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Description of natal down of the ostrich (Struthio camelus) and comparison with common quail (Coturnix coturnix): Developmental and evolutionary implications
Abstract
Natal down is a feather stage that differs in both form and function from the definitive feathers of adult birds. It has a simpler structure that has been speculated to be similar to the body coverings of non-avian dinosaurs. However, inference of the evolution of natal down has been limited by our understanding of its structural variation in extant birds. Most descriptive work has focused on neognathous birds, limiting our knowledge of the full diversity of feathers in extant taxa. Here, we describe the natal down of a post-hatch ostrich (Struthio camelus) and compare it to that of a post-hatch quail (Coturnix coturnix). We confirm the presence of featherless spaces (apteria) in S. camelus and the lack of barbules on the tips of natal down in both species. We also find differences between dorsal and ventral natal down structures, such as barbule density in S. camelus and the extent of the bare portion of the barb in both species. Surprisingly, we do not find that the neoptiles of either species follow the ideal morphologies for increasing insulation. Finally, we hypothesize that the different barb types present in S. camelus natal down result from a large addition of new barb ridges during development, which is not known except in feathers with a rachis. These results have implications for our understanding of how structure informs function and development in understudied feather types, such as those shared by non-avian dinosaurs.
... This work allowed refining phylogenetic relationships prior to the advent of genetic tools (Clench, 1995), and showed that within passerines, there is subtle variation in tract shape, but no variation in tract area (it covers 1/3rd of the dorsum in all Estrildid finches; 40). Strikingly, the rest of the avian phylogeny has been poorly covered: pterylography has been quantified with varying degrees of precision only in (i) the emu (Bailleul et al., 2019;Ho et al., 2019), ostrich (Bailleul et al., 2019;Ho et al., 2019;Urban, 2023), and Chilean Tinamou (Ho et al., 2019), all part of the ancestral-most super-order Paleognathae, (ii) a dozen species in the super-order Galloanserae, such as the North American grouse (Clark, 1899), mallard duck (Humphrey and Clark, 1961), domestic duck (Ho et al., 2019), Japanese quail (Haupaix et al., 2018), screamer (Demay, 1940), domestic chicken (Haupaix et al., 2018) and common pheasant (Haupaix et al., 2018), and (iii) very few non-passerine species in the Neoaves super-order, namely the eared dove (Clark, 1918), red-capped coua (Berger and Lunk, 1954), gentoo penguin (Bailleul et al., 2019), and pileated woodpecker (Lyman and Clark, 1893); (Fig. 2). Comparison between all studied species showed that the extent and shape of tracts varies greatly. ...
Globally, high elevation habitats have been independently colonized by taxa separated by millions of years of evolution. Mountains thus represent excellent systems to study how distantly related species adapt to the same environmental challenges. Cold temperatures influence the elevational distribution of birds along montane gradients. Yet the eco‐physiological adaptations that may explain this pattern, such as variation in insulative feather structure across high elevation and low elevation species has not been quantified. We used a comparative approach to understand if elevation, evolutionary history and body size drive variation in thermo‐insulative feather traits across 1715 specimens of 249 Himalayan passerines. Controlling for phylogenetic relationships between species, we found that the proportion of the feather's plumulaceous (downy) section increased with elevation. Body size also had a predictable effect on thermo‐insulative variables with small birds having relatively longer feathers and thus a more insulative plumage than large birds. We show that an increase in the proportion of the feather's downy section at colder temperatures is an evolutionarily widespread response across temperate and tropical taxa, and overall, smaller‐bodied birds tend to have longer and more insulative feathers. Our results reveal convergent patterns in feather structure variation as a response to cold temperatures across species separated by millions of years of evolution.
Down feathers are the first feather types that appear in both the phylogenetic and the ontogenetic history of birds. Although it is widely acknowledged that the primary function of downy elements is insulation, little is known about the interspecific variability in the structural morphology of these feathers, and the environmental factors that have influenced their evolution. Here, we collected samples of down and afterfeathers from 156 bird species and measured key morphological characters that define the insulatory properties of the downy layer. We then tested if habitat and climatic conditions could explain the observed between‐species variation in down feather structure. We show that habitat has a very strong and clearly defined effect on down feather morphology. Feather size, barbule length and nodus density all decreased from terrestrial towards aquatic birds, with riparian species exhibiting intermediate characters. Wintering climate, expressed as windchill (a combined measure of the ambient temperature and wind speed) had limited effects on down morphology, colder climate only being associated with higher nodus density in dorsal down feathers. Overall, an aquatic lifestyle selects for a denser plumulaceous layer, while the effect of harsh wintering conditions on downy structures appear limited. These results provide key evidence of adaptations to habitat at the level of the downy layer, both on the scale of macro‐ and micro‐elements of the plumage. Moreover, they reveal characters of convergent evolution in the avian plumage and mammalian fur, that match the varying needs of insulation in terrestrial and aquatic modes of life.
