Biology:Enantiornithes

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Short description: Extinct clade of dinosaurs

Enantiornithes
Temporal range:
Early Cretaceous[1]–Late Cretaceous, 136–66 Ma
Zhouornis hani.png
Fossil specimen of a bohaiornithid (Zhouornis hani)
Scientific classification e
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Clade: Saurischia
Clade: Theropoda
Clade: Avialae
Clade: Ornithothoraces
Clade: †Enantiornithes
Walker, 1981
Subgroups

and see text

The Enantiornithes, also known as enantiornithines or enantiornitheans in literature, are a group of extinct avialans ("birds" in the broad sense), the most abundant and diverse group known from the Mesozoic era.[3][4][5] Almost all retained teeth and clawed fingers on each wing, but otherwise looked much like modern birds externally. Over eighty species of Enantiornithes have been named, but some names represent only single bones, so it is likely that not all are valid. The Enantiornithes became extinct at the Cretaceous–Paleogene boundary, along with Hesperornithes and all other non-avian dinosaurs.

Discovery and naming

The first Enantiornithes to be discovered were incorrectly referred to modern bird groups. For example, the first known species of Enantiornithes, Gobipteryx minuta, was originally considered a paleognath related to ostriches and tinamou.[6] The Enantiornithes were first recognized as a distinct lineage, or "subclass" of birds, by Cyril A. Walker in 1981. Walker made this discovery based on some partial remains from the late Cretaceous period of what is now Argentina , which he assigned to a new genus, Enantiornis, giving the entire group its name. Since the 1990s, many more complete specimens of Enantiornithes have been discovered, and it was determined that a few previously described "birds" (e.g. Iberomesornis, Cathayornis, and Sinornis) were also Enantiornithes.

The name "Enantiornithes" means "opposite birds", from Ancient Greek enantios (ἐνάντιος) "opposite" + ornithes (ὄρνιθες) "birds" . The name was coined by Cyril Alexander Walker in his landmark paper which established the group.[7] In his paper, Walker explained what he meant by "opposite":

Perhaps the most fundamental and characteristic difference between the Enantiornithes and all other birds is in the nature of the articulation between the scapula [...] and the coracoid, where the 'normal' condition is completely reversed.[7]

This refers to an anatomical feature – the articulation of the shoulder bones – which has a concave-convex socket joint between the scapula (shoulder blade) and coracoid (the primary bone of the shoulder girdle in vertebrates other than mammals) that is the reverse of that of modern birds. Specifically, in the Enantiornithes, the scapula is concave and dish-shaped at this joint, and the coracoid is convex.[3]:249–50 In modern birds, the coracoscapular joint has a concave coracoid and convex scapula.[8][9]

Walker was not clear on his reasons for giving this name in the etymology section of his paper, and this ambiguity led to some confusion among later researchers. For example, Alan Feduccia stated in 1996:

Feduccia's point about the tarsometatarsus (the combined upper foot and ankle bone) is correct, but Walker did not use this reasoning in his original paper. Walker never described the fusion of the tarsometatarsus as opposite, but rather as "Only partial". Also, it is not certain that Enantiornithes had triosseal canals, since no fossil preserves this feature.[3]

As a group, the Enantiornithes are often referred to as "enantiornithines" in literature. However, several scientists have noted that this is incorrect, because following the standard rules for forming the names of animal groups, it implies reference only to the subfamily Enantiornithinae. Following the naming conventions used for modern birds as well as extinct groups, it has been pointed out that the correct term is "enantiornithean".[10][11]

Origin and range

Birds with characteristics of Enantiornithes found in Albian of Australia , Maastrichtian of South America, and Campanian of Mexico (Alexornis[12]), Mongolia and western edge of prehistoric Asia suggest a worldwide distribution of this group or in the relatively warm regions, at least.[13] Enantiornithes have been found on every continent except Antarctica. Fossils attributable to this group are exclusively Cretaceous in age, and it is believed that the Enantiornithes became extinct at the same time as their non-avialan dinosaur relatives. The earliest known Enantiornithes are from the Early Cretaceous of Spain (e.g. Noguerornis) and China (e.g. Protopteryx) and the latest from the Late Cretaceous of North and South America (e.g. Avisaurus and Enantiornis). The widespread occurrence of this group suggests that at least some Enantiornithes were able to cross oceans under their own power; they are the first known avialan lineage with a global distribution.

Description

A life restoration of Iberomesornis, a species of early Enantiornithes

Many fossils of Enantiornithes are very fragmentary, and some species are only known from a piece of a single bone. Almost all specimens that are complete, in full articulation, and with soft tissue preservation are known from Las Hoyas in Cuenca, Spain and the Jehol group in Liaoning (China). Extraordinary remains of Enantiornithes have also been preserved in Burmese amber deposits dated to 99 million years ago and include hatchlings described in 2017[14] and 2018,[15] as well as isolated body parts such as wings[16][17][18] and feet.[17][19] These amber remains are among the most well-preserved of any mesozoic dinosaur. Fossils of this clade have been found in both inland and marine sediments, suggesting that they were an ecologically diverse group.

Enantiornithes appear to have included waders, swimmers, granivores, insectivores, fishers, and raptors. The vast majority of Enantiornithes were small, between the size of a sparrow and a starling,[20] however display considerable variation in size with some species. The largest species in this clade include Pengornis houi,[21] Xiangornis shenmi,[22] Zhouornis hani,[20] and Mirarce eatoni,[23] (with the latter species being described as similar in size to modern turkeys,) although at least a few larger species may have also existed, including a potentially crane-sized species known only from footprints in the Eumeralla Formation (and possibly also represented in the Wonthaggi Formation by a single furcula).[24] Among the smallest described specimens are unnamed hatchlings, although the holotype specimens of Parvavis chuxiongensis[25] and Cratoavis cearensis[26] are comparable in size to small tits or hummingbirds.

Skull

A reconstruction of the skull of Bohaiornis, a bohaiornithid.

Given their wide range of habitats and diets, the skulls of Enantiornithes varied considerably between species. Skulls of Enantiornithes combined a unique suite of primitive and advanced features. As in more primitive avialans like Archaeopteryx, they retained several separate cranial bones, small premaxillae (bones of the snout tip) and most species had toothy jaws rather than toothless beaks. Only a few species, such as Gobipteryx minuta, were fully toothless and had beaks. They also had simple quadrate bones, a complete bar separating each orbit (eye hole) from each antorbital fenestra, and dentaries (the main toothed bones of the lower jaw) without forked rear tips. A squamosal bone is preserved in an indeterminate juvenile specimen, while a postorbital is preserved in Shenqiornis and Pengornis. In modern birds these bones are assimilated into the cranium. Some Enantiornithes may have had their temporal fenestrae (holes in the side of the head) merged into the orbits as in modern birds due to the postorbitals either not being present or not being long enough to divide the openings.[27] A quadratojugal bone, which in modern birds is fused to the jugal, is preserved in Pterygornis.[28] The presence of these primitive features of the skull would have rendered the Enantiornithes capable of only limited cranial kinesis (the ability to move the jaw independent of the cranium).[29]

Wing

A micro-CT scan of an amber-encased wing attributable to Enantiornithes showing rachises, skin, muscle and claws.

As a very large group of birds, the Enantiornithes displayed a high diversity of different body plans based on differences in ecology and feeding, reflected in an equal diversity of wing forms, many paralleling adaptions to different lifestyles seen in modern birds. In general, the wings of Enantiornithes were advanced compared to more primitive avialans like Archaeopteryx, and displayed some features related to flight similar to those found in the lineage leading to modern birds, the Ornithuromorpha. While most Enantiornithes had claws on at least some of their fingers, many species had shortened hands, a highly mobile shoulder joint, and proportional changes in the wing bones similar to modern birds. Like modern birds, Enantiornithes had alulas, or "bastard wings", small forward-pointing arrangements of feathers on the first digit that granted higher maneuverability in the air and aided in precise landings.[30]

Several wings with preserved feathers have been found in Burmese amber. These are the first complete Mesozoic dinosaur remains preserved this way (a few isolated feathers are otherwise known, unassigned to any species), and one of the most exquisitely preserved dinosaurian fossils known.[31] The preserved wings show variations in feather pigment and prove that Enantiornithes had fully modern feathers, including barbs, barbules, and hooklets, and a modern arrangement of wing feather including long flight feathers, short coverts, a large alula and an undercoat of down.[16]

One fossil of Enantiornithes shows wing-like feather tufts on its legs, similar to Archaeopteryx. The leg feathers are also reminiscent of the four-winged dinosaur Microraptor, however differ by the feathers being shorter, more disorganized (they do not clearly form a wing) and only extend down to the ankle rather than along the foot.[32]

Tail

thumb|Fossil skeleton of [[Rapaxavis pani (a longipterygid) with a preserved pygostyle]] Clarke et al. (2006) surveyed all fossils of Enantiornithes then known and concluded that none had preserved tail feathers that formed a lift-generating fan, as in modern birds. They found that all avialans outside of Euornithes (the clade they referred to as Ornithurae) with preserved tail feathers had only short coverts or elongated paired tail plumes. They suggested that the development of the pygostyle in Enantiornithes must have been a function of tail shortening, not the development of a modern tail feather anatomy. These scientists suggested that a fan of tail feathers and the associated musculature needed to control them, known as the rectrical bulb, evolved alongside a short, triangular pygostyle, like the ones in modern birds, rather than the long, rod- or dagger-shaped pygostyles in more primitive avialans like the Enantiornithes. Instead of a feather fan, most Enantiornithes had a pair of long specialized pinfeathers similar to those of the extinct Confuciusornis and certain extant birds-of-paradise.[33]

