreme curvature present in Fortunguavis suggests scansorial specialization in this species. These features hint at a unique ecology for this taxon and further increase the known diversity of body plans in Early Cretaceous enantiornithines.

Birds Drove Cope's Rule in Pterosaur Evolution

Competition and constraint drove Cope's rule in the evolution of giant flying reptiles

Authors:

Benson et al

Abstract:

The pterosaurs, Mesozoic flying reptiles, attained wingspans of more than 10 m that greatly exceed the largest birds and challenge our understanding of size limits in flying animals. Pterosaurs have been used to illustrate Cope’s rule, the influential generalization that evolutionary lineages trend to increasingly large body sizes. However, unambiguous examples of Cope’s rule operating on extended timescales in large clades remain elusive, and the phylogenetic pattern and possible drivers of pterosaur gigantism are uncertain. Here we show 70 million years of highly constrained early evolution, followed by almost 80 million years of sustained, multi-lineage body size increases in pterosaurs. These results are supported by maximum-likelihood modelling of a comprehensive new pterosaur data set. The transition between these macroevolutionary regimes is coincident with the Early Cretaceous adaptive radiation of birds, supporting controversial hypotheses of bird–pterosaur competition, and suggesting that evolutionary competition can act as a macroevolutionary driver on extended geological timescales.

Were There Multiple Species of Euryapteryx Moa?

Complex Species Status for Extinct Moa (Aves: Dinornithiformes) from the Genus Euryapteryx

Authors:

Huynen et al

Abstract:

The exact species status of New Zealand's extinct moa remains unknown. In particular, moa belonging to the genus Euryapteryx have been difficult to classify. We use the DNA barcoding sequence on a range of Euryapteryx samples in an attempt to resolve the species status for this genus. We obtained mitochondrial control region and the barcoding region from Cytochrome Oxidase Subunit I (COI) from a number of new moa samples and use available sequences from previous moa phylogenies and eggshell data to try and clarify the species status of Euryapteryx. Using the COI barcoding region we show that species status in Euryapteryx is complex with no clear separation between various individuals. Eggshell, soil, and bone data suggests that a Euryapteryx subspecies likely exists on New Zealand's North Island and can be characterized by a single mitochondrial control region SNP. COI divergences between Euryapteryx individuals from the south of New Zealand's South Island and those from the Far North of the North Island exceed 1.6% and are likely to represent separate species. Individuals from other areas of New Zealand were unable to be clearly separated based on COI differences possibly as a result of repeated hybridisation events. Despite the accuracy of the COI barcoding region to determine species status in birds, including that for the other moa genera, for moa from the genus Euryapteryx, COI barcoding fails to provide a clear result, possibly as a consequence of repeated hybridisation events between these moa. A single control region SNP was identified however that segregates with the two general morphological variants determined for Euryapteryx; a smaller subspecies restricted to the North Island of New Zealand, and a larger subspecies, found on both New Zealand's North and South Island.

Protoazin parisiensis: Eocene Paleogene Hoatzin Fossil From France

Earliest and first Northern Hemispheric hoatzin fossils substantiate Old World origin of a “Neotropic endemic”

Authors:

Mayr et al

Abstract:

The recent identification of hoatzins (Opisthocomiformes) in the Miocene of Africa showed part of the evolution of these birds, which are now only found in South America, to have taken place outside the Neotropic region. Here, we describe a new fossil species from the late Eocene of France, which constitutes the earliest fossil record of hoatzins and the first one from the Northern Hemisphere. Protoazin parisiensis gen. et sp. nov. is more closely related to South American Opisthocomiformes than the African taxon Namibiavis and substantiates an Old World origin of hoatzins, as well as a relictual distribution of the single extant species. Although recognition of hoatzins in Europe may challenge their presumed transatlantic dispersal, there are still no North American fossils in support of an alternative, Northern Hemispheric, dispersal route. In addition to Opisthocomiformes, other avian taxa are known from the Cenozoic of Europe, the extant representatives of which are only found in South America. Recognition of hoatzins in the early Cenozoic of Europe is of particular significance because Opisthocomiformes have a fossil record in sub-Saharan Africa, which supports the hypothesis that extinction of at least some of these “South American” groups outside the Neotropic region was not primarily due to climatic factors.

