The evolutionary origin and diversification of feathers

Progress on the evolutionary origin and diversification of feathers has been hampered by conceptual problems and by the lack of plesiomorphic feather fossils. Recently, both of these limitations have been overcome by the proposal of the developmental theory of the origin of feathers, and the discovery of primitive feather fossils on nonavian theropod dinosaurs. The conceptual problems of previous theories of the origin of feathers are reviewed, and the alternative developmental theory is presented and discussed. The developmental theory proposes that feathers evolved through a series of evolutionary novelties in developmental mechanisms of the follicle and feather germ. The discovery of primitive and derived fossil feathers on a diversity of coelurosaurian theropod dinosaurs documents that feathers evolved and diversified in nonavian theropods before the origin of birds and before the origin of flight. The morphologies of these primitive feathers are congruent with the predictions of the developmental theory. Alternatives to the theropod origin of feathers are critique and rejected. Hypotheses for the initial function of feathers are reviewed. The aerodynamic theory of feather origins is falsified, but many other functions remain developmentally and phylogenetically plausible. Whatever their function, feathers evolved by selection for a follicle that would grow an emergent tubular appendage. Feathers are inherently tubular structures. The homology of feathers and scales is weakly supported. Feathers are composed of a suite of evolutionary novelties that evolved by the duplication, hierarchical organization, interaction, dissociation, and differentiation of morphological modules. The unique capacity for modular subdivision of the tubular feather follicle and germ has fostered the evolution of numerous innovations that characterize feathers. The evolution of feather keratin and the molecular basis of feather development are also discussed.     

中国中生代的鸟类:介绍及综述

最近十来年 ,中国辽宁发现的早白垩世的鸟类化石超过了世界上其它任何一个地区。中国的中生代鸟类化石代表了始祖鸟化石之后鸟类历史上第一次显著的分异。它们不仅包括了带有明显恐龙祖先特征的长尾的鸟类 ,而且还包括了许多进步或特化的种类 ,如早白垩世最大的鸟类 ,最原始的反鸟类 ,以及保存最好的、飞行结构和现生鸟类几乎一样的今鸟类。这些早期鸟类在诸如飞行、大小和食性等所反映的演化、形态和生态学特征等方面出现了重大的分异。具有长尾骨骼的原始基干鸟类热河鸟和驰龙类具有的相似性 ,进一步支持了鸟类起源于恐龙的学说。中国发现的早白垩世的鸟类以及树栖的恐龙化石还为鸟类飞行的树栖起源假说提供了十分重要的证据。“恐龙下树”的假说结合了鸟类起源于恐龙的学说和鸟类飞行的树栖起源学说 ,因此也得到了化石证据的支持。由于多种恐龙带有羽毛 ,因此羽毛不一定代表了恒温。恒温的鸟类可能到了早白垩世的进步鸟类中才开始出现     

The early evolution of feathers: fossil evidence from Cretaceous amber of France

The developmental stages of feathers are of major importance in the evolution of body covering and the origin of avian flight. Until now, there were significant gaps in knowledge of early morphologies in theoretical stages of feathers as well as in palaeontological material. Here we report fossil evidence of an intermediate and critical stage in the incremental evolution of feathers which has been predicted by developmental theories but hitherto undocumented by evidence from both the recent and the fossil records. Seven feathers have been found in an Early Cretaceous (Late Albian, ca 100 Myr) amber of western France, which display a flattened shaft composed by the still distinct and incompletely fused bases of the barbs forming two irregular vanes. Considering their remarkably primitive features, and since recent discoveries have yielded feathers of modern type in some derived theropod dinosaurs, the Albian feathers from France might have been derived either from an early bird or from a non-avian dinosaur.     

Cross-Testing Adaptive Hypotheses: Phylogenetic Analysis and the Origin of Bird Flight

Adaptive scenarios in evolutionary biology have always beenbased on incremental improvements through a series of adaptivestages. But they have often been justified by appeal to assumptionsof how natural selection must work or by appeal to optimalityarguments or notions of evolutionary process. Cladistic methodology,though it cannot logically falsify hypotheses of process, provideshypotheses of evolutionary pattern independent of other considerationsand so provides a useful test of consilience with genealogy.I illustrate the cross-test of hypotheses of the evolution ofseveral functions and adaptations related to the origin of birdflight with independently derived phylogenetic analysis. Consiliencedoes not support ideas that the close ancestors of birds werearboreal or evolved flight from the trees, nor that they werephysiologically intermediate between typical reptiles and livingbirds, nor that feathers evolved for flight. Rather, the ancestorsof birds were terrestrial, they were fast-growing, active animals,and the original functions of feathers were in insulation andcoloration.     