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The rachises of extant feathers, composed of dense cortex and spongy internal medulla, are flexible and light, yet stiff enough to withstand the load required for flight, among other functions. Incomplete knowledge of early feathers prevents a full understanding of how cylindrical rachises have evolved. Bizarre feathers with unusually wide and flattened rachises, known as “rachis‐dominated feathers” (RDFs) have been observed in fossil non‐avian and avian theropods. Newly discovered RDFs embedded in early Late Cretaceous Burmese ambers (∼99 Ma) suggest the unusually wide and flattened rachises mainly consist of a dorsal cortex, lacking a medulla and a ventral cortex. Coupled with findings on extant feather morphogenesis, known fossil RDFs were categorized into three morphotypes based on their rachidial configurations. For each morphotype, potential developmental scenarios were depicted by referring to the rachidial development in chickens, and relative stiffness of each morphotype was estimated through functional simulations. The results suggest rachises of RDFs are developmentally equivalent to a variety of immature stages of cylindrical rachises. Similar rachidial morphotypes documented in extant penguins suggest that the RDFs are not unique to Mesozoic theropods, though they are likely to have evolved independently in extant penguins.
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The Jurassic stem bird Archaeopteryx is an iconic transitional fossil, with an intermediate morphology combining features of non-avian dinosaurs and crown Aves. Importantly, fossils of Archaeopteryx preserve not only the bones but also details of the plumage and therefore help shed light on the evolution of feathers, wings, and avian flight. Plumage is preserved in multiple individuals, allowing a detailed documentation of the feathers of the wings, tail, hindlimbs, and body. In some features, Archaeopteryx’ plumage is remarkably modern, yet in others, it is strikingly primitive. As in extant birds, remiges and coverts are enlarged and overlap to form airfoils. Remiges and rectrices exhibit asymmetrical, pennaceous vanes, with interlocking barbules. The hindlimbs bear large, vaned feathers as in Microraptor and Anchiornis. Rectrices are numerous and extend the full length of the tail to the hips. The plumage of crown Aves was assembled in a stepwise fashion from Anchiornis through Archaeopteryx, culminating in a modern arrangement in ornithothoracines. Subsequent stasis in feather and wing morphology likely reflects aerodynamic and developmental constraints. Feather morphology and arrangement in Archaeopteryx are consistent with lift-generating function, and the wing loading and aspect ratio are comparable to modern birds, consistent with gliding and perhaps flapping flight. The plumage of Archaeopteryx is intermediate between Anchiornis and more derived Pygostylia, suggesting a degree of flight ability intermediate between the two.
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.
Even in the absence of associated skeletal material, isolated feathers in amber remain of high scientific interest. The remarkable preservation of these delicate structures in amber implies a potential for significantly improving our knowledge of feather evolution and diversity. A large sample set of 150 Burmese amber specimens (Upper Cretaceous, ~99 Ma) containing feathers is herein described. Several structural types can be differentiated, including flight feathers, contour feathers, semiplumes, and filoplumes. In some cases, peculiar pigmentation patterns and structural features can be documented. Additionally, different developmental stages have been captured in this assemblage with some examples of erupting feathers or neoptile plumage. Comparisons with previous studies, including skeletal material described in amber, show that Enantiornithes and non avialan dinosaurs are most likely represented in the assemblage, as well as unknown taxa.
Exceptionally preserved fossil feathers and feather-like integumentary structures provide valuable insights into the early evolution of feathers and flight, but taphonomic biases often make interpretations at the microstructural and ultrastructural levels ambiguous. Maturation experiments have been demonstrated to be useful for investigating the taphonomic alterations of soft tissues, including feathers, during diagenesis. However, experimentally matured feathers resembling fossil feathers preserving keratinous matrix have not yet been obtained. Here we employ experimental maturation to obtain feathers corresponding to different degradation stages, and compare these matured feathers with untreated feathers and fossil feathers at the macroscopic, microstructural, and ultrastructural levels. Results show that several features of thermally matured feathers are similar to those found in fossil feathers. The fusion of barbules that occurred in thermally matured feathers suggests that such a process could occur during diagenesis, making barbules difficult to identify in fossil feathers. Under the most extreme experimental condition, the keratinous matrix can partially survive when the whole feather is turned into ash-like remains and many melanosomes are exposed. Moreover, our results show that the keratinous matrix immediately surrounding melanosomes appears to be more resistant to degradation than the unpigmented keratinous matrix, supporting the hypothesis that melanin can act as a fixative agent to prevent the degradation of keratin.
Birds are the most abundant terrestrial vertebrates and their diversity is greatly shaped by the feathers. How avian evolution is linked to feather evolution has long been a fascinating question. Numerous excellent studies have shed light on this complex relationship by investigating feather diversity and its underlying molecular mechanisms. However, most have focused on adult domestic birds, and the contribution of feather diversity to environmental adaptation has not been well-studied. In this review, we described bird diversity using the traditional concept of the altricial-precocial spectrum in bird hatchlings. We combined the spectrum with a recently published avian phylogeny to profile the spectrum evolution. We then focused on the discrete diagnostic character of the spectrum, the natal down, and propose a hypothesis for the precocial-to-altricial evolution. For the underlying molecular mechanisms in feather diversity and bird evolution, we reviewed the literature and constructed the known mechanisms for feather tract definition and natal down development. Finally, we suggested some future directions for research on altricial-precocial divergence, which may expand our understanding of the relationship between natal down diversity and bird evolution.