Hongshanornis longicresta: a Barremian/Aptian Cretaceous Ornithuromorph With a Modern Bird's Flight Profile

A new specimen of the Early Cretaceous bird Hongshanornis longicresta: insights into the aerodynamics and diet of a basal ornithuromorph

Authors:

Chiappe et al

Abstract:

The discovery of Hongshanornis longicresta, a small ornithuromorph bird with unusually long hindlimb proportions, was followed by the discovery of two closely related species, Longicrusavis houi and Parahongshanornis chaoyangensis. Together forming the Hongshanornithidae, these species reveal important information about the early diversity and morphological specialization of ornithuromorphs, the clade that contains all living birds. Here we report on a new specimen (DNHM D2945/6) referable to Hongshanornis longicresta that contributes significant information to better understand the morphology, trophic ecology, and aerodynamics of this species, as well as the taxonomy of the Hongshanornithidae. Most notable are the well-preserved wings and feathered tail of DNHM D2945/6, which afford an accurate reconstruction of aerodynamic parameters indicating that as early as 125 million years ago, basal ornithuromorphs had evolved aerodynamic surfaces comparable in size and design to those of many modern birds, and flight modes alike to those of some small living birds.

Parvavis chuxiongensis: a new Enantiornithe Bird From Upper Cretaceous China

The first enantiornithine bird from the Upper Cretaceous of China

Authors:

Wang et al

Abstract:

A new Late Cretaceous avian taxon, Parvavis chuxiongensis, gen. et sp. nov., is reported here based on an incomplete skeleton from Upper Cretaceous lake deposits in Yunnan Province, southern China. A phylogenetic analysis of 32 taxa and 242 morphological characters resulted in three most parsimonious trees, the strict consensus tree of which places Parvavis chuxiongensis within Enantiornithes. Histological study shows that the bones of Parvavis were composed of parallel-fibered bone tissue without lines of arrested growth, and indicated that growth rate had slowed but had not stopped at any stage prior to death. The bones also lack the rough surface texture seen in juvenile birds. Therefore, the new bird was probably close to adult body size at the time of death. However, the specimen is surprisingly small, highlighting the wide range of body sizes in Upper Cretaceous enantiornithines. The new specimen also represents both the first known bird from the Upper Cretaceous of China and the first Mesozoic bird from the south of China, and thus extends the temporal and geographic range of Mesozoic birds in China.

Embryonic Enantiornithine Bird Fossils From Campanian Cretaceous Mongolia

An embryonic enantiornithine bird and associated eggs from the cretaceous of Mongolia

Authors:

E. N. Kurochkin, S. Chatterjee and K. E. Mikhailov

Abstract:

Enantiornithes is the most speciose clade of Cretaceous birds, but many taxa are known from isolated postcranial skeletons. Two embryonic enantiornithine bird skeletons of Gobipipus reshetovi gen. et sp. nov. from the Upper Cretaceous (Campanian) Barun Goyot Formation of the Gobi Desert in Mongolia provide new insights into the anatomy, radiation, and mode of development of early avialans. In recent times, both enantiornithine and ornithuromorph birds are known from the Barun Goyot Formation as well as from the Djadokhta and Nemegt Formations. The 80-million-year-old Gobipipus skeletons encased within eggshells shows several features characteristic of enantiornithine birds. The wing skeleton and shoulder girdle show morphological features indicating that Gobipipus achieved sophisticated powered flight. Gobipipus reshetovi gen. et sp. nov. is quite distinct from the sympatric enantiornithine species Gobipteryx minuta from the same strata in many anatomical features. Phylogenetic analysis of 26 avialan ingroup taxa based on distribution of 202 characters indicate that Gobipipus is a basal member of enantiornithine birds along with Confuciusornis and shares more characters with ornithuromorphs than previously recognized. The embryonic nature of Gobipipus specimens sheds new light on the developmental history of enantiornithine birds. The well-ossified bones of the fore- and hind limbs, and fusion of many skeletal elements indicate a precocial mode of development in Gobipipus. Apparently Gobipipus hatchlings could walk away from the ground nests as soon as they emerged from their eggs. The asymmetry of egg poles are unique features of Gobipipus eggs (oogenus Gobioolithus) among Cretaceous avialans. The microstructure of the shell in Gobioolithus eggs with the embryos of Gobipipus is typical avian (of ornithoid basic type) and less ratite-like in morphology of the spongy layer than is that in the other possible egg-remains of enantiornitine birds (oofamily Laevisoolithidae).

