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61-486: Strisores ( / s t r aɪ ˈ s oʊ r iː z / stry- SOH -reez ), sometimes called nightbirds , is a clade of birds that includes the living families and orders Caprimulgidae (nightjars, nighthawks and allies), Nyctibiidae (potoos), Steatornithidae ( oilbirds ), Podargidae ( frogmouths ), Apodiformes (swifts and hummingbirds), as well as the Aegotheliformes (owlet-nightjars) whose distinctness

122-531: A Late Paleocene or Early Eocene genus of North America, cannot be assigned to any one strisore lineage with certainty but appears to be some ancestral form. Over some 20 million years, throughout the Eocene , the present-day diversity (as well as some entirely extinct lineages) slowly unfolds. By mid- Oligocene , some 30 million years ago, the crown lineages are present and adapting to their present-day ecological niches . These Paleogene birds strongly suggest that

183-525: A clade (from Ancient Greek κλάδος (kládos)  'branch'), also known as a monophyletic group or natural group , is a grouping of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree . In the taxonomical literature, sometimes the Latin form cladus (plural cladi ) is used rather than the English form. Clades are

244-479: A "ladder", with supposedly more "advanced" organisms at the top. Taxonomists have increasingly worked to make the taxonomic system reflect evolution. When it comes to naming , this principle is not always compatible with the traditional rank-based nomenclature (in which only taxa associated with a rank can be named) because not enough ranks exist to name a long series of nested clades. For these and other reasons, phylogenetic nomenclature has been developed; it

305-623: A clade can be described based on two different reference points, crown age and stem age. The crown age of a clade refers to the age of the most recent common ancestor of all of the species in the clade. The stem age of a clade refers to the time that the ancestral lineage of the clade diverged from its sister clade. A clade's stem age is either the same as or older than its crown age. Ages of clades cannot be directly observed. They are inferred, either from stratigraphy of fossils , or from molecular clock estimates. Viruses , and particularly RNA viruses form clades. These are useful in tracking

366-436: A complicated situation where some researchers currently use the resurrected name Strisores in a new sense, others expand the order Caprimulgiformes to include the 'traditional' apodiform families, whereas others use the superordinal name Caprimulgimorphae Cracraft, 2013, raising the 'traditional' caprimulgiform families to the rank of order. Proposed phylogenetic definitions of Strisores and Caprimulgimorphae treat Strisores as

427-622: A family within Apodiformes . Traditionally, Caprimulgiformes were regarded, on morphological grounds, as being midway between the owls (Strigiformes) and the swifts . Like the owls, they are nocturnal hunters with a highly developed sense of sight, and like the swifts they are excellent flyers with small, weak legs. At one time or another, they have been allied with owls, swifts, kingfishers , hoopoes , mousebirds , hornbills , rollers , bee-eaters , woodpeckers , trogons and hummingbirds . A close relationship to owls can be rejected since there

488-428: A host of prefixes have been defined to describe various branches of the phylogenetic tree relative to extant organisms. A pan-group or total group is the crown group and all organisms more closely related to it than to any other extant organisms. In a tree analogy, it is the crown group and all branches back to (but not including) the split with the closest branch to have living members. The Pan-Aves thus contain

549-422: A revised taxonomy based on a concept strongly resembling clades, although the term clade itself would not be coined until 1957 by his grandson, Julian Huxley . German biologist Emil Hans Willi Hennig (1913–1976) is considered to be the founder of cladistics . He proposed a classification system that represented repeated branchings of the family tree, as opposed to the previous systems, which put organisms on

610-432: A stem group allows the order of these acquisitions to be established, and thus the ecological and functional setting of the evolution of the major features of the group in question. Stem groups thus offer a route to integrate unique palaeontological data into questions of the evolution of living organisms. Furthermore, they show that fossils that were considered to lie in their own separate group because they did not show all

671-429: A suffix added should be e.g. "dracohortian". A clade is by definition monophyletic , meaning that it contains one ancestor which can be an organism, a population, or a species and all its descendants. The ancestor can be known or unknown; any and all members of a clade can be extant or extinct. The science that tries to reconstruct phylogenetic trees and thus discover clades is called phylogenetics or cladistics ,