Molting-the process replacing one plumage with another-is a critically important biological function in Aves. This process annually replaces the feather coat, damaged by normal wear and tear, produces ontogenetic changes in feathering, and produces alternate breeding plumages associated with reproductive activity in adults. Immature, growing feathers are encased in a keratinous sheath, giving them a narrow, tubular, and featureless appearance. The complete loss of the sheath indicates the feather is mature. Despite the wealth of integumentary data published from the Jehol Biota, immature feathers have never been definitively reported, although they may potentially be preserved in a juvenile specimen of the non-avian oviraptorosaur theropod dinosaur Similicaudipteryx from the 120 Ma Jiufotang Formation. A developing feather has been reported in a 99 Ma enantiornithine neonate preserved in Burmese amber, in which three-dimensional preservation makes interpretations of integumentary structures more straightforward. Here we report on probable immature feathers in four juvenile enantiornithines (Aves: Ornithothoraces) from the Jehol Group. As observed in developing feathers in extant birds, the purported immature fossil feathers appear proximally narrow and featureless with barbs protruding only distally. Based on our observations, we suggest that similar-appearing feather structures preserved on the manus and tibiotarsus in the holotype of the enantiornithine Cruralispennia multidonta may alternatively be interpreted as immature feathers. The presence of immature feathers in combination with sexually dimorphic ornamental feathers in juvenile enantiornithines suggests the complex molting patterns of Neornithes, in which such ornaments only appear after several years (following several molts) when reproductive activity is achieved, are limited to a subset of crownward avians.
We describe three-dimensionally preserved feathers in mid-Cretaceous Burmese amber that share macro-morphological similarities (e.g., proportionally wide rachis with a “medial stripe”) with lithic, two-dimensionally preserved rachis-dominated feathers, first recognized in the Jehol Biota. These feathers in amber reveal a unique ventrally concave and dorsoventrally thin rachis, and a dorsal groove (sometimes pigmented) that we identify as the “medial stripe” visible in many rachis-dominated rectrices of Mesozoic birds. The distally pennaceous portion of these feathers shows differentiated proximal and distal barbules, the latter with hooklets forming interlocking barbs. Micro-CT scans and transverse sections demonstrate the absence of histodifferentiated cortex and medullary pith of the rachis and barb rami. The highly differentiated barbules combined with the lack of obvious histodifferentiation of the barb rami or rachis suggests that these feathers could have been formed without the full suite and developmental interplay of intermediate filament alpha keratins and corneous beta-proteins that is employed in the cornification process of modern feathers. This study thus highlights how the development of these feathers might have differed from that of their modern counterparts, namely in the morphogenesis of the ventral components of the rachis and barb rami. We suggest that the concave ventral surface of the rachis of these Cretaceous feathers is not homologous with the ventral groove of modern rachises. Our study of these Burmese feathers also confirms previous claims, based on two-dimensional fossils, that they correspond to an extinct morphotype and it cautions about the common practice of extrapolating developmental aspects (and mechanical attributes) of modern feathers to those of stem birds (and their dinosaurian outgroups) because the latter need not to have developed through identical pathways.
The current study investigated the macroscopic and microscopic differences between pennaceous and plumulaceous feathers. The morphology of the barbules distinguished pennaceous and plumulaceous feathers, particularly the shape of barbules during their development. In pennaceous feathers, the initial barbules were large and elongated or pyriform in shape, while plumulaceous feathers had small, thin, elongated initial barbules. The spinous barbules were characteristic of pennaceous feathers. The histochemical reactivity of both feather types for Mallory trichrome, orange G, and acridine orange, safranin O, PAS, and methylene blue was determined. Keratin was detected by Mallory trichrome, orange G, and acridine orange. In conclusion, the histochemical properties of pennaceous and plumulaceous feathers of quail, particularly the distribution and nature of keratin during development, should be considered in future studies. The unique morphological features of pennaceous and plumulaceous feathers could be used as a guide for phylogenetic identification. Anat Rec, 2019. © 2019 American Association for Anatomy Anat Rec, 303:1865–1883, 2020. © 2019 American Association for Anatomy
We examined variation in five microscopic plumulaceous (downy) feather characters of eighteen species of dabbling (Anatini) and diving (Aythyini, Mergini) ducks to quantify the differences between these tribes, and to explain how the plumulaceous feather ultrastructure in ducks may be influenced by different ecological requirements. Over 75% of the variation in feather characters among these ducks was explained by the first two components of a principal components analysis (PCA). Component 1 explained 51% of the variation and was positively correlated with the characters that quantified the number of barbules with expanded nodes and the number of expanded nodes on barbules. The microscopic feather characters of dabbling ducks (Anatini) have triangular-shaped, expanded nodes on most proximal barbules, whereas diving ducks (Aythyini and Mergini) lack expanded nodes on some barbules. Anatini also have a greater density of expanded nodes per barbule, wider nodes, shorter distance between expanded nodes, and longer barbule length. Further analysis of node density across all taxa showed that as dive depths increase, the number of expanded nodes per barbule decreases, and in the deepest divers many of the barbules completely lack expanded nodes. The significantly greater density of expanded nodes in dabbling ducks suggests that the downy nodes may function to trap more air. Diving species have fewer expanded nodes, less buoyant plumage, and are more efficient at foraging in deeper water than dabbling ducks.