Miocene Neogene Kiwi Fossils Suggest Origins

Miocene fossils show that kiwi (Apteryx, Apterygidae) are probably not phyletic dwarves

Authors:

Worthy et al

Abstract:

Until now, kiwi (Apteryx, Apterygidae) have had no pre-Quaternary fossil record to inform on the timing of their arrival in New Zealand or on their inter-ratite relationships. Here we describe two fossils in a new genus of apterygid from Early Miocene sediments at St Bathans, Central Otago, minimally dated to 19–16 Ma. The new fossils indicate a markedly smaller and possibly volant bird, supporting a possible overwater dispersal origin to New Zealand of kiwi independent of moa. If the common ancestor of this early Miocene apterygid species and extant kiwi was similarly small and volant, then the phyletic dwarfing hypothesis to explain relatively small body size of kiwi compared with other ratites is incorrect. Apteryx includes five extant species distributed on North, South, Stewart and the nearshore islands of New Zealand. They are nocturnal, flightless and comparatively large birds, 1–3 kg, with morphological attributes that reveal an affinity with ratites, but others, such as their long bill, that differ markedly from all extant members of that clade. Although kiwi were long considered most closely related to sympatric moa (Dinornithiformes), all recent analyses of molecular data support a closer affinity to Australian ratites (Casuariidae). Usually assumed to have a vicariant origin in New Zealand (ca 80–60 Ma), a casuariid sister group relationship for kiwi, wherein the common ancestor was volant, would more easily allow a more recent arrival via overwater dispersal.

New Zealand Holocene Quaternary Giant Moas may not Have Been Robust Birds

Giant moa bird (Dinornis robustus, literally meaning 'robust strange bird') may not have actually had robust bones, according to new research conducted by The University of Manchester. The leg bones of one of the tallest birds that ever existed were actually rather like those of its modern (but distant) relatives, such as ostrich, emu and rhea, the studypublished in PLOS One today (18 December) shows.

The study, led by biomechanics researcher Charlotte Brassey, in collaboration with palaeobiologist Professor Richard Holdaway at the University of Canterbury in New Zealand, has found that the largest of the moa species had leg bones similar to those of modern flightless birds that can run fast, whereas a much smaller species of moa – from a different family - had an extremely robust skeleton.

Ms Brassey said:"Our research suggests that this group of birds came up with several different solutions to deal with the problem of supporting the large body necessary to process a diet of coarse vegetation.

"We know that these species of moa were living together in the same locations, at the same time. So we don't think the differences we're seeing in leg robustness are adaptations to a particular habitat type.

"Instead it seems they were perhaps engaging in different behaviours, although both could deal with extremely rough terrain."

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Tyrannosauroid Alioramus Retains Basal Sinuses Lost in Birds

The Cranial Pneumatic Sinuses of the Tyrannosaurid Alioramus (Dinosauria: Theropoda) and the Evolution of Cranial Pneumaticity in Theropod Dinosaurs

Authors:

Gold et al

Abstract:

Archosaurs and mammals exhibit skeletal pneumaticity, where bone is infilled by air-filled soft tissues. Some theropod dinosaurs possess extensively pneumatic skulls in which many of the individual bones are hollowed out by diverticula of three main cranial sinus systems: the paranasal, suborbital, and tympanic sinuses. Computed tomography (CT scanning) permits detailed study of the internal morphology of cranial sinuses. But only a few theropod specimens have yet been subjected to this type of analysis. We present CT scans of the remarkably preserved and disarticulated skull bones of the long-snouted tyrannosaurid theropod Alioramus. These scans indicate that Alioramus has extensive cranial pneumaticity, with pneumatic sinuses invading the maxilla, lacrimal, jugal, squamosal, quadrate, palatine, ectopterygoid, and surangular. Pneumaticity is not present, however, in the nasal, postorbital, quadratojugal, pterygoid, or angular. Comparisons between Alioramus and other theropods (most importantly the closely related Tyrannosaurus) show that the cranial sinuses of Alioramus are modified to fill the long-snouted skull of this taxon, and that Alioramus has an extreme degree of cranial pneumaticity compared to other theropods, which may be the result of the juvenile status of the specimen, a difference in feeding style between Alioramus and other theropods, or passive processes. Based on these comparisons, we provide a revised terminology of cranial pneumatic structures and review the distribution, variation, and evolution of cranial pneumaticity within theropod dinosaurs. This review illustrates that most theropods possess a common “groundplan” in which the maxilla and lacrimal are pneumatized, and that various theropods modify this groundplan by pneumatizing numerous other bones of the skull. Tyrannosaurids are very pneumatic compared to other theropods, particularly in the development of extensive ectopterygoid, quadrate, and palatine sinuses, as well as a pneumatic invasion into the surangular. Tyrannosauroids seem to retain many cranial sinuses, such as the jugal and nasal recesses, which are primitive for coelurosaurs but lost or apomorphically modified in taxa more closely related to birds.