被视为恐龙的鸟

尽管于 2 0世纪 2 0年代在亚洲地表层首次发现长有羽毛的恐龙和著名的“龙骨突位点” ,关于鸟类起源的争论仍没有休止。来自化石的证据表明 ,鸟类在进化分支上应归于兽脚类的特殊分支。本文主要阐明完好无损的化石揭示的鸟和非鸟类恐龙的亲密关系和鸟类羽毛及鸟类出现以前的羽毛的起源证据 ,分析体型缩小对飞行进化的重要意义及从新的角度论述鸟类如何飞上了天     

The relative timing of the origin of flight and endothermy: evidence from the comparative biology of birds and mammals.

Evidence from the comparative biology of living birds and mammals is used to address the question ‘which came first, flight or endothermy?’. Birds and mammals have evolved different solutions to the problems of high energy flow demanded by endothermy. The heavy apparatus needed for processing food to allow the rapid assimilation of energy is housed in the head of mammals, but low down in the bird's body. The primitive inefficient tidal-flow system of ventilation is simply enlarged in mammals, but is replaced in birds by a lighter uni-flow system through air sacs and parabronchi. Birds avoid the weight problems associated with the mammalian systems of viviparity and lactation by nourishing their young with large quantities of yolk within the egg and an unprocessed diet after hatching. The apparent adaptedness for flight of the avian systems suggests that in the animals ancestral to birds the adaptations for high energy flow were constrained from the start by the need for aerodynamic stability, i.e. flight was initiated before endothermy. The implications of this conclusion for the origin of flight and feathers are discussed.     

Respiratory and reproductive paleophysiology of dinosaurs and early birds

In terms of their diversity and longevity, dinosaurs and birds were/are surely among the most successful of terrestrial vertebrates. Unfortunately, interpreting many aspects of the biology of dinosaurs and the earliest of the birds presents formidable challenges because they are known only from fossils. Nevertheless, a variety of attributes of these taxa can be inferred by identification of shared anatomical structures whose presence is causally linked to specialized functions in living reptiles, birds, and mammals. Studies such as these demonstrate that although dinosaurs and early birds were likely to have been homeothermic, the absence of nasal respiratory turbinates in these animals indicates that they were likely to have maintained reptile-like (ectothermic) metabolic rates during periods of rest or routine activity. Nevertheless, given the metabolic capacities of some extant reptiles during periods of elevated activity, early birds were probably capable of powered flight. Similarly, had, for example, theropod dinosaurs possessed aerobic metabolic capacities and habits equivalent to those of some large, modern tropical latitude lizards (e.g., Varanus), they may well have maintained significant home ranges and actively pursued and killed large prey. Additionally, this scenario of active, although ectothermic, theropod dinosaurs seems reinforced by the likely utilization of crocodilian-like, diaphragm breathing in this group. Finally, persistent in vivo burial of their nests and apparent lack of egg turning suggests that clutch incubation by dinosaurs was more reptile- than birdlike. Contrary to previous suggestions, there is little if any reliable evidence that some dinosaur young may have been helpless and nestbound (altricial) at hatching.     

L’origine et l’évolution des oiseaux : 35 années de progrès

The origin and evolution of birds: 35 years of progress. Birds are dinosaurs – specifically, small feathered and flighted theropod dinosaurs that probably originated in Laurasia during the Late Jurassic over 140 million years ago. They are most closely related to other small theropods such as dromaeosaurs and troodontids, terrestrial predators that were fleet-footed hunters. The origin of birds is a classic example of two kinds of macroevolution: the phylogenetic origin of the group, and the sequential assembly of adaptations such as flight that are indelibly associated with birds. These adaptations were not assembled all at once. Rather, a great many characteristics associated with birds and flight first appeared in non-avian dinosaurs, where they were used for many purposes other than flight. These included insulation, brooding, and probably display and species recognition. Birds diversified steadily but gradually after their origin, which is identified with the origin of flight (Archaeopteryx); forelimb and other flight-associated features evolved more rapidly than features associated with the posterior skeleton. The first birds grew more slowly than extant birds do, and more like other small Mesozoic dinosaurs; like them, they probably matured sexually well before they completed their active skeletal growth. The origin of flight is not a problem of “trees down” or “ground up,” but rather an examination of the order in which diagnostic flight characters evolved, and what each stage can reveal about the functions and habits of bird outgroups at those evolutionary junctures.     