All roads lead to regulation
Species from widely divergent taxa can experience similar changes in traits. What underlying genetic drivers cause these parallel changes remains an open question. Sackton et al. looked across groups of birds that have repeatedly lost flight, the ratites and tinamous, and found that there is convergence in the regulatory regions associated with genes related to flight, but not within the protein coding regions. Changes within these regulatory regions influenced limb development and may represent quick paths toward convergent change across taxa.
Science , this issue p. 74
Feathers are arranged in a precise pattern in avian skin. They first arise during development in a row along the dorsal midline, with rows of new feather buds added sequentially in a spreading wave. We show that the patterning of feathers relies on coupled fibroblast growth factor (FGF) and bone morphogenetic protein (BMP) signalling together with mesenchymal cell movement, acting in a coordinated reaction-diffusion-taxis system. This periodic patterning system is partly mechanochemical, with mechanical-chemical integration occurring through a positive feedback loop centred on FGF20, which induces cell aggregation, mechanically compressing the epidermis to rapidly intensify FGF20 expression. The travelling wave of feather formation is imposed by expanding expression of Ectodysplasin A (EDA), which initiates the expression of FGF20. The EDA wave spreads across a mesenchymal cell density gradient, triggering pattern formation by lowering the threshold of mesenchymal cells required to begin to form a feather bud. These waves, and the precise arrangement of feather primordia, are lost in the flightless emu and ostrich, though via different developmental routes. The ostrich retains the tract arrangement characteristic of birds in general but lays down feather primordia without a wave, akin to the process of hair follicle formation in mammalian embryos. The embryonic emu skin lacks sufficient cells to enact feather formation, causing failure of tract formation, and instead the entire skin gains feather primordia through a later process. This work shows that a reaction-diffusion-taxis system, integrated with mechanical processes, generates the feather array. In flighted birds, the key role of the EDA/Ectodysplasin A receptor (EDAR) pathway in vertebrate skin patterning has been recast to activate this process in a quasi-1-dimensional manner, imposing highly ordered pattern formation.
Pterosaurs were the first vertebrates to achieve true flapping flight, but in the absence of living representatives, many questions concerning their biology and lifestyle remain unresolved. Pycnofibres—the integumentary coverings of pterosaurs—are particularly enigmatic: although many reconstructions depict fur-like coverings composed of pycnofibres, their affinities and function are not fully understood. Here, we report the preservation in two anurognathid pterosaur specimens of morphologically diverse pycnofibres that show diagnostic features of feathers, including non-vaned grouped filaments and bilaterally branched filaments, hitherto considered unique to maniraptoran dinosaurs, and preserved melanosomes with diverse geometries. These findings could imply that feathers had deep evolutionary origins in ancestral archosaurs, or that these structures arose independently in pterosaurs. The presence of feather-like structures suggests that anurognathids, and potentially other pterosaurs, possessed a dense filamentous covering that probably functioned in thermoregulation, tactile sensing, signalling and aerodynamics. © 2018, The Author(s), under exclusive licence to Springer Nature Limited.
Over the last 20 years, compression fossils of feathers surrounding dinosaurs have greatly expanded our understanding of the origin and evolution of feathers. One of the most peculiar feather morphotypes discovered to date are rachis dominated feathers (RDFs), which have also been referred to as proximally ribbon-like pennaceous feathers (PRPFs). These elongate feathers are only found in the tail plumage, typically occurring in pairs with both streamer (not proximally ribbon-like) and racket-plume morphologies recognized. Here we describe a large sample set of isolated and paired RDFs from Upper Cretaceous Burmese amber (~ 99 Ma). Amber preserves the finest details of these fragile structures in three dimensions, demonstrating that RDFs form a distinct feather morphotype with a ventrally open rachis, and with significant variability in pigmentation, microstructure, and symmetry.
Identifying feather morphology in extinct dinosaurs is challenging due to dense overlapping of filaments within fossilized plumage and the fact that some extinct feather morphologies are unlike those of extant birds or those predicted from an ‘evo-devo’ model of feather evolution. Here, we compare a range of dinosaur taxa with preserved integumentary appendages using high-resolution photographs to better understand fossil feather morphology and gain insight into their function and evolution. A specimen of the basal paravian Anchiornis possesses contour feathers disarticulated from the plumage, revealing a novel feather type: a ‘shaggy’, open-vaned, bifurcated feather with long barbs attached to a short rachis, which is much simpler than the contour feathers of most extant birds. In contrast, it is likely that the contour feathers of Sinosauropteryx were simpler than those seen in Anchiornis; a ‘tuft’ morphology of multiple barbs connected at their bases (e.g. via a shared follicle), but lacking a rachis, is tentatively inferred. However, conclusive morphological descriptions await the discovery of isolated Sinosauropteryx contour feathers. Paravian wing feathers also show potentially plesiomorphic traits. Comparison with Confuciusornis suggests that Anchiornis wing feathers were at least partially open-vaned. Combined with the interpretation of Anchiornis contour feathers, this suggests that differentiated barbicels are relatively derived compared to pennaceous feathers and the appearance of wings. ‘Shaggy’ contour feathers probably influenced thermoregulatory and water repellence abilities, and, in combination with open-vaned wing feathers, would have decreased aerodynamic efficiency. Simplified, open-vaned wing feathers were also observed on the oviraptorosaur Caudipteryx, consistent with, but not necessarily diagnostic of, its suggested flightlessness. Taken together, these observations have broad implications for how we depict a wide variety of dinosaurs and how we view the function and evolution of feathers.