Dinosaur Cove Site in Albanian Cretaceous Australia Has Landing Trackways From Flying Birds

Two fossilized footprints found at Dinosaur Cove in Victoria, Australia, were likely made by birds during the Early Cretaceous, making them the oldest known bird tracks in Australia.

The journal Palaeontology is publishing an analysis of the footprints led by Anthony Martin, a paleontologist at Emory University in Atlanta who specializes in trace fossils, which include tracks, burrows and nests. The study was co-authored by Patricia Vickers-Rich and Michael Hall of Monash University in Victoria and Thomas Rich of the Museum Victoria in Melbourne.

Much of the rocky coastal strata of Dinosaur Cove in southern Victoria were formed in river valleys in a polar climate during the Early Cretaceous. A great rift valley formed as the ancient supercontinent Gondwana broke up and Australia separated from Antarctica.

"These tracks are evidence that we had sizeable, flying birds living alongside other kinds of dinosaurs on these polar, river floodplains, about 105 million years ago," Martin says.

The thin-toed tracks in fluvial sandstone were likely made by two individual birds that were about the size of a great egret or a small heron, Martin says. Rear-pointing toes helped distinguish the tracks as avian, as opposed to a third nearby fossil track that was discovered at the same time, made by a non-avian theropod.

A long drag mark on one of the two bird tracks particularly interested Martin.

"I immediately knew what it was – a flight landing track – because I've seen many similar tracks made by egrets and herons on the sandy beaches of Georgia," Martin says.

Martin often leads student field trips to Georgia's coast and barrier islands, where he studies modern-day tracks and other life traces, to help him better identify fossil traces.

The ancient landing track from Australia "has a beautiful skid mark from the back toe dragging in the sand, likely caused as the bird was flapping its wings and coming in for a soft landing," Martin says. Fossils of landing tracks are rare, he adds, and could add to our understanding of the evolution of flight.

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Did Jeholornis Have TWO Tails?

The early bird gets two tails? A 120-million-year-old bird sported a long tail and a second, unexpected tail frond, paleontologists suggest. The discovery points to a complicated evolutionary path for the tails we see in birds today.

The second-oldest known bird, Jeholornis, lived in what is today China, along with a trove of other feathered dinosaurs discovered in the region over the last decade. Fossils show that Jeholornis was turkey-size, had claws on its wing forelimbs, and possessed three small teeth in its lower jaw. It was also thought to sport only a long fan-feathered tail at its back end. Now, however, paleontologists are claiming discovery of a second tail frond adorning the bird.

"The 'two-tail' plumage of Jeholornis is unique," according to the study, which was led by Jingmai O’Connor of the Chinese Academy of Sciences in Beijing. The report of the discovery of the tail frond was published in the Proceedings of the National Academy of Sciences.

Of 11 Jeholornis fossils that retain evidence of ancient plumage, 6 have signs of this frond of 11 feathers, which would have jutted above the bird's back at a jaunty, upright angle in a "visually striking" manner, according to the study.

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Therapsid-Archosaur Wars Continue: Flying Archosaur Steals Meal From Top Predator Therapsid

Eagle Attacks and Takes Sika Deer Staked Out for Amur Tiger.