However, further discoveries showed that at least among basal Enantiornithes, tail anatomy was more complex than previously thought. One genus, Shanweiniao, was initially interpreted as having at least four long tail feathers that overlapped each other[34] and might have formed a lift-generating surface similar to the tail fans of Euronithes,[35] though a later study indicates that Shanweiniao was more likely to have rachis-dominated tail feathers similar to feathers present in Paraprotopteryx.[36] Chiappeavis, a primitive pengornithid, had a fan of tail feathers similar to that of more primitive avialans like Sapeornis, suggesting that this might have been the ancestral condition, with pinfeathers being a feature evolved several times in early avialans for display purposes.[36] Another species of Enantiornithes, Feitianius, also had an elaborate fan of tail feathers. More importantly, soft tissue preserved around the tail was interpreted as the remains of a rectrical bulb, suggesting that this feature was not in fact restricted to species with modern-looking pygostyles, but might have evolved much earlier than previously thought and been present in many Enantiornithes.[37] At least one genus of Enantiornithes, Cruralispennia, had a modern-looking pygostyle but lacked a tail fan.[38]

Biology

Diet

Given the wide diversity of skull shape among Enantiornithes, many different dietary specializations must have been present among the group. Some, like Shenqiornis, had large, robust jaws suitable for eating hard-shelled invertebrates. The short, blunt teeth of Pengornis were likely used to feed on soft-bodied arthropods.[27] The strongly hooked talons of Bohaiornithidae suggest that they were predators of small to medium-sized vertebrates, but their robust teeth instead suggest a diet of hard-shelled animals.[2]

A few specimens preserve actual stomach contents. Unfortunately, none of these preserve the skull, so direct correlation between their known diet and snout/tooth shape cannot be made. Eoalulavis was found to have the remains of exoskeletons from aquatic crustaceans preserved in its digestive tract,[39] and Enantiophoenix preserved corpuscles of amber among the fossilized bones, suggesting that this animal fed on tree sap, much like modern sapsuckers and other birds. The sap would have fossilized and become amber.[40] However, more recently it has been suggested that the sap moved post-mortem, hence not representing true stomachal contents. Combined with the putative fish pellets of Piscivorenantiornis turning out to be fish excrement, the strange stomachal contents of some species turning out to be ovaries and the supposed gastroliths of Bohaiornis being random mineral precipitates, only the Eoalulavis displays actual stomach contents.[41]

A study on paravian digestive systems indicates that known Enantiornithes lacked a crop and a gizzard, didn't use gastroliths and didn't eject pellets. This is considered at odds with the high diversity of diets that their different teeth and skull shapes imply,[42] though some modern birds have lost the gizzard and rely solely on strong stomachal acids.[43] An example was discovered with what was suspected to be gastroliths in the what would have been the fossil's stomach, re-opening the discussion of the use of gastroliths by Enantiornithes. X-ray and scanning microscope inspection of the rocks determined that they were actually chalcedony crystals, and not gastroliths.[44]

Longipterygidae is the most extensively studied family in terms of diet due to their rather unusual rostral anatomy, with long jaws and few teeth arranged at the jaw ends. They have variously been interpreted as piscivores,[45] probers akin to shorebirds[46] and as arboreal bark-probers.[47] A 2022 study however does find them most likely to be generalistic insectivores (sans possibly Shengjingornis due to its larger size, poorly preserved skull and unusual pedal anatomy), being too small for specialised carnivory and herbivory; the atypical rostrum is tentatively speculated to be unrelated to feeding ecology.[48]

Predation

A fossil from Spain reported by Sanz et al. in 2001 included the remains of four hatchling skeletons of three different species of Enantiornithes. They are substantially complete, very tightly associated, and show surface pitting of the bones that indicates partial digestion. The authors concluded that this association was a regurgitated pellet and, from the details of the digestion and the size, that the hatchlings were swallowed whole by a pterosaur or small theropod dinosaur. This was the first evidence that Mesozoic avialans were prey animals, and that some Mesozoic pan-avians regurgitated pellets like owls do today.[49]

Life history

Fossilized eggs of Gobipteryx minuta, Dinosaurium (Prague)

Known fossils of Enantiornithes include eggs,[50][51] embryos,[52] and hatchlings.[53] An embryo, still curled in its egg, has been reported from the Yixian Formation.[54] Juvenile specimens can be identified by a combination of factors: rough texture of their bone tips indicating portions which were still made of cartilage at the time of death, relatively small breastbones, large skulls and eyes, and bones which had not yet fused to one another.[55] Some hatchling specimens have been given formal names, including "Liaoxiornis delicatus"; however, Luis Chiappe and colleagues considered the practice of naming new species based on juveniles detrimental to the study of Enantiornithes, because it is nearly impossible to determine which adult species a given juvenile specimen belongs to, making any species with a hatchling holotype a nomen dubium.[55]

Together with hatchling specimens of the Mongolian Gobipteryx[56] and Gobipipus,[57][58] these finds demonstrate that hatchling Enantiornithes had the skeletal ossification, well-developed wing feathers, and large brain which correlate with precocial or superprecocial patterns of development in birds of today. In other words, Enantiornithes probably hatched from the egg already well developed and ready to run, forage, and possibly even fly at just a few days old.[55]

Findings suggests Enantiornithes, especially the toothed species, had a longer incubation time than modern birds.[59][60]

Analyses of Enantiornithes bone histology have been conducted to determine the growth rates of these animals. A 2006 study of Concornis bones showed a growth pattern different from modern birds; although growth was rapid for a few weeks after hatching, probably until fledging, this small species did not reach adult size for a long time, probably several years.[61] Other studies have all supported the view that growth to adult size was slow, as it is in living precocial birds (as opposed to altricial birds, which are known to reach adult size quickly).[39] Studies of the rate of bone growth in a variety of Enantiornithes has shown that smaller species tended to grow faster than larger ones, the opposite of the pattern seen in more primitive species like Jeholornis and in non-avialan dinosaurs.[62] Some analyses have interpreted the bone histology to indicate that Enantiornithes may not have had fully avian endothermy, instead having an intermediate metabolic rate.[63] However a 2021 study rejects the idea that they had less endothermic metabolisms than modern birds.[64]

Evidence of colonial nesting has been found in Enantiornithes, in sediments from the Late Cretaceous (Maastrichtian) of Romania.[65] Evidence from nesting sites shows that Enantiornithes buried their eggs like modern megapodes, which is consistent with their inferred superprecocial adaptations.[66]

A 2020 study on a juvenile's feathers further stresses the ontological similarities to modern megapodes, but cautions several differences such as the arboreal nature of most Enantiornithes as opposed to the terrestrial lifestyle of megapodes.[67]

It has been speculated that superprecociality in Enantiornithes might have prevented them from developing specialised toe arrangements seen in modern birds like zygodactyly.[68]

Although the vast majority of histology studies and known remains of Enantiornithes point to superprecociality being the norm, one specimen, MPCM-LH-26189, seems to represent an altricial juvenile, implying that like modern birds Enantiornithes explored multiple reproductive strategies.[69]

Flight

Because many Enantiornithes lacked complex tails and possessed radically different wing anatomy compared to modern birds, they have been the subject of several studies testing their flight capabilities.

Traditionally, they have been considered inferior flyers, due to the shoulder girdle anatomy being assumed to be more primitive and unable to support a ground-based launching mechanism,[70] as well as due to the absence of rectrices in many species.[33][35][71]

However, several studies have shown that they were efficient flyers, like modern birds, possessing a similarly complex nervous system and wing feather ligaments. Additionally, the lack of a complex tail appears to not have been very relevant for avian flight as a whole - some extinct birds like lithornids also lacked complex tail feathers but were good flyers,[72] and they appear to have been capable of a ground based launching.[73]

Enantiornithes resemble Ornithuromorphs in many anatomical features of the flight apparatus, but a sternal keel is absent in the basal-most members, only a single basal taxon appears to have had a triosseal canal, and their robust pygostyle seems unable to support the muscles that control the modern tail feathers involved in flight.[74] Though some basal Enantiornithes exhibit ancestral flight apparatuses, by the end of the Mesozoic many Enantiornithes had several features convergent with the Neornithes including a deeply keeled sternum, a narrow furcula with a short hypocleidium, and ulnar quill knobs that indicate increased aerial abilities.[75][76]

At least Elsornis appears to have become secondarily flightless.[77]

Classification

Some researchers classify Enantiornithes, along with the true birds, in the class Aves. Others use the more restrictive crown group definition of Aves (which only includes neornithes, anatomically modern birds), and place Enantiornithes in the more inclusive group Avialae. Enantiornithes were more advanced than Archaeopteryx, Confuciusornis, and Sapeornis, but in several respects they were more primitive than modern birds, perhaps following an intermediate evolutionary path.