Studies modelling heat transfer of bird plumage design suggest that insulative properties can be attributed to the density and structure of the downy layer, whereas waterproofing is the result of the outer layer, comprised of contour feathers. In this study, we test how habitat and thermal condition affect feather mass and density of body feathers (contour, semiplume and downy feathers) measured on the ventral and dorsal sides of the body, using a phylogenetic comparative analysis of 152 bird species.
Our results demonstrate that feather mass and the density of downy feathers are higher in species that inhabit colder environments, whereas total feather density is higher of species breeding under intermediate temperatures compared to the ones breeding under more extreme conditions. The density of contour feathers, depending on the body region, is either quadratically related or negatively correlated with minimum winter temperature.
The density of contour and downy feathers, measured on both sides of the body, is higher in aquatic than in terrestrial birds. However, among the former, diving behaviour does not select for further increases in body feather mass or density.
The results of this study provides key insights into how the plumage of birds is adapted to different environments and lifestyles and provides a basis for understanding the diverse range and the evolution of variation in these characteristics.
A plain language summary is available for this article.
Genuine fossils with exquisitely preserved plumage from the Late Jurassic and Early Cretaceous of northeastern China have recently revealed that bird-like theropod dinosaurs had long pennaceous feathers along their hindlimbs and may have used their four wings to glide or fly. Thus, it has been postulated that early bird flight might initially have involved four wings (Xu et al. Nature 421:335–340, 2003; Hu et al. Nature 461:640–643, 2009; Han et al. Nat Commun 5:4382, 2014). Here, we describe Serikornis sungei gen. et sp. nov., a new feathered theropod from the Tiaojishan Fm (Late Jurassic) of Liaoning Province, China. Its skeletal morphology suggests a ground-dwelling ecology with no flying adaptations. Our phylogenetic analysis places Serikornis, together with other Late Jurassic paravians from China, as a basal paravians, outside the Eumaniraptora clade. The tail of Serikornis is covered proximally by filaments and distally by slender rectrices. Thin symmetrical remiges lacking barbules are attached along its forelimbs and elongate hindlimb feathers extend up to its toes, suggesting that hindlimb remiges evolved in ground-dwelling maniraptorans before being co-opted to an arboreal lifestyle or flight.
Burmese amber has recently provided some detailed glimpses of plumage, soft tissues, and osteology of juvenile enantiornithine birds, but these insights have been restricted to isolated wing apices. Here we describe nearly half of a hatchling individual, based on osteological and soft tissue data obtained from the skull, neck, feet, and wing, and identified as a member of the extinct avian clade Enantiornithes. Preserved soft tissue provides the unique opportunity to observe the external opening of the ear, the eyelid, and fine details of tarsal scutellation. The new amber specimen yields the most complete view of hatchling plumage and integument yet to be recovered from the Cretaceous, including details of pterylosis, feather microstructure, and pigmentation patterns. The hatchling was encapsulated during the earliest stages of its feather production, providing a point for comparisons to other forms of body fossils, as well as isolated feathers found in Cretaceous ambers. The plumage preserves an unusual combination of precocial and altricial features unlike any living hatchling bird, having functional remiges combined with sparse body feathers. Unusual feather morphotypes on the legs, feet, and tail suggest that first generation feathers in the Enantiornithes may have been much more like contour feathers than the natal down observed in many modern birds. However, these regions also preserve filamentous feathers that appear comparable to the protofeathers observed in more primitive theropods. Overall, the new specimen brings a new level of detail to our understanding of the anatomy of the juvenile stages of the most species-rich clade of pre-modern birds and contributes to mounting data that enantiornithine development drastically differed from that of Neornithes.
Insulation is an essential component of nest structure that helps provide incubation requirements for birds. Many species of waterfowl breed in high latitudes where rapid heat loss can necessitate a high energetic input from parents and use down feathers to line their nests. Common Eider (Somateria mollissima) nest down has exceptional insulating properties but the microstructural mechanisms behind the feather properties have not been thoroughly examined.
Here, we hypothesized that insulating properties of nest down are correlated to down feather (plumule) microstructure. We tested the thermal efficiency (fill power) and cohesion of plumules from nests of two Icelandic colonies of wild Common Eiders and compared them to properties of plumules of wild Greylag Goose (Anser anser). We then used electron microscopy to examine the morphological basis of feather insulating properties. We found that Greylag Goose down has higher fill power (i.e. traps more air) but much lower cohesion (i.e. less prone to stick together) compared to Common Eider down. These differences were related to interspecific variation in feather microstructure. Down cohesion increased with the number of barbule microstructures (prongs) that create strong points of contact among feathers. Eider down feathers also had longer barbules than Greylag Goose down feathers, likely increasing their air-trapping capacity. Feather properties of these two species might reflect the demands of their contrasting evolutionary history. In Greylag Goose, a temperate, terrestrial species, plumule microstructure may optimize heat trapping. In Common Eiders, a diving duck that nests in arctic and subarctic waters, plumule structure may have evolved to maximize cohesion over thermal insulation, which would both reduce buoyancy during their foraging dives and enable nest down to withstand strong arctic winds.