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Bird-like Track Fossils in the Triassic Claim RETRACTED

In this Letter, we considered the bird-like footprints from the former Santo Domingo Formation of northwest Argentina to be of Late Triassic age. Recent radiometric dating1 of the sedimentary sequence containing these bird-like footprints (renamed as the Laguna Brava Formation) indicated a Late Eocene age. Further geological studies2 suggest that the region suffered a complex deformation during the Andean orogeny, including block rotation. In consequence, our previous inferences about the possible implications of this finding for the fossil record of Aves are no longer supported.

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De-Extinction: Bring Back Birds of Our Dreams

A New Large Juvenile Enantiornithine Bird of the Cretaceous Jehol Biota of China

A large bird from the Early Cretaceous of China: new information on the skull of enantiornithines

Authors:

1. Zihui Zhang
2. Luis M. Chiappe
3. Gang Han
4. Anusuya Chinsamy

Affiliations:

blocked by lack of access...

Abstract:

We describe the anatomy and bone histology of an enantiornithine specimen from the Early Cretaceous Jehol Group of northeastern China. CNUVB-0903 is larger than most Early Cretaceous enantiornithine birds and also different from all other named taxa. Thus, we erect the new species Zhouornis hani for the new specimen. CNUVB-0903 preserves a suite of new morphologies for the clade. Noteworthy are those of the braincase and occipital region of the skull, which was previously poorly known for enantiornithines. The morphology and placement of the large basipterygoid processes and the well-developed basisphenoid recess—comparable in morphology to those of non-avian dinosaurs—highlights the evolutionary conservatism of the enantiornithine skull. The histological characterization of CNUVB-0903 indicates that it was not yet a full-grown individual at the time of death. This, combined with the comparatively large size of the skeleton, supports previous evidence indicating that early in their history, enantiornithines were able to achieve relatively large sizes.

Digit Loss in Extent Archosaurs and Evolutionary Implications

Digit loss in archosaur evolution and the interplay between selection and constraints

Authors:

1. Merijn A. G. de Bakker (a)
2. Donald A. Fowler (a)
3. Kelly den Oude (a)
4. Esther M. Dondorp (a)
5. M. Carmen Garrido Navas (a)
6. Jaroslaw O. Horbanczuk (b)
7. Jean-Yves Sire (c)
8. Danuta Szczerbińska (d)
9. Michael K. Richardson (a)

Affiliations:

a. Department of Integrative Zoology, Institute of Biology, Leiden University, Sylvius Laboratory, Sylviusweg 72, 2333BE Leiden, the Netherlands

b. Institute of Genetics and Animal Breeding, Polish Academy of Sciences, 05-552 Jastrzębiec, Poland

c. Evolution & Développement du squelette, UMR 7138, Université Pierre et Marie Curie, 7 Quai Saint-Bernard, Bat A2, Case 5, 75005 Paris, France

d. Department of Poultry and Ornamental Bird Breeding, Western Pomeranian University of Technology, 20 Judyma Street, 71-466 Szczecin, Poland

Abstract:

Evolution involves interplay between natural selection and developmental constraints. This is seen, for example, when digits are lost from the limbs during evolution. Extant archosaurs (crocodiles and birds) show several instances of digit loss under different selective regimes, and show limbs with one, two, three, four or the ancestral number of five digits. The ‘lost’ digits sometimes persist for millions of years as developmental vestiges. Here we examine digit loss in the Nile crocodile and five birds, using markers of three successive stages of digit development. In two independent lineages under different selection, wing digit I and all its markers disappear. In contrast, hindlimb digit V persists in all species sampled, both as cartilage, and as Sox9- expressing precartilage domains, 250 million years after the adult digit disappeared. There is therefore a mismatch between evolution of the embryonic and adult phenotypes. All limbs, regardless of digit number, showed similar expression of sonic hedgehog (Shh). Even in the one-fingered emu wing, expression of posterior genes Hoxd11 and Hoxd12 was conserved, whereas expression of anterior genes Gli3 and Alx4 was not. We suggest that the persistence of digit V in the embryo may reflect constraints, particularly the conserved posterior gene networks associated with the zone of polarizing activity (ZPA). The more rapid and complete disappearance of digit I may reflect its ZPA-independent specification, and hence, weaker developmental constraints. Interacting with these constraints are selection pressures for limb functions such as flying and perching. This model may help to explain the diverse patterns of digit loss in tetrapods. Our study may also help to understand how selection on adults leads to changes in development.