A consensus of scientific analyses indicates that Enantiornithes is one of two major groups within the larger group Ornithothoraces. The other ornithothoracine group is Euornithes or Ornithuromorpha, which includes all living birds as a subset. This means that Enantiornithes were a successful branch of avialan evolution, but one that diversified entirely separately from the lineage leading to modern birds.[3] One study has however found that the shared sternal anatomy was acquired independently and such a relationship needs to be reexamined.[78]

Enantiornithes classification and taxonomy has historically been complicated by a number of factors. In 2010, paleontologists Jingmai O'Connor and Gareth Dyke outlined a number of criticisms against the prevailing practices of scientists failing to describe many specimens in enough detail for others to evaluate thoroughly. Some species have been described based on specimens which are held in private collections, making further study or review of previous findings impossible. Because it is often unfeasible for other scientists to study each specimen in person given the worldwide distribution of the Enantiornithes, and due to the many uninformative descriptions which have been published on possibly important specimens, many of these specimens become "functional nomina dubia".[79] Furthermore, many species have been named based on extremely fragmentary specimens, which would not be very informative scientifically even if they were described sufficiently. Over one-third of all named species are based on only a fragment of a single bone. O'Connor and Dyke argued that while these specimens can help expand knowledge of the time span or geographic range of the Enantiornithes and it is important to describe them, naming such specimens is "unjustifiable".[79]

Relationships

Enantiornithes is the sister group to Euornithes, and together they form a clade called Ornithothoraces (though see above). Most phylogenetic studies have recovered Enantiornithes as a monophyletic group distinct from the modern birds and their closest relatives. The 2002 phylogenetic analysis by Clarke and Norell, though, reduced the number of Enantiornithes autapomorphies to just four.[80]

Enantiornithes systematics are highly provisional and notoriously difficult to study, due to their small size[26] and the fact that Enantiornithes tend to be extremely homoplastic, or very similar to each other in most of their skeletal features due to convergent evolution rather than common ancestry.[36] What appears fairly certain by now is that there were subdivisions within Enantiornithes possibly including some minor basal lineages in addition to the more advanced Euenantiornithes. The details of the interrelationship of all these lineages, indeed the validity of most, is disputed, although the Avisauridae, for one example, seem likely to constitute a valid group. Phylogenetic taxonomists have hitherto been very reluctant to suggest delimitations of clades of Enantiornithes.[81]

One such delineation named the Euenantiornithes, was defined by Chiappe (2002) as comprising all species closer to Sinornis than to Iberomesornis. Because Iberomesornis is often found to be the most primitive or basal member of the Enantiornithes, Euenantiornithes may be an extremely inclusive group, made up of all Enantiornithes except for Iberomesornis itself. Despite being in accordance with phylogenetic nomenclature, this definition of Euenantiornithes was severely criticized by some researchers, such as Paul Sereno, who called it "a ill-defined clade [...] a good example of a poor choice in a phylogenetic definition".[81]

The cladogram below was found by an analysis by Wang et al. in 2015, updated from a previous data set created by Jingmai O'Connor.[28]

Ornithothoraces
 

Euornithes Y martini.png

 
†Enantiornithes
 

†Protopteryx Protopteryx Silhouette.png

 
 

†Pengornithidae Chiappeavis magnapremaxillo.png

 
 

†Longipterygidae Longipteryx restoration.jpg

 
†Euenantiornithes
 

†Eocathayornis

 
 
 

†Elsornis

 
 
 

†Dunhuangia

 
 

†Fortunguavis

 
 
 

†Bohaiornithidae Zhouornis hani.png

 
 
 

†Pterygornis

 
 
 

†Eoenantiornis

 
 
 

†Cathayornis

 
 
 

†Vescornis

 
 

†Neuquenornis

 
 

†Gobipteryx

 
 

†EoalulavisEoaluavis life restoration.png

 
 
 

†Qiliania

 
 

†Concornis

 
 
 
 
 
 
 
 
 
 
 
 

The cladogram below is from Wang et al., 2022, and includes most named taxa and recovers several previously-named clades. Letters on branches indicate the positions of "wildcard" taxa, those which have been recovered in multiple disparate positions.[82]

Enantiornithes
 

Cruralispennia

 
 
 

Protopteryx

 
 
 
 

Elsornis

 
 

Flexomornis

 
 
 
Longipterygidae
 

Iberomesornis

 
bh
bh

Longirostravis

 
 

Rapaxavis

 
 
 
Euenantiornithes
c
e

Sinornis

 
e
 

Enantiornis

 
 
 

Halimornis

 
 
 

Concornis

 
 

Neuquenornis

 
 
 
 
c
b
 

Eoalulavis

 
 

Liaoningornis

 
 
 
 

Gobipteryx

 
 

Hebeiornis

 
 
 
 
 
 

Eocathayornis

 
 
 
 

Qiliania

 
Avisauridae
 
 

Intiornis

 
 

Soroavisaurus

 
 
 
 

Avisaurus

 
 
 

Gettyia

 
n

Mirarce

 
 
 
e
 
 
 

Feitianius

 
 
 
fn

Longusunguis

 
fn
n

Dunhuangia

 
fn

Musivavis

 
 
 
 
Pengornithidae
 

Pengornis

 
m
 

Yuanchuavis

 
 
 

Eopengornis

 
 

Parapengornis

 
 
 
m
 
 
 

Grabauornis

 
 
 

Parvavis

 
 
 

Cratoavis

 
 

Gretcheniao

 
 
 
 
 
 
 

Huoshanornis

 
 

Shengjingornis

 
 
 
 

Zhouornis

 
Bohaiornithidae
 

Bohaiornis

 
ij
 

Eoenantiornis

 
 
 

Fortunguavis

 
 
 

Mystiornis

 
 
 

Shenqiornis

 
 

Sulcavis

 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
l

Key to letters:

b = Boluochia
c = Cathayornis
e = Enantiophoenix
f = Houornis
h = Longipteryx
i = Parabohaiornis
j = Pterygornis
l = Vorona
m = Yuanjiawaornis
n = Yungavolucris

List of genera

Incertae sedis

Enantiornithes taxonomy is difficult to evaluate, and as a result few clades within the group are consistently found by phylogenetic analyses. Most Enantiornithes are not included in any specific family, and as such are listed here. Many of these have been considered Euenantiornithes, although the controversy behind this name means that it is not used consistently in studies of Enantiornithes.[citation needed]