Body shape is a fundamental expression of organismal biology, but its quantitative
reconstruction in fossil vertebrates is rare. Due to the absence of fossilized soft tissue
evidence, the functional consequences of basal paravian body shape and its implications
for the origins of avians and flight are not yet fully understood. Here we reconstruct the
quantitative body outline of a fossil paravian Anchiornis based on high-definition images
of soft tissues revealed by laser-stimulated fluorescence. This body outline confirms
patagia-bearing arms, drumstick-shaped legs and a slender tail, features that were probably
widespread among paravians. Finely preserved details also reveal similarities in propatagial
and footpad form between basal paravians and modern birds, extending their record to
the Late Jurassic. The body outline and soft tissue details suggest significant functional
decoupling between the legs and tail in at least some basal paravians. The number of
seemingly modern propatagial traits hint that feathering was a significant factor in how basal
paravians utilized arm, leg and tail function for aerodynamic benefit.
In the two decades since the discovery of feathered dinosaurs [1-3], the range of plumage known from non-avialan theropods has expanded significantly, confirming several features predicted by developmentally informed models of feather evolution [4-10]. However, three-dimensional feather morphology and evolutionary patterns remain difficult to interpret, due to compression in sedimentary rocks [9, 11]. Recent discoveries in Cretaceous amber from Canada, France, Japan, Lebanon, Myanmar, and the United States [12-18] reveal much finer levels of structural detail, but taxonomic placement is uncertain because plumage is rarely associated with identifiable skeletal material [14]. Here we describe the feathered tail of a non-avialan theropod preserved in mid-Cretaceous (∼99 Ma) amber from Kachin State, Myanmar [17], with plumage structure that directly informs the evolutionary developmental pathway of feathers. This specimen provides an opportunity to document pristine feathers in direct association with a putative juvenile coelurosaur, preserving fine morphological details, including the spatial arrangement of follicles and feathers on the body, and micrometer-scale features of the plumage. Many feathers exhibit a short, slender rachis with alternating barbs and a uniform series of contiguous barbules, supporting the developmental hypothesis that barbs already possessed barbules when they fused to form the rachis [19]. Beneath the feathers, carbonized soft tissues offer a glimpse of preservational potential and history for the inclusion; abundant Fe(2+) suggests that vestiges of primary hemoglobin and ferritin remain trapped within the tail. The new finding highlights the unique preservation potential of amber for understanding the morphology and evolution of coelurosaurian integumentary structures.
1. Body feathers ensure both waterproofing and insulation in waterbirds, but how natural variation in the morphological properties of these appendages relates to environmental constraints remains largely unexplored. Here, we test how habitat and thermal condition affect the morphology of body feathers using a phylogenetic comparative analysis of five structural traits [i.e., total feather length, the lengths of the pennaceous (distal) and plumulaceous (proximal) sections, barb density, and pennaceous barbule density] from a sample of 194 European bird species.
2. Body feather total length is shorter in aquatic than in terrestrial birds, and this difference between groups is due to the shorter plumulaceous feather section in aquatic birds. Indeed, a reduced plumulaceous section in feather length probably reflects the need to limit air trapped in the plumage to adjust the buoyancy of aquatic birds. In contrast, the high pennaceous barbule density of aquatic birds compared to their terrestrial counterparts reflects water resistance of the plumage in contact with water.
3. Our results show that birds living in environments with low ambient temperature have long plumulaceous feather lengths, low barb density, and low pennaceous barbule density. Data also suggest that plumage probably has limited function in reducing the heat absorption of species living in hot environments.
4. Our results have broad implications for understanding the suite of selection pressures driving the evolution of body feather functional morphology. It remains to be tested, however, how other feather traits, such as the density of plumage (feathers per unit area) and the relative number of different feather types, for example downy feathers, are distributed amongst birds with different water resistance and thermoinsulative needs.
Our knowledge of Cretaceous plumage is limited by the fossil record itself: compression fossils surrounding skeletons lack the finest morphological details and seldom preserve visible traces of colour, while discoveries in amber have been disassociated from their source animals. Here we report the osteology, plumage and pterylosis of two exceptionally preserved theropod wings from Burmese amber, with vestiges of soft tissues. The extremely small size and osteological development of the wings, combined with their digit proportions, strongly suggests that the remains represent precocial hatchlings of enantiornithine birds. These specimens demonstrate that the plumage types associated with modern birds were present within single individuals of Enantiornithes by the Cenomanian (99 million years ago), providing insights into plumage arrangement and microstructure alongside immature skeletal remains. This finding brings new detail to our understanding of infrequently preserved juveniles, including the first concrete examples of follicles, feather tracts and apteria in Cretaceous avialans.
Birds can be classified into altricial and precocial. The hatchlings of altricial birds are almost naked, whereas those of
precocial birds are covered with natal down. This regulatory divergence is thought to reflect environmental adaptation, but
the molecular basis of the divergence is unclear. To address this issue, we chose the altricial zebra finch and the precocial
chicken as the model animals. We noted that zebra finch hatchlings show natal down growth suppressed anterior dorsal (AD)
skin but partially down-covered posterior dorsal (PD) skin. Comparing the transcriptomes of AD and PD skins, we found that
the feather growth promoter SHH (sonic hedgehog) was expressed higher in PD skin than in AD skin. Moreover, the data suggested that the FGF/MAPK signaling pathway is involved
in natal down growth suppression and that FGF16 (fibroblast growth factor 16) is a candidate upstream signaling suppressor. Ectopic expression of FGF16 on chicken leg skin showed downregulation of SHH, upregulation of the feather growth suppressor FGF10, and suppression of feather bud elongation, similar to the phenotype found in zebra finch embryonic AD skin. Therefore, we
propose that FGF16 related signals suppress natal down elongation and cause the naked AD skin in zebra finch. Our study provides insights into
the regulatory divergence in natal down formation between precocial and altricial birds.