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732-499: Is also used with a similar meaning in other fields besides biology, such as historical linguistics ; see Cladistics § In disciplines other than biology . The term "clade" was coined in 1957 by the biologist Julian Huxley to refer to the result of cladogenesis , the evolutionary splitting of a parent species into two distinct species, a concept Huxley borrowed from Bernhard Rensch . Many commonly named groups – rodents and insects , for example – are clades because, in each case,

793-471: Is in turn included in the mammal, vertebrate and animal clades. The idea of a clade did not exist in pre- Darwinian Linnaean taxonomy , which was based by necessity only on internal or external morphological similarities between organisms. Many of the better known animal groups in Linnaeus's original Systema Naturae (mostly vertebrate groups) do represent clades. The phenomenon of convergent evolution

854-618: Is no consensus phylogeny. Stem arthropods constitute a group that has seen attention in connection with the Burgess Shale fauna. Several of the finds , including the enigmatic Opabinia and Anomalocaris have some, though not all, features associated with arthropods , and are thus considered stem arthropods. The sorting of the Burgess Shale fauna into various stem groups finally enabled phylogenetic sorting of this enigmatic assemblage and also allowed for identifying velvet worms as

915-515: Is responsible for many cases of misleading similarities in the morphology of groups that evolved from different lineages. With the increasing realization in the first half of the 19th century that species had changed and split through the ages, classification increasingly came to be seen as branches on the evolutionary tree of life . The publication of Darwin's theory of evolution in 1859 gave this view increasing weight. In 1876 Thomas Henry Huxley , an early advocate of evolutionary theory, proposed

976-425: Is shown below: † Archaeopteryx other extinct groups Neornithes (modern birds, some extinct like the dodo) In this diagram, the clade labelled "Neornithes" is the crown group of birds: it includes the most recent common ancestor of all living birds and its descendants, living or not. Although considered to be birds (i.e. members of the clade Aves), Archaeopteryx and other extinct groups are not included in

1037-489: Is still controversial. As an example, see the full current classification of Anas platyrhynchos (the mallard duck) with 40 clades from Eukaryota down by following this Wikispecies link and clicking on "Expand". The name of a clade is conventionally a plural, where the singular refers to each member individually. A unique exception is the reptile clade Dracohors , which was made by haplology from Latin "draco" and "cohors", i.e. "the dragon cohort "; its form with

1098-536: Is strong molecular evidence that owls are members of a clade, called Telluraves , that excludes Caprimulgiformes. Based on analysis of DNA sequence data – notably β- fibrinogen intron 7 – Fain and Houde considered the families of the Caprimulgiformes to be members of the proposed clade Metaves , which also includes the hoatzin , tropicbirds , sandgrouse , pigeons , kagu , sunbittern , mesites , flamingos , grebes and swifts and hummingbirds . Metaves

1159-453: Is thought by some to make the Cambrian explosion easier to understand without invoking unusual evolutionary mechanisms; however, application of the stem group concept does nothing to ameliorate the difficulties that phylogenetic telescoping poses to evolutionary theorists attempting to understand both macroevolutionary change and the abrupt character of the Cambrian explosion . Overemphasis on

1220-561: The ossa maxillaria separated by a large cleft, a mandible with short pars symphysialis , and rami mandibulae slender in their distal half. The taxonomy of this group of birds has a long and complicated history. Jean Cabanis originally coined the name Strisores in 1847 as an order encompassing a much broader group of birds subdivided into two 'tribes': the Macrochires ( hummingbirds , swifts , and nightjars , including oilbirds and potoos , but notably excluding frogmouths ) and

1281-684: The Amphibolae ( hoatzin , mousebirds , and turacos ). Hermann Burmeister later excluded the taxa in Cabanis' Amphibolae from Strisores, but added kingfishers and motmots . Subsequent authors used either definition according to their own judgement, with Baird following Cabanis', and Cooper following Burmeister's usage. In 1867, Thomas Henry Huxley proposed the name Cypselomorphae for hummingbirds, swifts, and nightjars (including owlet-nightjars and potoos), however, he considered frogmouths and oilbirds unrelated due to aspects of their skull morphology. In