Name Year Formation Location Notes Images
Abavornis 1998 Bissekty Formation (Late Cretaceous, Turonian to Coniacian)  Uzbekistan One of many fragmentary Bissekty Enantiornithes, known only from coracoids
Alethoalaornis 2007 Jiufotang Formation (Early Cretaceous, Aptian)  China Poorly known
Alexornis 1976 La Bocana Roja Formation (Late Cretaceous, Campanian)  Mexico One of the first Enantiornithes known. Once thought to be an ancient relative of rollers and woodpeckers Alexornis antecedens.png
Avimaia 2019 Xiagou Formation (Late Cretaceous, Aptian)  China One specimen from this genus died with an unlaid egg in its body Fossil-AvimaiaSchweitzeraeWithUnlaidEgg.jpg
Bauxitornis 2010 Csehbánya Formation (Late Cretaceous, Santonian)  Hungary Fragmentary but unique in the structure of its tarsometatarsus Bauxitornis.jpg
Brevirostruavis 2021 Jiufotang Formation (Early Cretaceous, Aptian)  China Possessed an enlarged hyoid that suggests a feeding specialization similar to hummingbirds, honeyeaters, and woodpeckers Brevirostruavis macrohyoideus restoration by Lauren Helton.png
Castignovolucris 2023 Unnamed formation (Late Cretaceous, Campanian)  France May have been the size of a Canada goose
Catenoleimus 1998 Bissekty Formation (Late Cretaceous, Turonian to Coniacian)  Uzbekistan One of many fragmentary Bissekty Enantiornithes, known only from a coracoid
Cathayornis 1992 Jiufotang Formation (Early Cretaceous, Aptian)  China One of the first Jehol biota Enantiornithes described. Known from many species, although some are now placed into their own genera. May have had a similar appearance and lifestyle to a pitta Cathayornis - early cretaceous Liaoning IMG 5191 Beijing Museum of Natural History.jpg
Concornis 1992 Las Hoyas (Early Cretaceous, Barremian)  Spain One of the most complete Las Hoyas Enantiornithes Concornis lacustris 438.jpg
Cratoavis[83] 2015 Santana Formation (Early Cretaceous, Aptian)  Brazil A very well-preserved South American member of the group, complete with ribbon-like tail feathers
Cruralispennia[38] 2017 Huajiying Formation (Early Cretaceous, Hauterivian)  China Had an unusual ornithuromorph-like pygostyle and brush-like thigh feathers. One of the oldest Enantiornithes Ncomms14141-f1.jpg
Cuspirostrisornis 1997 Jiufotang Formation (Early Cretaceous, Aptian)  China Originally mistakenly believed to have possessed a pointed beak Cuspirostrornis houi life restoration.jpg
Dalingheornis 2006 Yixian Formation (Early Cretaceous, Aptian)  China Was well-adapted for climbing due to its heterodactyl feet, like those of a trogon
Dunhuangia[84] 2015 Xiagou Formation (Early Cretaceous, Aptian)  China A genus of Enantiornithes from the Changma basin, an area which is unusually dominated by ornithuromorphs
Elbretornis 2009 Lecho Formation (Late Cretaceous, Maastrichtian)  Argentina Only known from wing bones. May be synonymous with other Lecho formation Enantiornithes
Elektorornis 2019 Burmese Amber (Late Cretaceous, Cenomanian)  Myanmar Known from a foot preserved in amber with an elongated middle toe
Enantiornis 1981 Lecho Formation (Late Cretaceous, Maastrichtian)  Argentina Although only known from a few bones, this genus is the namesake of Enantiornithes. It was also one of the largest and last representative of the group prior to their extinction
Eoalulavis 1996 Las Hoyas (Early Cretaceous, Barremian)  Spain Preserves feathers including an alula, a specialized type of feather which controls air flow over the wing
Eocathayornis 2002 Jiufotang Formation (Early Cretaceous, Aptian)  China Once considered to be a basal close relative of Cathayornis, although now considered to be more distantly related
Eoenantiornis 1999 Yixian Formation (Early Cretaceous, Aptian)  China Well-preserved but inconsistent in phylogenetic placement Eoenantiornis-Paleozoological Museum of China.jpg
Evgenavis 2014 Ilek Formation (Early Cretaceous, Barremian)  Russia Known only from a tarsometatarsus which shares some features with those of Enantiornithes
Explorornis 1998 Bissekty Formation (Late Cretaceous, Turonian to Coniacian)  Uzbekistan One of many fragmentary Bissekty Enantiornithes, known only from coracoids
Falcatakely 2020 Maevarano Formation (Late Cretaceous, Maastrichtian)  Madagascar Developed a massive snout with only a single tooth, despite retaining a "primitive" skull arrangement in contrast to modern birds Falcatakely restoration.jpg
Feitianius[37] 2015 Xiagou Formation (Early Cretaceous, Aptian)  China Possessed an elaborate set of tail feathers, unlike the paired ribbon-like feathers of most Enantiornithes 200px
Flexomornis 2010 Woodbine Formation (Late Cretaceous, Cenomanian)  United States ( Texas) One of the oldest North American avialans found, albeit known only from fragmentary remains
Fortipesavis 2021 Burmese amber (Late Cretaceous, Cenomanian)  Myanmar Had an enlarged outer toe that may have been an adaptation for perching
Fortunguavis[85] 2014 Jiufotang Formation (Early Cretaceous, Aptian)  China Had robust bones, including feet and claws which may have been adapted for climbing trees
Grabauornis[86] 2015 Yixian Formation (Early Cretaceous, Barremian)  China The proportions of the wings of this genus of Enantiornithes as well as the presence of an alula suggest that it was a good flier
Gracilornis 2011 Jiufotang Formation (Early Cretaceous, Aptian)  China A possible relative of Cathayornis with characteristically slender bones
Gurilynia 1999 Nemegt Formation (Late Cretaceous, Maastrichtian)  Mongolia A poorly known genus of Enantiornithes, but evidently a large and late-surviving member of the group
Hollanda[87] 2010 Barun Goyot Formation (Late Cretaceous, Campanian)  Mongolia Originally identified as an ornithuromorph but since reinterpreted as a genus of Enantiornithes closely related to Lectavis.[88] Hollanda luceria 2021.jpg
Holbotia[89] 2015 Andaikhudag Formation (Early Cretaceous, Aptian)  Mongolia Considered a small pterosaur since its discovery in 1977 until it received a formal description in 2015. Possessed unique neck vertebrae and a primitive palate
Houornis 1997 Jiufotang Formation (Early Cretaceous, Aptian)  China Once considered to be dubious or a species of Cathayornis, although a 2015 study considered it to be a valid genus[90]
Huoshanornis 2010 Jiufotang Formation (Early Cretaceous, Aptian)  China May have been a very maneuverable flier due to the structure of its hand and sternum
Iberomesornis 1992 Las Hoyas (Early Cretaceous, Barremian)  Spain One of the first genera of Enantiornithes known from decent remains. Also one of the oldest and most primitive members of the group Iberomesornis-model.jpg
Incolornis 1998 Bissekty Formation (Late Cretaceous, Turonian to Coniacian)  Uzbekistan One of many fragmentary Bissekty Enantiornithes, known only from coracoids. One species was once considered to belong to Enantiornis
Junornis[91] 2017 Yixian Formation (Early Cretaceous, Aptian)  China So well preserved that its flight pattern could be reconstructed using the proportions of its feathers and wings Junornis.PNG
Kizylkumavis 1984 Bissekty Formation (Late Cretaceous, Turonian to Coniacian)  Uzbekistan One of the many fragmentary Bissekty Enantiornithes, known only from a humerus fragment Kizylkumavis cretacea.jpg
Largirostrornis 1997 Jiufotang Formation (Early Cretaceous, Aptian)  China Possibly related to Cuspirostrisornis or a synonym of Cathayornis
Lectavis 1993 Lecho Formation (Late Cretaceous, Maastrichtian)  Argentina A large and long-legged member of the group, proportionally similar to modern shorebirds
Lectavis bretincola reconstruction by Lauren Helton.jpg
Lenesornis 1996 Bissekty Formation (Late Cretaceous, Turonian to Coniacian)  Uzbekistan One of many fragmentary Bissekty Enantiornithes, known only from a synsacrum fragment. Originally considered to belong to Ichthyornis
Liaoningornis 1996 Yixian Formation (Early Cretaceous, Aptian)  China Originally believed to be an ornithuran, but now considered a relative of Eoalulavis
Longchengornis 1997 Jiufotang Formation (Early Cretaceous, Aptian)  China May have been a synonym of Cathayornis
Martinavis 2007 Grès à Reptiles Formation, Lecho Formation (Late Cretaceous, Maastrichtian)  France  Argentina Although known only from humeri, this genus was large and lived in a broad range
Microenantiornis 2017 Jiufotang Formation (Early Cretaceous, Aptian)  China A small member of the group which possessed several primitive and derived features compared to other Enantiornithes
Mirusavis 2020 Yixian Formation (Early Cretaceous, Barremian to Aptian)  China Holotype was a small osteologically immature female preserved with medullary bone tissue
Monoenantiornis[92] 2016 Yixian Formation (Early Cretaceous, Aptian)  China Known from a juvenile specimen which depicts how various features developed in Enantiornithes as they age
Musivavis 2022 Jiufotang Formation (Early Cretaceous, Aptian)  China Most similar to bohaiornithids but also has features of other groups of Enantiornithes
Nanantius 1986 Toolebuc Formation (Early Cretaceous, Albian)  Australia Fragmentary, but may have been a seabird because remains from this genus have been found as ichthyosaur gut content
Noguerornis 1989 El Montsec (Early Cretaceous, Barremian)  Spain Preserves impressions of a propatagium, a skin flap on the shoulder which forms part of a wing
Orienantius 2018 Huajiying Formation (Early Cretaceous, Hauterivian)  China Many soft tissue details of specimens from this genus were revealed by UV light
Otogornis 1993 Yijinholuo Formation (Early Cretaceous)  China Poorly known
Paraprotopteryx 2007 Qiaotou member of the Huajiying Formation (Early Cretaceous, Aptian?)  China Seemingly had four ribbon-like tail feathers instead of only two as in most Enantiornithes
Parvavis[93] 2014 Jiangdihe Formation (Late Cretaceous, Turonian to Santonian)  China Small but fully mature at the time of its death. One of only a few Chinese Enantiornithes dated to the Late Cretaceous Parvavis.jpg
Piscivorenantiornis[94] 2017 Jiufotang Formation (Early Cretaceous, Aptian)  China Known from a disarticulated skeleton preserved overlying a piece of stomach content composed of fish bones, which may have been its last meal
Protopteryx 2000 Huajiying Formation (Early Cretaceous, Hauterivian)  China One of the oldest and most primitive members of the group Protopteryx fengningensis (BMNHC Ph1060A) NMNS.jpg
Pterygornis[28] 2016 Jiufotang Formation (Early Cretaceous, Aptian)  China One disarticulated skeleton from this genus possesses well-preserved bones of the skull, including a quadratojugal
Qiliania 2011 Xiagou Formation (Early Cretaceous, Aptian)  China Some of this genus's remains include well-preserved hindlimbs. the species names, Q. graffini, is named after Greg Graffin from the band Bad Religion
Sazavis 1989 Bissekty Formation (Late Cretaceous, Turonian to Coniacian)  Uzbekistan One of many fragmentary Bissekty Enantiornithes, known only from a tibiotarsus (shin bone) Sazavis prisca.png
Shangyang 2019 Jiufotang Formation (Early Cretaceous, Aptian)  China Unusually, the premaxillae of this genus were fused
Sinornis 1992 Jiufotang Formation (Early Cretaceous, Aptian)  China One of the first Jehol biota Enantiornithes described. Similar to Cathayornis but usually considered to be distinct Sinornis2.jpg
Xiangornis 2012 Jiufotang Formation (Early Cretaceous, Aptian)  China The hand of this genus was similar to that of ornithuromorphs, likely through convergent evolution. A large member of the group
Yatenavis[95] 2022 Chorrillo Formation (Late Cretaceous, Maastrichtian)  Argentina The southernmost known member of Enantiornithes and one of the youngest members of the group
Yuanjiawaornis[96] 2015 Jiufotang Formation (Early Cretaceous, Aptian)  China One of the largest Enantiornithes known from decent remains
Yungavolucris 1993 Lecho Formation (Late Cretaceous, Maastrichtian)  Argentina Had a large and unusually wide tarsometatarsal (ankle bone) Yungavolucris holotype tarsometatarsus.png
Yuornis 2021 Qiupa Formation (Late Cretaceous, Maastrichtian)  China A large, toothless genus of Enantiornithes, one of the most-well preserved members from the Late Cretaceous. Yuornis skull diagram.png

Longipterygidae

The Longipterygidae was a family of long-snouted early Cretaceous Enantiornithes, with teeth only at the tips of the snout. They are generally considered to be fairly basal members of the group.[34]