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Antarctic penguins survive some of the harshest conditions on the planet. Emperor penguins breed on the sea ice where temperatures drop below 2408C and forage in21.88Cwaters. Their ability to maintain 388C body temperature in these conditions is due in large part to their feathered coat. Penguins have been reported to have the highest contour feather density of any bird, and both filoplumes and plumules (downy feathers) are reported absent in penguins. In studies modelling the heat transfer properties and the potential biomimetic applications of penguin plumage design, the insulative properties of penguin plumage have been attributed to the single afterfeather attached to contour feathers. This attribution of the afterfeather as the sole insulation component has been repeated in subsequent studies. Our results demonstrate the presence of both plumules and filoplumes in the penguin body plumage. The downy plumules are four times denser than afterfeathers and play a key, previously overlooked role in penguin survival. Our study also does not support the report that emperor penguins have the highest contour feather density. © 2015, The Author(s) Published by the Royal Society. All rights reserved.
Despite a wealth of fossils of Mesozoic birds revealing evidence of plumage and other soft-tissue structures, the epidermal and dermal anatomy of their wing's patagia remain largely unknown. We describe a distal forelimb of an enantiornithine bird from the Lower Cretaceous limestones of Las Hoyas, Spain, which reveals the overall morphology of the integument of the wing and other connective structures associated with the insertion of flight feathers. The integumentary anatomy, and myological and arthrological organization of the new fossil is remarkably similar to that of modern birds, in which a system of small muscles, tendons and ligaments attaches to the follicles of the remigial feathers and maintains the functional integrity of the wing during flight. The new fossil documents the oldest known occurrence of connective tissues in association with the flight feathers of birds. Furthermore, the presence of an essentially modern connective arrangement in the wing of enantiornithines supports the interpretation of these primitive birds as competent fliers.
This is the first re-appraisal in 50 years of concepts of development made in birds. This book is a case study in evolutionary diversification of life histories. Although birds have a rather uniform body plan and physiology, they exhibit marked variation in development type, parental care, and rate of growth. Altricial birds are fully dependent on their parents for warmth and nutrition and begin posthatching life in a more or less embryonic condition. At the other extreme, such superprecocial species as the megapodes are independent of all parental care from hatching, and the neonate, able to fly, resembles an adult bird. This book thus attempts to present an integrative perspective of organism biology, ecology, and evolution.
The discovery of Sinosauropteryx in 1996 marks the beginning of a new era in the research on the origin and early evolution of feathers. Subsequent discoveries of dinosaur fossils preserving feathers and feather-like integumentary appendages from both the Jurassic and Cretaceous deposits of China and other countries demonstrate a longer and more complex evolutionary history of feathers before the origin of birds than was previously thought. Currently, there are still many issues that continue to be debated or remain unresolved, such as at what point in phylogeny the first feathers originated (e.g., at the base of Avemetatarsalia vs. within Theropoda), how major microstructural features of feathers (e.g., barbules) evolved, whether early feathers developed ontogenetically in the same way as modern feathers, whether major features of feathers appeared incrementally in evolution or some appeared simultaneously (e.g., whether the feather follicle is a starting point for major feather features or these features appeared gradually), and what the primary functions of various early feathers were (e.g., display vs. insulation function for simple filamentous feathers and display vs. aerodynamic function for vaned feathers), among others. While insights from other fields such as developmental biology will help to address these various controversial issues pertaining to feather evolution, and a multidisciplinary approach is necessary to tackle and provide the full story of feather evolution, fossil specimens will continue to provide key data for the reconstruction and documentation of the evolutionary history of feathers, including evolutionary experiments and forms that no longer occur in nature.
Exceptionally preserved Mesozoic feathered dinosaur fossils (including birds) are famous, but recognized from only very few localities worldwide, and are especially rare in the Southern Hemisphere. Here we report an assemblage of non-avian and avian dinosaur feathers from an Early Cretaceous polar (around 70°S) environment in what is now southeastern Australia. The recovered remains incorporate small (10–30 mm long) basal paravian-like tufted body feathers, open-vaned contour feathers, and asymmetrical bird-like wing feathers that possess high-angled barbs with possible remnants of barbicels — amongst the geologically oldest observed to date. Such morphological diversity augments scant skeletal evidence for a range of insulated non-avian theropods and birds inhabiting extreme southern high-latitude settings during the Mesozoic. Although some of these fossil feathers exhibit what may be residual patterning, most are uniformly toned and preserve rod-shaped microbodies, as well as densely-packed microbody imprints on the barbules that are structurally consistent with eumelanosomes. Geochemical analysis detected no identifiable residual biomolecules, which we suspect were lost via hydrolysis and oxidization during diagenesis and weathering. Nevertheless, an originally dark pigmentation can be reasonably inferred from these melanic traces, which like the coloured feathers of modern birds, might have facilitated crypsis, visual communication and/or thermoregulation in a cold polar habitat.