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1342-520: The Early Eocene (though this is somewhat uncertain), seems to be a basal form that at times has been allied with the oilbird and the potoos, but cannot be assigned to either with certainty. In the consensus scenario, it would represent a record of the initial divergence of the three lineages. This agrees with fossils suggesting that the basal divergence of the owlet-nightjar and apodiform branch also occurred during that time. In addition, Eocypselus ,

1403-487: The crown group and Caprimulgimorphae as the total group . This allows both names to be valid, with similar but not identical meanings. Strisores has a well-represented fossil record, with fossils of most major strisorean lineages known from the Paleogene . Chen et al. (2019) included 14 fossil lineages in their analysis. Nonetheless, it supports the emerging consensus phylogeny well. The genus Paraprefica , probably from

1464-405: The dinosaurs and the pterosaurs . The last common ancestor of birds and crocodilians—the first crown group archosaur—was neither bird nor crocodilian and possessed none of the features unique to either. As the bird stem group evolved, distinctive bird features such as feathers and hollow bones appeared. Finally, at the base of the crown group, all traits common to extant birds were present. Under

1525-466: The last common ancestor of the crown group and their closest living relatives. It follows from the definition that all members of a stem group are extinct. The "stem group" is the most used and most important of the concepts linked to crown groups, as it offers a means to reify and name paraphyletic assemblages of fossils that otherwise do not fit into systematics based on living organisms. While often attributed to Jefferies (1979), Willmann (2003) traced

1586-426: The lungfish , our nearest relatives among the fishes. In addition to a series of lobe-finned fishes , they also include some of the early labyrinthodonts . Exactly what labyrinthodonts are in the stem group tetrapods rather than the corresponding crown group is uncertain, as the phylogeny of early tetrapods is not well understood. This example shows that crown and stem group definitions are of limited value when there

1647-454: The "Metaves" must originate quite some time before the Paleogene , and they reconciled this with the fossil record. While the relationships of cypselomorphs are a subject of ongoing debate, the phylogeny of the individual lineages is better resolved. Much of the remaining uncertainty regards minor details. Initial mtDNA cytochrome b sequence analysis agreed with earlier morphological and DNA-DNA hybridization studies insofar as that

1708-400: The "crown" and "stem" group terminology was coined by R. P. S. Jefferies in 1979. Though formulated in the 1970s, the term was not commonly used until its reintroduction in 2000 by Graham Budd and Sören Jensen . It is not necessary for a species to have living descendants in order for it to be included in the crown group. Extinct side branches on the family tree that are descended from

1769-434: The 1880s Anton Reichenow continued to use Strisores in a similar sense as Huxley's Cypselomorphae (this time also excluding the owlet-nightjars ), but by the late 19th Century, Strisores had fallen into disuse, and this remained the case through the 20th Century. By the early 21st century, analyses of anatomical morphology and molecular phylogenomics demonstrated that the order Caprimulgiformes as had been used for much of

1830-445: The 20th century (oilbirds, potoos, nightjars, frogmouths, and owlet-nightjars) is actually paraphyletic respective to Apodiformes (hummingbirds, swifts, and treeswifts ), with apodiform birds nesting deeply within caprimulgiformes and a sister taxon to the owlet-nightjars. The discovery has led to a challenge of reconciling a Linnean hierarchy with phylogenetic relationships while still maintaining nomenclatural stability, resulting in

1891-521: The Crocodilia branch. Basal branch names such as Avemetatarsalia are usually more obscure. However, not so advantageous are the facts that "Pan-Aves" and "Aves" are not the same group, the circumscription of the concept of "Pan-Aves" (synonymous with Avemetatarsalia) is only evident by examination of the above tree, and calling both groups "birds" is ambiguous. Stem mammals are those in the lineage leading to living mammals, together with side branches, from

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1952-514: The closest living relatives of arthropods. Stem priapulids are other early Cambrian to middle Cambrian faunas, appearing in Chengjiang to Burgess Shale. The genus Ottoia has more or less the same build as modern priapulids , but phylogenetic analysis indicates that it falls outside the crown group, making it a stem priapulid. The name plesion has a long history in biological systematics, and plesion group has acquired several meanings over

2013-582: The crown group, as they fall outside the Neornithes clade, being descended from an earlier ancestor. An alternative definition does not require any members of a crown group to be extant, only to have resulted from a "major cladogenesis event". The first definition forms the basis of this article. Often, the crown group is given the designation "crown-", to separate it from the group as commonly defined. Both birds and mammals are traditionally defined by their traits, and contain fossil members that lived before