Name Year Formation Location Notes Images
Boluochia 1995 Jiufotang Formation (Early Cretaceous, Aptian)  China Originally mistakenly believed to have possessed a hooked beak
Camptodontornis 2010 Jiufotang Formation (Early Cretaceous, Aptian)  China Originally called Camptodontus, although that genus name is occupied by a beetle
Dapingfangornis 2006 Jiufotang Formation (Early Cretaceous, Aptian)  China May have had a thorn-like structure on its forehead
Longipteryx 2001 Jiufotang Formation (Early Cretaceous, Aptian)  China The most common and well-known member of the family Longipteryx chaoyangensis - early cretaceous Liaoning IMG 5197 Beijing Museum of Natural History.jpg
Longirostravis 2004 Yixian Formation (Early Cretaceous, Aptian)  China Like other longipterygids, it possessed a thin snout which may have been used for probing for invertebrates in mud or bark Longirostravis-Tianjin Natural History Museum.jpg
Rapaxavis 2009 Jiufotang Formation (Early Cretaceous, Aptian)  China Specialized for perching due to the structure of its feet Rapaxavis.jpg
Shanweiniao 2009 Yixian Formation (Early Cretaceous, Aptian)  China Acquired multiple tail feathers which may have been capable of generating lift as in modern birds Shanweiniao cooperorum.jpg
Shengjingornis 2012 Jiufotang Formation (Early Cretaceous, Aptian)  China A large member of the family

Pengornithidae

The Pengornithidae was a family of large early Enantiornithes. They had numerous small teeth and numerous primitive features which are lost in most other Enantiornithes.[1] Mostly known from the early Cretaceous of China, with putative Late Cretaceous taxa from Madagascar.[97]

Name Year Formation Location Notes Images
Chiappeavis[36] 2015 Jiufotang Formation (Early Cretaceous, Aptian)  China Possessed a fan-shaped tail composed of many feathers Chiappeavis magnapremaxillo.png
Eopengornis 2014 Huajiying Formation (Early Cretaceous, Hauterivian)  China The oldest known member of the family, and one of the oldest putative genera of Enantiornithes known. Possessed extremely well-preserved tail ribbons
Parapengornis[98] 2015 Jiufotang Formation (Early Cretaceous, Aptian)  China Proposed to have a woodpecker-like lifestyle due to features of the foot and tail Parapengornis holotype.png
Pengornis 2008 Jiufotang Formation (Early Cretaceous, Aptian)  China The first pengornithid discovered, and also one of the largest members of the Enantiornithes known from decent remains
Yuanchuavis 2021 Jiufotang Formation (Early Cretaceous, Aptian)  China Possesses an elaborate "pintail" tail fan longer than its body, which may have had a display function Yuanchuavis kompsosoura.png

Bohaiornithidae

Bohaiornithids were large but geologically short-lived early Enantiornithes, with long, hooked talons and robust teeth with curved tips. They may have been equivalent to birds of prey, although this interpretation is open to much debate.[2] The monophyly of this group is doubtful, and it may actually be an evolutionary grade.[99]

Name Year Formation Location Notes Images
Beiguornis 2022 Longjiang Formation (Early Cretaceous, Aptian)  China Had a short but robust manual ungual
Bohaiornis 2011 Jiufotang Formation (Early Cretaceous, Aptian)  China Originally considered to have been preserved with gastroliths, although later these were found to be mineral concretions Bohaiornis skull reconstruction.png
Gretcheniao 2019 Yixian Formation (Early Cretaceous, Barremian)  China Adapted for flapping, rather than soaring, flight. Its describers suggest paraphyly or polyphyly of Bohaiornithidae Gretcheniao sinensis holotype.png
Linyiornis[100] 2016 Jiufotang Formation (Early Cretaceous, Aptian)  China A possible member of the family, known from a well-preserved skeleton complete with structures believed to be developing eggs
Longusunguis 2014 Jiufotang Formation (Early Cretaceous, Aptian)  China A fairly typical member of the family Longusunguis skull.png
Parabohaiornis 2014 Jiufotang Formation (Early Cretaceous, Aptian)  China A close relative of Bohaiornis
Shenqiornis 2010 Qiaotou member of the Huajiying Formation (Early Cretaceous, Aptian?)  China The first known member of the family, although not considered a close relative of Bohaiornis until a few years later. Preserves a large postorbital bone
Sulcavis 2013 Yixian Formation (Early Cretaceous, Aptian)  China A close relative of Shenqiornis with grooved enamel on its teeth, unique among fossil birds Sulcavis skull.png
Zhouornis 2013 Jiufotang Formation (Early Cretaceous, Aptian)  China A large member of the family with a well-preserved braincase Zhouornis hani.png

Gobipterygidae

Some members of the group are obscure or poorly described and may be synonymous with its type species, Gobipteryx minuta.

Name Year Formation Location Notes Images
Gobipteryx 1974 Barun Goyot Formation (Late Cretaceous, Campanian)  Mongolia A toothless advanced genus of Enantiornithes, possessing a robust beak which convergently evolved with those of modern birds
Jibeinia 1997 Qiaotou member of the Huajiying Formation (Early Cretaceous, Aptian?)  China Poorly known and described from a skeleton which has now been lost. May have been synonymous with Vescornis
Vescornis 2004 Qiaotou member of the Huajiying Formation (Early Cretaceous, Aptian?)  China A small and short-snouted genus of Enantiornithes which may be synonymous with Jibeinia

Avisauridae

Avisauridae is subjected to two differing definitions of varying inclusiveness. The more inclusive definition, which follows Cau & Arduini (2008), is used here. Avisaurids were a long-lasting and widespread family of Enantiornithes, which are mainly distinguished by specific features of their tarsometatarsals (ankle bones). The largest and most advanced members of the group survived in North and South America up until the end of the Cretaceous, yet are very fragmentary compared to some earlier taxa.

Name Year Formation Location Notes Images
Avisaurus 1985 Hell Creek Formation (Late Cretaceous, Maastrichtian)  United States ( Montana) The eponymous avisaurid, as well as one of the largest members of the family. Originally considered a non-avialan dinosaur Avisaurus and Brachychampsa by tomozsaurus.jpg
Elsornis 2007 Djadochta Formation (Late Cretaceous, Campanian)  Mongolia Although incomplete, its skeleton possesses three-dimensional preservation. Possibly flightless due to its wing proportions
Enantiophoenix 2008 Ouadi al Gabour Formation (Late Cretaceous, Cenomanian)  Lebanon Was once believed to have fed on tree sap as it was preserved in association with amber beads, however this was later determined as an artefact of preservation and not an indicator of diet
Gettyia 2018 Two Medicine Formation (Late Cretaceous, Campanian)  United States ( Montana) A new genus for Avisaurus gloriae
Halimornis 2002 Mooreville Chalk Formation (Late Cretaceous, Campanian)  United States ( Alabama) Would have lived in a coastal environment
Intiornis 2010 Las Curtiembres Formation (Late Cretaceous, Campanian)  Argentina Although closely related to some of the largest avisaurids, members of this genus were very small birds Intiornis holotype left foot.png
Mirarce 2018 Kaiparowits Formation (Late Cretaceous, Campanian)  United States ( Utah) The most complete known North American avisaurid Mirarce eatoni.jpg
Mystiornis 2011 Ilek Formation (Early Cretaceous, Barremian to Aptian)  Russia Possesses a myriad of features from various groups in Paraves, although most closely resembles avisaurids among sampled groups
Neuquenornis 1994 Bajo de la Carpa Formation (Late Cretaceous, Santonian)  Argentina Possessed long wings and a reverse hallux, indicating good flight and perching abilities
Soroavisaurus 1993 Lecho Formation (Late Cretaceous, Maastrichtian)  Argentina A very close relative of Avisaurus

Dubious genera and notable unnamed specimens

  • Gobipipus reshetovi: Described in 2013 from embryo specimens within eggshells from the Barun Goyot Formation of Mongolia. These specimens were very similar to embryonic Gobipteryx specimens, although the describers of Gobipipus (a set of controversial paleontologists including Evgeny Kurochkin and Sankar Chatterjee) consider it distinct.[101]
  • Hebeiornis fengningensis: A synonym of Vescornis due to having been described from the same specimen. Despite having been described in 1999, 5 years prior to the description of Vescornis, the description was so poor compared to the description of Vescornis that the latter name is considered to take priority by most authors. As a result, the name Hebeiornis is considered a nomen nudum ("naked name").
  • "Proornis" is an informally-named bird from North Korea. It may not be a member of Enantiornithes.
  • Liaoxiornis delicatus: Described in 1999 from a specimen of Enantiornithes found in the Yixian Formation. This specimen was originally considered to be a tiny adult, but later found to be a hatchling. Other specimens have henceforth been assigned to the genus. Due to a lack of distinguishing feature, many paleontologists have considered this genus an undiagnostic nomen dubium.
  • "Wasaibpanchi": A supposed member of Enantiornithes from Pakistan; the describing paper is of dubious status.
  • LP-4450: A juvenile of an indeterminate specimen of Enantiornithes from the El Montsec Formation of Spain . Its 2006 description studied the histology of the skeleton, while later studies reported a squamosal bone present in the specimen but unknown in other Enantiornithes.
  • IVPP V 13939: Briefly described in 2004, this Yixian Enantiornithes specimen had advanced pennaceous feathers on its legs, similar to (albeit shorter than) those of other paravians such as Microraptor and Anchiornis.[32]
  • DIP-V-15100 and DIP-V-15101: Two different wings from hatchling specimens which were described in 2015. They attracted a significant amount of media attention upon their description. They were preserved in exceptional details due to having been trapped within Burmese amber for approximately 99 million years.[16]
  • HPG-15-1: A partial corpse of an Enantiornithes hatchling also preserved in Burmese amber. Although indeterminate, it attracted even more media attention than the two wings upon its description in 2017.[14]
  • CUGB P1202: An indeterminate juvenile bohaiornithid from the Jiufotang Formation. A 2016 analysis of its feathering found elongated putative melanosomes, suggesting that a large portion of its feathering was iridescent.
  • DIP-V-15102: Another corpse of an indeterminate hatchling preserved in Burmese amber. Described in early 2018.[15]
  • MPCM-LH-26189 a/b: A partial skeleton of a hatchling from Las Hoyas in Spain, including both slab and counter-slab components. Its 2018 description revealed how various features developed in Enantiornithes as they aged. Such features include the ossification of the sternum from various smaller bones, and the fusion of tail vertebrae into a pygostyle.