The Palaeognathae comprise the flightless ratites and the volant tinamous, and together with the Neognathae constitute the extant members of class Aves. It is commonly believed that Palaeognathae originated in Gondwana since most of the living species are found in the Southern Hemisphere [1–3]. However, this hypothesis has been questioned because the fossil paleognaths are mostly from the Northern Hemisphere in their earliest time (Paleocene) and possessed many putative ancestral characters [4]. Uncertainties regarding the origin and evolution of Palaeognathae stem from the difficulty in estimating their divergence times [1, 2] and their remarkable morphological convergence. Here, we recovered nuclear genome fragments from extinct elephant birds, which enabled us to reconstruct a reliable phylogenomic time tree for the Palaeognathae. Based on the tree, we identified homoplasies in morphological traits of paleognaths and reconstructed their morphology-based phylogeny including fossil species without molecular data. In contrast to the prevailing theories, the fossil paleognaths from the Northern Hemisphere were placed as the basal lineages. Combined with our stable divergence time estimates that enabled a valid argument regarding the correlation with geological events, we propose a new evolutionary scenario that contradicts the traditional view. The ancestral Palaeognathae were volant, as estimated from their molecular evolutionary rates, and originated during the Late Cretaceous in the Northern Hemisphere. They migrated to the Southern Hemisphere and speciated explosively around the Cretaceous-Paleogene boundary. They then extended their distribution to the Gondwana-derived landmasses, such as New Zealand and Madagascar, by overseas dispersal. Gigantism subsequently occurred independently on each landmass.
Avian integument is thin, elastic, and loosely attached to the body, giving birds the freedom of movement needed for flight, Its epidermis is both keratinized and lipogenic, and the skin as a whole acts as a sebaceous secretory organ. The skin is covered by feathers over most of the body, but many birds show colored bare skin or integumentary outgrowths on the head and neck, Heavily cornified epidermis covers the beak, claws, spurs, and the scales on the legs and feet. These structures (except the back of the leg and underside of the foot) contain beta-keratin like that in reptilan scales. Most birds have sebaceous secretory glands at the base of the tail and in the ear canals, Feathers are the most numerous, elaborate, and diverse of avian integunentary derivatives. Their diversity is due to the possibilities inherent in their basic plan of a shaft with two orders of branches and the use of modified beta-keratin as a strong, light, and plastic building material. The evolution of feathers in birds has been accompanied by the development of complex systems for producing colors and patterns, the innovations of feather arrangement and follicles with their musculature and innervation, and the process and control of molting.
In colour polymorphic species morphs are considered to be adaptations to different environments, where they have evolved and are maintained because of their differential sensitivity to the environment. In cold environments the plumage insulation capacity is essential for survival and it has been proposed that plumage colour is associated with feather structure and thereby the insulation capacity of the plumage. We studied the structure of contour feathers in the colour polymorphic tawny owl (Strix aluco). A previous study of tawny owls in the same population has found strong selection against the brown morph in cold and snowy winters whereas this selection pressure is absent in mild winters. We predicted that grey morphs have a denser and more insulative plumage, enabling them to survive better in cold climate compared to brown ones. The insulative plumulaceous part of the dorsal contour feathers was larger and the fine structure of the plumulaceous part of the feather was denser in grey tawny owls than in brown ones. In the ventral contour feathers the plumulaceous part of the feather was denser in females than in males and in older birds without any differences between morphs. Our study suggests that insulative microscopical feather structures differ between colour morphs and we propose that feather structure may be a trait associated with morph-specific survival in cold environments.This article is protected by copyright. All rights reserved.
Numerical cladistic analysis of 73 cranial and postcranial characters has resulted in a highly corroborated hypothesis describing the phylogenetic pattern of early avian evolution. Using “non-avian theropod” dinosaurs as a comparative outgroup and root for the tree, the analysis confirmed Archaeopteryx to be the sister-group of all remaining avian taxa, or Ornithurae. This latter taxon is subdivided into two lineages, the Hesperornithiformes and the Carinatae. The carinates, in turn, were also resolved into two sister-groups, the Ichthyornithiformes and the modern birds, or Neornithes. This paper provides morphological data corroborating the divergence of the two basal clades of the Neornithes: the Palaeognathae (tinamous and ratites) and Neognathae (all other modern birds). The phylogenetic relationships of four important Cretaceous taxa were also investigated, but these fossil taxa were too fragmentary to determine their phylogenetic position unambiguously. Alexornis and Ambiortus are both carinates, but their relationships cannot be resolved in greater detail. The relationships of the Enantiornithes may lie within the Carinatae or these two taxa may be sister-groups. Gobipteryx is a neornithine and possibly the sister-group of the Palaeognathae.
This analysis indicates that major patterns of morphological change took place at the time of origin of the ancestors of the Ornithurae and the Carinatae. Ornithurine innovations included major changes throughout the skeleton, whereas those of the carinates, while substantial, were primarily restricted to the pectoral girdle and forelimb. The phylogenetic results, in conjunction with the known ages of fossil taxa, indicate that the early lineages of birds very likely arose in the Jurassic. The early cladistic events within the neornithine lineage are also more ancient than generally recognized, and may well extend back to the early Cretaceous.