2074-483: The diagnostic features of a living clade, can nevertheless be related to it by lying in its stem group. Such fossils have been of particular importance in considering the origins of the tetrapods , mammals , and animals . The application of the stem group concept also influenced the interpretation of the organisms of the Burgess shale . Their classification in stem groups to extant phyla, rather than in phyla of their own,

2135-940: The distribution of fossils, the Paleogene radiation seems to have originated in Asia , which at that time became a highly fragmented landscape as the Himalayas lifted up and the Turgai Strait started to disappear. Several fossil taxa are tentatively placed here as basal or incertae sedis Strisores contains the extant orders Aegotheliformes , Apodiformes (with families Apodidae , Hemiprocnidae , and Trochilidae ), Caprimulgiformes , Nyctibiiformes , Podargiformes , Steatornithiformes . Apodidae and Hemiprocnidae are grouped together as Apodi, Apodi and Trochilidae are grouped together as Apodiformes , and Apodiformes and Aegotheliformes are grouped together as Daedalornithes. The classification of

2196-509: The divergence of the lineage from the Sauropsida to the last common ancestor of the living mammals. This group includes the synapsids as well as mammaliaforms like the morganucodonts and the docodonts ; the latter groups have traditionally and anatomically been considered mammals even though they fall outside the crown group mammals. Stem tetrapods are the animals belonging to the lineage leading to tetrapods from their divergence from

2257-505: The extinct moa ) The crown group here is Neornithes , all modern bird lineages back to their last common ancestor. The closest living relatives of birds are crocodilians . If we follow the phylogenetic lineage leading to Neornithes to the left, the line itself and all side branches belong to the stem birds until the lineage merges with that of the crocodilians. In addition to non-crown group primitive birds like Archaeopteryx , Hesperornis and Confuciusornis , stem group birds include

2318-442: The frogmouths seem quite distinct among the remaining Caprimulgiformes, but their exact placement cannot be resolved based on osteological data alone. Even the study of Ericson et al. could not properly resolve the oilbird's and frogmouths' relationships beyond the fact that they are quite certainly well distinct. It robustly supported, however, the idea that the owlet-nightjars should be considered closer to Caprimulgiformes, unlike

2379-432: The full bifurcating phylogeny. Stem birds perhaps constitute the most cited example of a stem group, as the phylogeny of this group is fairly well known. The following cladogram, based on Benton (2005), illustrates the concept: Crocodilia Pterosauria Hadrosauridae Stegosauria Sauropoda Tyrannosauridae Archaeopteryx Neognathae (including the extinct dodo ) Paleognathae (including

2440-451: The fundamental unit of cladistics , a modern approach to taxonomy adopted by most biological fields. The common ancestor may be an individual, a population , or a species ( extinct or extant ). Clades are nested, one in another, as each branch in turn splits into smaller branches. These splits reflect evolutionary history as populations diverged and evolved independently. Clades are termed monophyletic (Greek: "one clan") groups. Over

2501-546: The group consists of a common ancestor with all its descendant branches. Rodents, for example, are a branch of mammals that split off after the end of the period when the clade Dinosauria stopped being the dominant terrestrial vertebrates 66 million years ago. The original population and all its descendants are a clade. The rodent clade corresponds to the order Rodentia, and insects to the class Insecta. These clades include smaller clades, such as chipmunk or ant , each of which consists of even smaller clades. The clade "rodent"

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2562-522: The last common ancestors of the living groups or, like the mammal Haldanodon , were not descended from that ancestor although they lived later. Crown-Aves and Crown-Mammalia therefore differ slightly in content from the common definition of Aves and Mammalia. This has caused some confusion in the literature. The cladistic idea of strictly using the topology of the phylogenetic tree to define groups necessitates other definitions than crown groups to adequately define commonly discussed fossil groups. Thus,

2623-590: The last few decades, the cladistic approach has revolutionized biological classification and revealed surprising evolutionary relationships among organisms. Increasingly, taxonomists try to avoid naming taxa that are not clades; that is, taxa that are not monophyletic . Some of the relationships between organisms that the molecular biology arm of cladistics has revealed include that fungi are closer relatives to animals than they are to plants, archaea are now considered different from bacteria , and multicellular organisms may have evolved from archaea. The term "clade"