References

  1. ↑ 1.0 1.1 1.2 Wang, X.; O'Connor, J. K.; Zheng, X.; Wang, M.; Hu, H.; Zhou, Z. (2014). "Insights into the evolution of rachis dominated tail feathers from a new basal enantiornithine (Aves: Ornithothoraces)". Biological Journal of the Linnean Society 113 (3): 805–819. doi:10.1111/bij.12313. 
  2. ↑ 2.0 2.1 2.2 Wang, Min; Zhou, Zhong-He; O'Connor, Jingmai K.; Zelenkov, Nikita V. (2014). "A new diverse enantiornithine family (Bohaiornithidae fam. nov.) from the Lower Cretaceous of China with information from two new species". Vertebrata PalAsiatica 52 (1): 31–76. http://www.ivpp.cas.cn/cbw/gjzdwxb/xbwzxz/201401/P020140121386256443882.pdf. 
  3. ↑ 3.0 3.1 3.2 3.3 Chiappe, Luis M.; Walker, Cyril A. (2002). "Skeletal Morphology and Systematics of the Cretaceous Euenantiornithes (Ornithothoraces: Enantiornithes)". in Chiappe, Luis M.; Witmer, Lawrence M.. Mesozoic Birds: Above the Heads of Dinosaurs. University of California Press. pp. 240–67. ISBN 978-0-520-20094-4. https://books.google.com/books?id=2MQeh1KCp7sC&pg=PA240. 
  4. ↑ Chiappe, Luis M. (2007). Glorified Dinosaurs: The Origin and Early Evolution of Birds. Hoboken, New Jersey: John Wiley and Sons. ISBN 978-0-471-24723-4. [page needed]
  5. ↑ O'Connor, Jingmai K.; Chiappe, Luis M.; Gao, Chunling; Zhao, Bo (September 2011). "Anatomy of the Early Cretaceous Enantiornithine Bird Rapaxavis pani". Acta Palaeontologica Polonica 56 (3): 463–475. doi:10.4202/app.2010.0047. 
  6. ↑ Elzanowski, Andrzej (1974). "Preliminary note on the palaeognathous bird from the Upper Cretaceous of Mongolia". Palaeontologia Polonica 29: 103–9. http://www.palaeontologia.pan.pl/Archive/1974_30_103-109_32-33.pdf. 
  7. ↑ 7.0 7.1 Walker, C.A. (1981). "New subclass of birds from the Cretaceous of South America". Nature 292 (5818): 51–3. doi:10.1038/292051a0. Bibcode: 1981Natur.292...51W. 
  8. ↑ Hope, Sylvia (2002). "The Mesozoic Radiation of Neornithes". in Chiappe, Luis M.; Witmer, Lawrence M.. Mesozoic Birds: Above the Heads of Dinosaurs. University of California Press. pp. 339–88. ISBN 978-0-520-20094-4. 
  9. ↑ Mayr, G. (2021). "The coracoscapular joint of neornithine birds—extensive homoplasy in a widely neglected articular surface of the avian pectoral girdle and its possible functional correlates". Zoomorphology 140 (2): 217–28. doi:10.1007/s00435-021-00528-2. 
  10. ↑ Harris, Jerald D.; Lamanna, Matthew C.; You, Hai-lu; Ji, Shu-an; Ji, Qiang (18 May 2006). "A second enantiornithean (Aves: Ornithothoraces) wing from the Early Cretaceous Xiagou Formation near Changma, Gansu Province, People's Republic of China". Canadian Journal of Earth Sciences 43 (5): 547–554. doi:10.1139/e06-007. Bibcode: 2006CaJES..43..547H. 
  11. ↑ You, Hai-lu; Lamanna, Matthew C.; Harris, Jerald D.; Chiappe, Luis M.; O'Connor, Jingmai; Ji, Shu-an; Lü, Jun-chang; Yuan, Chong-xi et al. (16 June 2006). "A Nearly Modern Amphibious Bird from the Early Cretaceous of Northwestern China". Science 312 (5780): 1640–1643. doi:10.1126/science.1126377. PMID 16778053. Bibcode: 2006Sci...312.1640Y. 
  12. ↑ Pierce Brodkorb (1976). Discovery of a Cretaceous bird, apparently ancestral to the orders Coraciiformes and Piciformes (Aves: Carinatae) [File size 70 MB]. 27. Smithsonian Contributions to Paleobiology. pp. 67–73. http://www.sil.si.edu/smithsoniancontributions/Paleobiology/pdf_hi/SCtP-0027.pdf. 
  13. ↑ L. A. Nesov, A. A. Yarkov (1989). Новые птицы мел-палеогена СССР и некоторые замечания по истории возникновения и эволюции класса (New Cretaceous-Paleogene birds of USSR and some remarks about history of the origin and evolution of class). 197. USSR Academy of scienses. p. 82. http://museionsajtyarkova.ru/downloads/publication/Novye%20pticy%20mela-paleogena.pdf. 
  14. ↑ 14.0 14.1 Xing, Lida; O'Connor, Jingmai K.; McKellar, Ryan C.; Chiappe, Luis M.; Tseng, Kuowei; Li, Gang; Bai, Ming (September 2017). "A mid-Cretaceous enantiornithine (Aves) hatchling preserved in Burmese amber with unusual plumage". Gondwana Research 49: 264–277. doi:10.1016/j.gr.2017.06.001. Bibcode: 2017GondR..49..264X. 
  15. ↑ 15.0 15.1 Xing, Lida; O'Connor, Jingmai K.; McKellar, Ryan C.; Chiappe, Luis M.; Bai, Ming; Tseng, Kuowei; Zhang, Jie; Yang, Haidong et al. (February 2018). "A flattened enantiornithine in mid-Cretaceous Burmese amber: morphology and preservation". Science Bulletin 63 (4): 235–243. doi:10.1016/j.scib.2018.01.019. PMID 36659012. Bibcode: 2018SciBu..63..235X. 
  16. ↑ 16.0 16.1 16.2 Xing, Lida; McKellar, Ryan C.; Wang, Min; Bai, Ming; O'Connor, Jingmai K.; Benton, Michael J.; Zhang, Jianping; Wang, Yan et al. (28 June 2016). "Mummified precocial bird wings in mid-Cretaceous Burmese amber". Nature Communications 7 (1): 12089. doi:10.1038/ncomms12089. PMID 27352215. Bibcode: 2016NatCo...712089X. 
  17. ↑ 17.0 17.1 Xing, Lida; McKellar, Ryan C.; O'Connor, Jingmai K.; Bai, Ming; Tseng, Kuowei; Chiappe, Luis M. (30 January 2019). "A fully feathered enantiornithine foot and wing fragment preserved in mid-Cretaceous Burmese amber". Scientific Reports 9 (1): 927. doi:10.1038/s41598-018-37427-4. PMID 30700773. Bibcode: 2019NatSR...9..927X. 
  18. ↑ Xing, Lida; McKellar, Ryan C.; O'Connor, Jingmai K. (June 2020). "An unusually large bird wing in mid-Cretaceous Burmese amber". Cretaceous Research 110: 104412. doi:10.1016/j.cretres.2020.104412. Bibcode: 2020CrRes.11004412X. 
  19. ↑ Xing, Lida; O'Connor, Jingmai K.; Chiappe, Luis M.; McKellar, Ryan C.; Carroll, Nathan; Hu, Han; Bai, Ming; Lei, Fumin (2019-07-22). "A New Enantiornithine Bird with Unusual Pedal Proportions Found in Amber" (in English). Current Biology 29 (14): 2396–2401.e2. doi:10.1016/j.cub.2019.05.077. ISSN 0960-9822. PMID 31303484. https://www.cell.com/current-biology/abstract/S0960-9822(19)30691-8. 
  20. ↑ 20.0 20.1 Zhang, Zihui; Chiappe, Luis M.; Han, Gang; Chinsamy, Anusuya (2013). "A large bird from the Early Cretaceous of China: new information on the skull of enantiornithines". Journal of Vertebrate Paleontology 33 (5): 1176–89. doi:10.1080/02724634.2013.762708. Bibcode: 2013JVPal..33.1176Z. 
  21. ↑ Zhou, Zhonghe; Clarke, Julia; Zhang, Fucheng (May 2008). "Insight into diversity, body size and morphological evolution from the largest Early Cretaceous enantiornithine bird". Journal of Anatomy 212 (5): 565–77. doi:10.1111/j.1469-7580.2008.00880.x. PMID 18397240. 
  22. ↑ Hu, Dongyu; Xu, Xing; Hou, Lianhai; Sullivan, Corwin (2012). "A New Enantiornithine Bird from the Lower Cretaceous of Western Liaoning, China, and Its Implications for Early Avian Evolution". Journal of Vertebrate Paleontology 32 (3): 639–45. doi:10.1080/02724634.2012.652321. Bibcode: 2012JVPal..32..639H. 
  23. ↑ Atterholt, Jessie; Hutchison, J. Howard; O'Connor, Jingmai K. (13 November 2018). "The most complete enantiornithine from North America and a phylogenetic analysis of the Avisauridae". PeerJ 6: e5910. doi:10.7717/peerj.5910. PMID 30479894. 
  24. ↑ Martin, Anthony J.; Vickers-Rich, Patricia; Rich, Thomas H.; Hall, Michael; Angielczyk, Kenneth (January 2014). "Oldest known avian footprints from Australia: Eumeralla Formation (Albian), Dinosaur Cove, Victoria". Palaeontology 57 (1): 7–19. doi:10.1111/pala.12082. Bibcode: 2014Palgy..57....7M. 