Origin of feathered flight

The origin of flight in birds and theropod dinosaurs is a many-sided and debatable problem. We develop a new approach to the resolution of this problem, combining terrestrial and arboreal hypotheses of the origin of flight. The bipedalism was a key adaptation for the development of flight in both birds and theropods. The bipedalism dismissed the forelimbs from the supporting function and promoted transformation into wings. For the development of true flapping avian flight, a key role was played by the initial universal anisodactylous foot of birds. This foot pattern provided a firm support on both land and trees. Theropod dinosaurs, archaeopteryxes, and some other early feathered creatures had a pamprodactylous foot and, hence, they developed only gliding descent. Early birds descended by flattering parachuting with the use of incipient wings; this gave rise to true flight. Among terrestrial vertebrates, only bats, pterosaurians, and birds developed true flapping flight, although they followed different morphofunctional pathways when solving this task. However, it remains uncertain what initiated the adaptation of the three groups for the air locomotion. Nevertheless, the past decade has provided unexpectedly abundant paleontological data, which facilitate the resolution of this question with reference to birds.     

Do feathered dinosaurs exist? Testing the hypothesis on neontological and paleontological evidence

The origin of birds and avian flight from within the archosaurian radiation has been among the most contentious issues in paleobiology. Although there is general agreement that birds are related to theropod dinosaurs at some level, debate centers on whether birds are derived directly from highly derived theropods, the current dogma, or from an earlier common ancestor lacking suites of derived anatomical characters. Recent discoveries from the Early Cretaceous of China have highlighted the debate, with claims of the discovery of all stages of feather evolution and ancestral birds (theropod dinosaurs), although the deposits are at least 25 million years younger than those containing the earliest known bird Archaeopteryx. In the first part of the study we examine the fossil evidence relating to alleged feather progenitors, commonly referred to as protofeathers, in these putative ancestors of birds. Our findings show no evidence for the existence of protofeathers and consequently no evidence in support of the follicular theory of the morphogenesis of the feather. Rather, based on histological studies of the integument of modern reptiles, which show complex patterns of the collagen fibers of the dermis, we conclude that "protofeathers" are probably the remains of collagenous fiber "meshworks" that reinforced the dinosaur integument. These "meshworks" of the skin frequently formed aberrant patterns resembling feathers as a consequence of decomposition. Our findings also draw support from new paleontological evidence. We describe integumental structures, very similar to "protofeathers," preserved within the rib area of a Psittacosaurus specimen from Nanjing, China, an ornithopod dinosaur unconnected with the ancestry of birds. These integumental structures show a strong resemblance to the collagenous fiber systems in the dermis of many animals. We also report the presence of scales in the forearm of the theropod ornithomimid (bird mimic) dinosaur, Pelecanimimus, from Spain. In the second part of the study we examine evidence relating to the most critical character thought to link birds to derived theropods, a tridactyl hand composed of digits 1-2-3. We maintain the evidence supports interpretation of bird wing digit identity as 2,3,4, which appears different from that in theropod dinosaurs. The phylogenetic significance of Chinese microraptors is also discussed, with respect to bird origins and flight origins. We suggest that a possible solution to the disparate data is that Aves plus bird-like maniraptoran theropods (e.g., microraptors and others) may be a separate clade, distinctive from the main lineage of Theropoda, a remnant of the early avian radiation, exhibiting all stages of flight and flightlessness.     