2684-518: The latter term coined by Ernst Mayr (1965), derived from "clade". The results of phylogenetic/cladistic analyses are tree-shaped diagrams called cladograms ; they, and all their branches, are phylogenetic hypotheses. Three methods of defining clades are featured in phylogenetic nomenclature : node-, stem-, and apomorphy-based (see Phylogenetic nomenclature§Phylogenetic definitions of clade names for detailed definitions). The relationship between clades can be described in several ways: The age of

2745-628: The living birds and all (fossil) organisms more closely related to birds than to crocodilians (their closest living relatives). The phylogenetic lineage leading back from Neornithes to the point where it merges with the crocodilian lineage, along with all side branches, constitutes pan-birds. In addition to non-crown group primitive birds like Archaeopteryx , Hesperornis and Confuciusornis , therefore, pan-group birds would include all dinosaurs and pterosaurs as well as an assortment of non-crocodilian animals like Marasuchus . Pan-Mammalia consists of all mammals and their fossil ancestors back to

2806-549: The methodologically weaker studies of Mariaux & Braun (1996) and Fain and Houde (2004). Alternatively, Mayr's phylogenetic taxon Cypselomorphae might be placed at order rank and substitute the two present orders Caprimulgiformes and Apodiformes. Such a group would be fairly uninformative as regards its evolutionary history, as it has to include some very plesiomorphic and some extremely derived lineages (such as hummingbirds) to achieve monophyly. Reddy et al . (2017) included hummingbirds and swifts in Caprimulgiformes, preserving

2867-423: The monophyly of the order. The following cladogram follows the results of Mayr's (2002) phylogenetic study, which used a parsimony analysis of 25 morphological characters: Steatornithidae Podargidae Caprimulgidae Nyctibiidae Aegothelidae Trochilidae Hemiprocnidae Apodidae Subsequent molecular work has converged on two alternative topologies (topologies 1 and 2 below) that differ in

2928-442: The most recent common ancestor of living members will still be part of a crown group. For example, if we consider the crown-birds (i.e. all extant birds and the rest of the family tree back to their most recent common ancestor), extinct side branches like the dodo or great auk are still descended from the most recent common ancestor of all living birds , so fall within the bird crown group. One very simplified cladogram for birds

2989-464: The narrower one. Often, an (extinct) grouping is identified as belonging together. Later, it may be realized other (extant) groupings actually emerged within such grouping, rendering them a stem grouping. Cladistically , the new groups should then be added to the group, as paraphyletic groupings are not natural. In any case, stem groupings with living descendants should not be viewed as a cohesive group, but their tree should be further resolved to reveal

3050-416: The oilbird and the frogmouths seemed rather distinct. The other lineages appeared to form a clade , but this is now known to have been caused by methodological limitations. The Aegothelidae ( owlet-nightjars ) with about a dozen living species in one genus are apparently closer to the Apodiformes ; these and the Caprimulgiformes are closely related, being grouped together as Cypselomorphae . The oilbird and

3111-570: The order Caprimulgiformes identical to the clade Strisores. Authorities that favor the use of Strisores for this group (e.g., Yuri et al. 2013 and Chen et al. 2019) adopt a sensu stricto definition of the order, limiting to the family Caprimulgidae . They also elevate many (or even all) of the families traditionally placed in Caprimulgiformes to ordinal rank. This requires recognizing at least three additional orders: Nyctibiiformes , Steatornithiformes , and Podargiformes . Owlet-nightjars can be placed in their own order ( Aegotheliformes ) or viewed as

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3172-424: The origin of the stem group concept to Austrian systematist Othenio Abel (1914), and it was discussed and diagrammed in English as early as 1933 by A. S. Romer . Alternatively, the term "stem group" is sometimes used in a wider sense to cover any members of the traditional taxon falling outside the crown group. Permian synapsids like Dimetrodon or Anteosaurus are stem mammals in the wider sense but not in

3233-493: The phylogenetic split from the remaining amniotes (the Sauropsida ). Pan-Mammalia is thus an alternative name for Synapsida . A stem group is a paraphyletic assemblage composed of the members of a pan-group or total group, above, minus the crown group itself (and therefore minus all living members of the pan-group). This leaves primitive relatives of the crown groups , back along the phylogenetic line to (but not including)