  25. ↑ Wang, Min; Zhou, Zhonghe; Xu, Guanghui (2014-01-01). "The first enantiornithine bird from the Upper Cretaceous of China". Journal of Vertebrate Paleontology 34 (1): 135–145. doi:10.1080/02724634.2013.794814. ISSN 0272-4634. Bibcode: 2014JVPal..34..135W. 
  26. ↑ 26.0 26.1 Carvalho, Ismar; Novas, Fernando; Agnolin, Federico; Isasi, Marcelo; Freitas, Francisco; Andrade, José (2015-06-05). "A new genus and species of enantiornithine bird from the Early Cretaceous of Brazil". Brazilian Journal of Geology 45 (2): 161–171. doi:10.1590/23174889201500020001. https://www.researchgate.net/publication/279516967. 
  27. ↑ 27.0 27.1 O'Connor, Jingmai K.; Chiappe, Luis M. (28 February 2011). "A revision of enantiornithine (Aves: Ornithothoraces) skull morphology". Journal of Systematic Palaeontology 9 (1): 135–157. doi:10.1080/14772019.2010.526639. Bibcode: 2011JSPal...9..135O. 
  28. ↑ 28.0 28.1 28.2 Wang, Min; Hu, Han; Li, Zhiheng (21 August 2015). "A new small enantiornithine bird from the Jehol Biota, with implications for early evolution of avian skull morphology". Journal of Systematic Palaeontology 14 (6): 481–497. doi:10.1080/14772019.2015.1073801. 
  29. ↑ Wang, Min; Hu, Han (January 2017). "A Comparative Morphological Study of the Jugal and Quadratojugal in Early Birds and Their Dinosaurian Relatives". The Anatomical Record 300 (1): 62–75. doi:10.1002/ar.23446. PMID 28000410. 
  30. ↑ Chiappe, Luis M. (2009). "Downsized Dinosaurs: The Evolutionary Transition to Modern Birds". Evolution: Education and Outreach 2 (2): 248–56. doi:10.1007/s12052-009-0133-4. 
  31. ↑ Becker, Rachel (28 June 2016). "Bird wings trapped in amber are a fossil first from the age of dinosaurs". Nature. doi:10.1038/nature.2016.20162. 
  32. ↑ 32.0 32.1 Zhang, Fucheng; Zhou, Zhonghe (October 2004). "Palaeontology: Leg feathers in an Early Cretaceous bird". Nature 431 (7011): 925. doi:10.1038/431925a. PMID 15496911. Bibcode: 2004Natur.431..925Z. 
  33. ↑ 33.0 33.1 Clarke, Julia A.; Zhou, Zhonghe; Zhang, Fucheng (March 2006). "Insight into the evolution of avian flight from a new clade of Early Cretaceous ornithurines from China and the morphology of Yixianornis grabaui". Journal of Anatomy 208 (3): 287–308. doi:10.1111/j.1469-7580.2006.00534.x. PMID 16533313. 
  34. ↑ 34.0 34.1 O'Connor, Jingmai K.; Wang, Xuri; Chiappe, Luis M.; Gao, Chunling; Meng, Qingjin; Cheng, Xiaodong; Liu, Jinyuan (12 March 2009). "Phylogenetic support for a specialized clade of Cretaceous enantiornithine birds with information from a new species". Journal of Vertebrate Paleontology 29 (1): 188–204. doi:10.1080/02724634.2009.10010371. Bibcode: 2009JVPal..29..188O. 
  35. ↑ 35.0 35.1 Chiappe, Luis M.; Bo, Zhao; O'Connor, Jingmai K.; Chunling, Gao; Xuri, Wang; Habib, Michael; Marugan-Lobon, Jesus; Qingjin, Meng et al. (2014). "A new specimen of the Early Cretaceous bird Hongshanornis longicresta: insights into the aerodynamics and diet of a basal ornithuromorph". PeerJ 2: e234. doi:10.7717/peerj.234. PMID 24482756. 
  36. ↑ 36.0 36.1 36.2 36.3 O'Connor, Jingmai K.; Wang, Xiaoli; Zheng, Xiaoting; Hu, Han; Zhang, Xiaomei; Zhou, Zhonghe (January 2016). "An Enantiornithine with a Fan-Shaped Tail, and the Evolution of the Rectricial Complex in Early Birds". Current Biology 26 (1): 114–119. doi:10.1016/j.cub.2015.11.036. PMID 26748849. 
  37. ↑ 37.0 37.1 O'Connor, Jingmai K.; Li, Da-Qing; Lamanna, Matthew C.; Wang, Min; Harris, Jerald D.; Atterholt, Jessie; You, Hai-Lu (30 December 2015). "A new Early Cretaceous enantiornithine (Aves, Ornithothoraces) from northwestern China with elaborate tail ornamentation". Journal of Vertebrate Paleontology 36 (1): e1054035. doi:10.1080/02724634.2015.1054035. 
  38. ↑ 38.0 38.1 Wang, Min; O'Connor, Jingmai K.; Pan, Yanhong; Zhou, Zhonghe (2017-01-31). "A bizarre Early Cretaceous enantiornithine bird with unique crural feathers and an ornithuromorph plough-shaped pygostyle" (in En). Nature Communications 8: 14141. doi:10.1038/ncomms14141. PMID 28139644. Bibcode: 2017NatCo...814141W. 
  39. ↑ 39.0 39.1 Sanz, José L.; Chiappe, Luis M.; Buscalioni, Angela D. (1995). "The Osteology of Concornis lacustris (Aves: Enantiornithes) from the Lower Cretaceous of Spain and a Reexamination of its Phylogenetic Relationships". American Museum Novitates (3133): 1–23. 
  40. ↑ Dalla Vecchia, Fabio M.; Chiappe, Luis M. (2003). "First avian skeleton from the Mesozoic of northern Gondwana". Journal of Vertebrate Paleontology 22 (4): 856–60. doi:10.1671/0272-4634(2002)022[0856:FASFTM2.0.CO;2]. 
  41. ↑ O'Connor, Jingmai K. (1 January 2019). "The trophic habits of early birds". Palaeogeography, Palaeoclimatology, Palaeoecology 513: 178–195. doi:10.1016/j.palaeo.2018.03.006. Bibcode: 2019PPP...513..178O. 
  42. ↑ O'Connor, Jingmai K.; Zhou, Zhonghe; Smith, Andrew (6 November 2019). "The evolution of the modern avian digestive system: insights from paravian fossils from the Yanliao and Jehol biotas". Palaeontology 63 (1): 13–27. doi:10.1111/pala.12453. 
  43. ↑ Houston, David C.; Copsey, J. A. (1994). "Bone digestion and intestinal morphology of the Bearded Vulture". The Journal of Raptor Research 28 (2): 73–78. https://www.biodiversitylibrary.org/page/51322199. 
  44. ↑ "Short Crystal: Quartz and the Fossilized Bird | GeoRarities" (in en). 2021-03-12. https://georarities.com/2021/03/12/short-crystal-quartz-and-the-fossilized-bird/. 
  45. ↑ O’Connor, Jingmai K.; Zhou, Zhonghe; Zhang, Fucheng (28 February 2011). "A reappraisal of Boluochia zhengi (Aves: Enantiornithes) and a discussion of intraclade diversity in the Jehol avifauna, China". Journal of Systematic Palaeontology. 9 (1): 51–63. doi:10.1080/14772019.2010.512614. S2CID 84817636.
  46. ↑ Lianhai Hou; LuisM. Chiappe; Fucheng Zhang; Cheng-Ming Chuong (2004). "New Early Cretaceous fossil from China documents a novel trophic specialization for Mesozoic birds". Naturwissenschaften. 91 (1): 22–25. Bibcode:2004NW.....91...22H. doi:10.1007/s00114-003-0489-1. PMC 4382005. PMID 14740099.
  47. ↑ Morschhauser, E. M.; Varricchio, D.J.; Gao, C.; Liu, J.; Wang, Z.; Cheng, X. & Meng, Q. (2009). "Anatomy of the Early Cretaceous bird Rapaxavis pani, a new species from Liaoning Province, China". Journal of Vertebrate Paleontology. 29 (2): 545–554. doi:10.1671/039.029.0210. S2CID 84643293.
  48. ↑ Miller, Case Vincent; Pittman, Michael; Wang, Xiaoli; Zheng, Xiaoting; Bright, Jen A. (2022). "Diet of Mesozoic toothed birds (Longipterygidae) inferred from quantitative analysis of extant avian diet proxies". BMC Biology 20 (1): 101. doi:10.1186/s12915-022-01294-3. PMID 35550084. 
  49. ↑ Sanz, José L.; Chiappe, Luis M.; Fernádez-Jalvo, Yolanda; Ortega, Francisco; Sánchez-Chillón, Begoña; Poyato-Ariza1, Francisco J.; Pérez-Moreno, Bernardino P. (February 2001). "An early Cretaceous pellet". Nature 409 (6823): 998–1000. doi:10.1038/35059172. PMID 11234054. Bibcode: 2001Natur.409..998S. 
  50. ↑ Mikhailov, Konstantin E. (1991). "Classification of fossil eggshells of amniotic vertebrates". Acta Palaeontologica Polonica 36 (2): 193–238. https://www.app.pan.pl/archive/published/app36/app36-193.pdf. 
  51. ↑ Mikhailov, Konstantin E. (1996). "New Genera of Fossil Eggs from the Upper Cretaceous of Mongolia". Paleontological Journal 30 (2): 246–8. 