Mesozoic birds of China-a synoptic review

A synoptic review of the discoveries and studies of Chinese Mesozoic birds is provided in this paper.40Ar/39Ar dating of several bird-bearing deposits in the Jehol Group has established a geochronological framework for the study of the early avian radiation.Chinese Mesozoic birds had lasted for at least 11 Ma during about 131 Ma and 120 Ma (Barremian to Aptian)of the middle and late Early Cretaceous,respectively.In order to further evaluate the change of the avian diversity in the Jehol Biota,six new orders and families are erected based on known genera and species,which brings the total number of orders of Chinese Mesozoic birds to 15 and highlights a remarkable radiation ever since the first appearante of birds in the Late Jurassic.Chinese Early Cretaceous birds had experienced a significant differentiation in morphology,flight,diet and habitat.Further examination of the foot of Jeholornis suggests this bird might not have possessed a fully reversed hallux.However,the attachment of metatarsal Ⅰ to the medial side of metatarsal Ⅱ does not preclude trunk climbing,a pre-adaptation for well developed perching life of early birds.Arboreality had proved to be a key adaptation in the origin and early evolution of bird flight,and the adaptation to lakeshore environment had played an equally important role in the origin of omithurine birds and their near-modern flight skill.Many Chinese Early Cretaceous birds had preserved the direct evidence of their diet,showing that the most primitive birds were probably mainly insectivorous and that specialized herbivorous or carnivorous (e.g.,piscivorous)dietary adaptation had appeared only in later advanced forms.The only known Early Cretaceous bird embryo fossil has shown that precocial birds had occurred prior to altricial birds in avian history,and the size of the embryo and other analysis indicate it probably had a short incubation period.Leg feathers probably have a wide range of distribution in early birds,further suggesting that leg feathers had played a key role in the beginning stage of the flight of birds.Finally,the Early Cretaceous avian radiation can be better understood against the background of their unique ecosystem.The advantage of birds in the competitions with other vertebrate groups such as pterosaurs had probably not only resulted in the rapid differentiation and radiation of birds but also the worldwide spreading of pterosaurs and other vertebrates from East Asia in the Early Cretaceous.     

Linking the molecular evolution of avian beta (β) keratins to the evolution of feathers

Feathers of today's birds are constructed of beta (β)-keratins, structural proteins of the epidermis that are found solely in reptiles and birds. Discoveries of "feathered dinosaurs" continue to stimulate interest in the evolutionary origin of feathers, but few studies have attempted to link the molecular evolution of their major structural proteins (β-keratins) to the appearance of feathers in the fossil record. Using molecular dating methods, we show that before the appearance of Anchiornis (～155 Million years ago (Ma)) the basal β-keratins of birds began diverging from their archosaurian ancestor ～216?Ma. However, the subfamily of feather β-keratins, as found in living birds, did not begin diverging until ～143?Ma. Thus, the pennaceous feathers on Anchiornis, while being constructed of avian β-keratins, most likely did not contain the feather β-keratins found in the feathers of modern birds. Our results demonstrate that the evolutionary origin of feathers does not coincide with the molecular evolution of the feather β-keratins found in modern birds. More likely, during the Late Jurassic, the epidermal structures that appeared on organisms in the lineage leading to birds, including early forms of feathers, were constructed of avian β-keratins other than those found in the feathers of modern birds. Recent biophysical studies of the β-keratins in feathers support the view that the appearance of the subfamily of feather β-keratins altered the biophysical nature of the feather establishing its role in powered flight.     

The colour of fossil feathers

Feathers are complex integumentary appendages of birds and some other theropod dinosaurs. They are frequently coloured and function in camouflage and display. Previous investigations have concluded that fossil feathers are preserved as carbonized traces composed of feather-degrading bacteria. Here, an investigation of a colour-banded feather from the Lower Cretaceous Crato Formation of Brazil revealed that the dark bands are preserved as elongate, oblate carbonaceous bodies 1-2mum long, whereas the light bands retain only relief traces on the rock matrix. Energy dispersive X-ray analysis showed that the dark bands preserve a substantial amount of carbon, whereas the light bands show no carbon residue. Comparison of these oblate fossil bodies with the structure of black feathers from a living bird indicates that they are the eumelanin-containing melanosomes. We conclude that most fossil feathers are preserved as melanosomes, and that the distribution of these structures in fossil feathers can preserve the colour pattern in the original feather. The discovery of preserved melanosomes opens up the possibility of interpreting the colour of extinct birds and other dinosaurs.     

Dinosaur protofeathers: pushing back the origin of feathers into the Middle Triassic?

Reports of primordial feathers (protofeathers) in dinosaurs have received widespread interest. Recently, it was proposed that a novel protofeather in the theropod dinosaur Beipiaosaurus completes the transitional series in the evolution of the feather and provides the first evidence of filamentous feathers as display in nonavian theropods. A more far-reaching evolutionary ramification is the claim that these structures push the origin of monofilamentous integumentary structures into the Middle Triassic or earlier. I discuss problems with the analyses within the broader context of studies concerning the hypothesis of protofeathers, and show that affinity between the integumentary structures in Beipiaosaurus and feathers is improbable. The scientific methodology is questioned by its failure to make phenomena perceivable by objective means, by questionable rationalizations in critical issues, and by lack of consideration of exceptions to the postulated thesis. The notion that primordial feathers occurred in a clade more inclusive than the Coelurosauria and that it is supported by the presence of integumental structures in Psittacosaurus is analyzed and rejected.     