3294-1207: The placement of the root. Although Braun et al. (2019) suggested that topology 1 was favored in large-scale analyses of non-coding data were analyzed and that topology 2 was favored in large-scale analyses of coding data (e.g., Prum et al . (2015)) subsequent analyses of datasets with many non-coding loci have also recovered topology 2. Thus, topology 2 should be viewed as the best-corroborated hypothesis at this time. Topology 1: phylogeny according to Reddy et al . (2017), which analyzed 54 nuclear loci (mostly introns ): Steatornithidae Nyctibiidae Caprimulgidae Podargidae Aegothelidae Hemiprocnidae Apodidae Trochilidae Topology 2: phylogeny according to Prum et al . (2015) (259 "anchored hybrid enrichment" loci, which are mostly coding exons), Chen et al. (2019) (combined analysis of 2289 ultra-conserved elements [UCEs] and 117 morphological characters and including fossil taxa), and White and Braun (2019) (based on analyses of multiple UCE datasets, ranging in size from 2289 to 4243 loci): Caprimulgidae Steatornithidae Nyctibiidae Podargidae Aegothelidae Hemiprocnidae Clade In biological phylogenetics ,

3355-596: The spread of viral infections . HIV , for example, has clades called subtypes, which vary in geographical prevalence. HIV subtype (clade) B, for example is predominant in Europe, the Americas and Japan, whereas subtype A is more common in east Africa. Total group The concept was developed by Willi Hennig , the formulator of phylogenetic systematics , as a way of classifying living organisms relative to their extinct relatives in his "Die Stammesgeschichte der Insekten", and

3416-587: The two main extant lineages of strisores separated about 60-55 mya ( Selandian - Thanetian ), and that some time around the Lutetian - Bartonian boundary, some 40 mya, the common ancestors of Nyctibiidae, Caprimulgidae and eared nightjars diverged from those of oilbird and frogmouths. The relationships of the Early Eocene Parvicuculus and Procuculus from the southern North Sea basin are unresolved, but they bear some similarities to strisores. By

3477-403: The various birds that make up the order has long been controversial and difficult, particularly in the case of the nightjars and the paraphyly of the traditional Caprimulgiformes in relation to " Apodiformes ", traditionally considered a separate order. The IUCN adopts the following classification of Order Caprimulgiformes, which follows recent phylogenetic studies: The IUCN definition renders

3538-597: The widely used total-group perspective, the Crocodylomorpha would become synonymous with the Crocodilia, and the Avemetatarsalia would become synonymous with the birds, and the above tree could be summarized as Crocodilia Birds An advantage of this approach is that declaring Theropoda to be birds (or Pan-aves ) is more specific than declaring it to be a member of the Archosauria, which would not exclude it from

3599-487: The years. One use is as "nearby group" (plesion means close to in Greek ), i.e. sister group to a given taxon , whether that group is a crown group or not. The term may also mean a group, possibly paraphyletic , defined by primitive traits (i.e. symplesiomorphies ). It is generally taken to mean a side branch splitting off earlier on the phylogenetic tree than the group in question. Placing fossils in their right order in

3660-734: Was also found by the expanded study of Ericson et al. (2006), but support for the clade was extremely weak. While only the latter study recovered monophyly of the Cypselomorphae (see below) within Metaves, the former was based on only a single locus and could not resolve their relationships according to standard criteria of statistical confidence. No morphological synapomorphies have been found that uniquely unite Metaves (or Caprimulgiformes for that matter), but numerous unlinked nuclear genes independently support their monophyly either in majority or whole. Ericson et al. (2006) concluded that if valid,

3721-733: Was only recently realized. The Apodiformes (which include the " Trochiliformes " of the Sibley-Ahlquist taxonomy ) and the Aegotheliformes form the Daedalornithes . The material evidence for this group is very equivocal; the most ancient Strisores are quite nondescript tree-dwellers but already tend towards peculiarly apomorphic feet, and no Cretaceous fossils are known. Torpor and other metabolic peculiarities are frequently found in this group, perhaps more often than in any other bird lineage. The synapomorphies that define this clade are

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