Gravity-defying Behaviors: Identifying Models for Protoaves

Most current phylogenetic hypotheses based upon cladistic methodologyassert that birds are the direct descendants of derived maniraptorantheropod dinosaurs, and that the origin of avian flight necessarilydeveloped within a terrestrial context (i.e., from the "groundup"). Most theoretical aerodynamic and energetic models or chronologicallyappropriate fossil data do not support these hypotheses forthe evolution of powered flight. The more traditional modelfor the origin of flight derives birds from among small arborealearly Mesozoic archosaurs ("thecodonts"). According to thismodel, protoavian ancestors developed flight in the trees viaa series of intermediate stages, such as leaping, parachuting,gliding, and flapping. This model benefits from the assemblageof living and extinct arboreal vertebrates that engage in analogousnon-powered aerial activities using elevation as a source ofgravitational energy. Recent reports of "feathered theropods"notwithstanding, the evolution of birds from any known groupof maniraptoran theropods remains equivocal.     

The evolution of endothermy in terrestrial vertebrates: Who? When? Why?

Avian and mammalian endothermy results from elevated rates of resting, or routine, metabolism and enables these animals to maintain high and stable body temperatures in the face of variable ambient temperatures. Endothermy is also associated with enhanced stamina and elevated capacity for aerobic metabolism during periods of prolonged activity. These attributes of birds and mammals have greatly contributed to their widespread distribution and ecological success. Unfortunately, since few anatomical/physiological attributes linked to endothermy are preserved in fossils, the origin of endothermy among the ancestors of mammals and birds has long remained obscure. Two recent approaches provide new insight into the metabolic physiology of extinct forms. One addresses chronic (resting) metabolic rates and emphasizes the presence of nasal respiratory turbinates in virtually all extant endotherms. These structures are associated with recovery of respiratory heat and moisture in animals with high resting metabolic rates. The fossil record of nonmammalian synapsids suggests that at least two Late Permian lineages possessed incipient respiratory turbinates. In contrast, these structures appear to have been absent in dinosaurs and nonornithurine birds. Instead, nasal morphology suggests that in the avian lineage, respiratory turbinates first appeared in Cretaceous ornithurines. The other approach addresses the capacity for maximal aerobic activity and examines lung structure and ventilatory mechanisms. There is no positive evidence to support the reconstruction of a derived, avian-like parabronchial lung/air sac system in dinosaurs or nonornithurine birds. Dinosaur lungs were likely heterogenous, multicameral septate lungs with conventional, tidal ventilation, although evidence from some theropods suggests that at least this group may have had a hepatic piston mechanism of supplementary lung ventilation. This suggests that dinosaurs and nonornithurine birds generally lacked the capacity for high, avian-like levels of sustained activity, although the aerobic capacity of theropods may have exceeded that of extant ectotherms. The avian parabronchial lung/air sac system appears to be an attribute limited to ornithurine birds.     

Mesozoic birds of China—a synoptic review

A synoptic review of the discoveries and studies of Chinese Mesozoic birds is provided in this paper. ⁴⁰Ar/³⁹Ar dating of several bird-bearing deposits in the Jehol Group has established a geochronological framework for the study of the early avian radiation. Chinese Mesozoic birds had lasted for at least 11 Ma during about 131 Ma and 120 Ma (Barremian to Aptian) of the middle and late Early Cretaceous, respectively. In order to further evaluate the change of the avian diversity in the Jehol Biota, six new orders and families are erected based on known genera and species, which brings the total number of orders of Chinese Mesozoic birds to 15 and highlights a remarkable radiation ever since the first appearance of birds in the Late Jurassic. Chinese Early Cretaceous birds had experienced a significant differentiation in morphology, flight, diet and habitat. Further examination of the foot of Jeholornis suggests this bird might not have possessed a fully reversed hallux. However, the attachment of metatarsal I to the medial side of metatarsal II does not preclude trunk climbing, a pre-adaptation for well developed perching life of early birds. Arboreality had proved to be a key adaptation in the origin and early evolution of bird flight, and the adaptation to lakeshore environment had played an equally important role in the origin of ornithurine birds and their near-modern flight skill. Many Chinese Early Cretaceous birds had preserved the direct evidence of their diet, showing that the most primitive birds were probably mainly insectivorous and that specialized herbivorous or carnivorous (e.g., piscivorous) dietary adaptation had appeared only in later advanced forms. The only known Early Cretaceous bird embryo fossil has shown that precocial birds had occurred prior to altricial birds in avian history, and the size of the embryo and other analysis indicate it probably had a short incubation period. Leg feathers probably have a wide range of distribution in early birds, further suggesting that leg feathers had played a key role in the beginning stage of the flight of birds. Finally, the Early Cretaceous avian radiation can be better understood against the background of their unique ecosystem. The advantage of birds in the competitions with other vertebrate groups such as pterosaurs had probably not only resulted in the rapid differentiation and radiation of birds but also the worldwide spreading of pterosaurs and other vertebrates from East Asia in the Early Cretaceous. Selected from Vertebrata PalAsiatica 2006, 44 (1): 74–98     

Testing co-evolutionary hypotheses over geological timescales: interactions between Mesozoic non-avian dinosaurs and cycads

The significance of co‐evolution over ecological timescales is well established, yet it remains unclear to what extent co‐evolutionary processes contribute to driving large‐scale evolutionary and ecological changes over geological timescales. Some of the most intriguing and pervasive long‐term co‐evolutionary hypotheses relate to proposed interactions between herbivorous non‐avian dinosaurs and Mesozoic plants, including cycads. Dinosaurs have been proposed as key dispersers of cycad seeds during the Mesozoic, and temporal variation in cycad diversity and abundance has been linked to dinosaur faunal changes. Here we assess the evidence for proposed hypotheses of trophic and evolutionary interactions between these two groups using diversity analyses, a new database of Cretaceous dinosaur and plant co‐occurrence data, and a geographical information system (GIS) as a visualisation tool. Phylogenetic evidence suggests that the origins of several key biological properties of cycads (e.g. toxins, bright‐coloured seeds) likely predated the origin of dinosaurs. Direct evidence of dinosaur–cycad interactions is lacking, but evidence from extant ecosystems suggests that dinosaurs may plausibly have acted as seed dispersers for cycads, although it is likely that other vertebrate groups (e.g. birds, early mammals) also played a role. Although the Late Triassic radiations of dinosaurs and cycads appear to have been approximately contemporaneous, few significant changes in dinosaur faunas coincide with the late Early Cretaceous cycad decline. No significant spatiotemporal associations between particular dinosaur groups and cycads can be identified – GIS visualisation reveals disparities between the spatiotemporal distributions of some dinosaur groups (e.g. sauropodomorphs) and cycads that are inconsistent with co‐evolutionary hypotheses. The available data provide no unequivocal support for any of the proposed co‐evolutionary interactions between cycads and herbivorous dinosaurs – diffuse co‐evolutionary scenarios that are proposed to operate over geological timescales are plausible, but such hypotheses need to be firmly grounded on direct evidence of interaction and may be difficult to support given the patchiness of the fossil record.     

鸟类起源于基干的初龙类

中亚三叠纪和中国早白垩世的新的重要化石发现或许可以帮助解决有关鸟类起源的争议。鸟类的飞行可能起源于三叠纪一些小型的、四脚滑翔的初龙类。羽毛的起源最初是为了滑翔而不是保温。“手盗龙类”实际上起源于鸟类 ,并至少发育了初级飞羽 ,具备滑翔的能力     

The species recognition hypothesis explains exaggerated structures in non-avialan dinosaurs better than sexual selection does

Several hypotheses have been proposed to explain “bizarre structures” in dinosaurs and other extinct animals (e.g., mechanical function and several kinds of intra- and interspecific display). Recent evidence and tests for species recognition as a possible driver of these structures have been proposed, in particular as an alternative to traditional hypotheses of function and sexual selection, which have fallen short. Advocates of sexual selection and mechanical function have advanced untested hypotheses claiming that species recognition cannot be an important process in evolution. We address these claims and show that they are based on misreading of the evidence and of previous literature. We also acknowledge that there have been historically differing definitions of sexual selection, which have greatly impeded understanding of the whole phenomenon of mate attraction and choice. Particularly in fossil animals, it is impossible to accept any hypothesis as the “default” that does not require evidence or testing to establish it.