  52. ↑ Elżanowski, Andrzej (1981). "Embryonic bird skeletons from the late Cretaceous of Mongolia". Palaeontologia Polonica 42: 147–79. http://www.palaeontologia.pan.pl/Archive/1981-42_147-179_40-45.pdf. 
  53. ↑ Sanz, José L.; Chiappe, Luis M.; Pérez-Moreno, Bernardino P.; Moratalla, José J.; Hernández-Carrasquilla, Francisco; Buscalioni, Angela D.; Ortega, Francisco; Poyato-Ariza, Francisco J. et al. (June 6, 1997). "A Nestling Bird from the Lower Cretaceous of Spain: Implications for Avian Skull and Neck Evolution". Science 276 (5318): 1543–6. doi:10.1126/science.276.5318.1543. 
  54. ↑ Zhou, Zhonghe; Zhang, Fucheng (October 22, 2004). "A Precocial Avian Embryo from the Lower Cretaceous of China". Science 306 (5696): 653. doi:10.1126/science.1100000. PMID 15499011. 
  55. ↑ 55.0 55.1 55.2 Chiappe, Luis M.; Shu'an, Ji; Qiang, Ji (2007). "Juvenile Birds from the Early Cretaceous of China: Implications for Enantiornithine Ontogeny". American Museum Novitates (3594): 1–46. doi:10.1206/0003-0082(2007)3594[1:JBFTEC2.0.CO;2]. https://www.biodiversitylibrary.org/bibliography/169246. 
  56. ↑ Elżanowski, Andrzej (1995). "Cretaceous birds and avian phylogeny". Courier Forschungsinstitut Senckenberg 181: 37–53. 
  57. ↑ Kurochkin, E. N.; Chatterjee, S.; Mikhailov, K. E. (December 2013). "An embryonic enantiornithine bird and associated eggs from the cretaceous of Mongolia". Paleontological Journal 47 (11): 1252–1269. doi:10.1134/S0031030113110087. Bibcode: 2013PalJ...47.1252K. 
  58. ↑ Kurochkin, E. N.; Chatterjee, S.; Mikhailov, K. E. (19 December 2013). "An embryonic enantiornithine bird and associated eggs from the cretaceous of Mongolia". Paleontological Journal 47 (11): 1252–1269. doi:10.1134/S0031030113110087. Bibcode: 2013PalJ...47.1252K. 
  59. ↑ Dinosaur incubation periods directly determined from growth-line counts in embryonic teeth show reptilian-grade development - PNAS
  60. ↑ The origin of the bird's beak: new insights from dinosaur incubation periods
  61. ↑ Cambra-Moo, Oscar; Buscalioni, Ángela Delgado; Cubo, Jorge; Castanet, Jacques; Loth, Marie-Madeleine; de Margerie, Emmanuel; de Ricqlès, Armand (2006). "Histological observations of Enantiornithine bone (Saurischia, Aves) from the Lower Cretaceous of Las Hoyas (Spain)". Comptes Rendus Palevol 5 (5): 685–91. doi:10.1016/j.crpv.2005.12.018. Bibcode: 2006CRPal...5..685C. 
  62. ↑ O'Connor, Jingmai K.; Wang, Min; Zheng, Xiao-Ting; Wang, Xiao-Li; Zhou, Zhong-He (2014). "The histology of two female Early Cretaceous birds". Vertebrata PalAsiatica 52 (1): 112–28. http://www.ivpp.cas.cn/cbw/gjzdwxb/xbwzxz/201401/P020140121384211870999.pdf. 
  63. ↑ Chiappe, L.M. (1995). "The phylogenetic position of the Cretaceous birds of Argentina: Enantiornithes and Patagopteryx deferrariisi". Courier Forschungsinstitut Senckenberg 181: 55–63. 
  64. ↑ Cubo, Jorge; Buscalioni, Angela D.; Legendre, Lucas J.; Bourdon, Estelle; Sanz, Jose L.; Ricqlès, Armand (2021). "Palaeohistological inferences of resting metabolic rates in Concornis and Iberomesornis (Enantiornithes, Ornithothoraces) from the Lower Cretaceous of las Hoyas (Spain)". Palaeontology 65. doi:10.1111/pala.12583. https://onlinelibrary.wiley.com/doi/10.1111/pala.12583. 
  65. ↑ Dyke, Gareth; Vremir, Mátyás; Kaiser, Gary; Naish, Darren (June 2012). "A drowned Mesozoic bird breeding colony from the Late Cretaceous of Transylvania". Die Naturwissenschaften 99 (6): 435–42. doi:10.1007/s00114-012-0917-1. PMID 22575918. Bibcode: 2012NW.....99..435D. 
  66. ↑ Fernández, Mariela S.; García, Rodolfo A.; Fiorelli, Lucas; Scolaro, Alejandro; Salvador, Rodrigo B.; Cotaro, Carlos N.; Kaiser, Gary W.; Dyke, Gareth J. et al. (17 April 2013). "A Large Accumulation of Avian Eggs from the Late Cretaceous of Patagonia (Argentina) Reveals a Novel Nesting Strategy in Mesozoic Birds". PLOS ONE 8 (4): e61030. doi:10.1371/journal.pone.0061030. PMID 23613776. Bibcode: 2013PLoSO...861030F. 
  67. ↑ O'Connor, Jingmai K.; Falk, Amanda; Wang, Min; Zheng, Xiao-Ting (2020). "First report of immature feathers in juvenile Enantiornithes from the Early Cretaceous Jehol avifauna". Vertebrata PalAsiatica 58: 24–44. doi:10.19615/j.cnki.1000-3118.190823. 
  68. ↑ Clark, Alexander D.; O'Connor, Jingmai K. (15 June 2021). "Exploring the Ecomorphology of Two Cretaceous Enantiornithines With Unique Pedal Morphology". Frontiers in Ecology and Evolution 9: 654156. doi:10.3389/fevo.2021.654156. 
  69. ↑ Kaye, Thomas G.; Pittman, Michael; Marugán-Lobón, Jesús; Martín-Abad, Hugo; Sanz, José Luis; Buscalioni, Angela D. (21 March 2019). "Fully fledged enantiornithine hatchling revealed by Laser-Stimulated Fluorescence supports precocial nesting behavior". Scientific Reports 9 (1): 5006. doi:10.1038/s41598-019-41423-7. PMID 30899080. Bibcode: 2019NatSR...9.5006K. 
  70. ↑ Padian, Kevin; Chiappe, Luis M. (11 January 2007). "The origin and early evolution of birds". Biological Reviews 73 (1): 1–42. doi:10.1111/j.1469-185x.1997.tb00024.x. http://doc.rero.ch/record/16121/files/PAL_E3214.pdf. 
  71. ↑ Zhou, Shuang; Zhou, Zhong-He; O'Connor, Jingmai K. (2012). "A new basal beaked ornithurine bird from the Lower Cretaceous of Western Liaoning, China". Vertebrata PalAsiatica 50 (1): 9–24. http://www.ivpp.cas.cn/cbw/gjzdwxb/xbwzxz/201202/P020120209570168040733.pdf. 
  72. ↑ Houde, Peter W. (1988). "Paleognathous Birds from the Early Tertiary of the Northern Hemisphere". Publications of the Nuttall Ornithological Club (Cambridge Massachusetts, USA: Nuttall Ornithological Club) 22
  73. ↑ Navalón, Guillermo; Marugán-Lobón, Jesús; Chiappe, Luis M.; Luis Sanz, José; Buscalioni, Ángela D. (6 October 2015). "Soft-tissue and dermal arrangement in the wing of an Early Cretaceous bird: Implications for the evolution of avian flight". Scientific Reports 5 (1): 14864. doi:10.1038/srep14864. PMID 26440221. Bibcode: 2015NatSR...514864N. 
  74. ↑ Brusatte, Stephen L.; O'Connor, Jingmai K.; Jarvis, Erich D. (5 October 2015). "The Origin and Diversification of Birds". Current Biology 25 (19): R888–898. doi:10.1016/j.cub.2015.08.003. ISSN 1879-0445. PMID 26439352. https://pubmed.ncbi.nlm.nih.gov/26439352/. Retrieved 12 November 2023. 
  75. ↑ Atterholt, Jessie; Hutchison, J. Howard; O'Connor, Jingmai K. (2018). "The most complete enantiornithine from North America and a phylogenetic analysis of the Avisauridae". PeerJ 6: e5910. doi:10.7717/peerj.5910. ISSN 2167-8359. PMID 30479894. 
  76. ↑ Wang, Xia; McGowan, Alistair J.; Dyke, Gareth J.; Turvey, Samuel T. (7 December 2011). "Avian Wing Proportions and Flight Styles: First Step towards Predicting the Flight Modes of Mesozoic Birds". PLOS ONE 6 (12): e28672. doi:10.1371/journal.pone.0028672. PMID 22163324. Bibcode: 2011PLoSO...628672W. 
  77. ↑ Chiappe, Luis M.; Suzuki, Shigeru; Dyke, Gareth J.; Watabe, Mahito; Tsogtbaatar, K.; Barsbold, Rinchen (January 2007). "A new Enantiornithine bird from the Late Cretaceous of the Gobi desert". Journal of Systematic Palaeontology 5 (2): 193–208. doi:10.1017/S1477201906001969. Bibcode: 2007JSPal...5..193C. 
  78. ↑ Zheng, Xiaoting; Wang, Xiaoli; O'Connor, Jingmai; Zhou, Zhonghe (9 October 2012). "Insight into the early evolution of the avian sternum from juvenile enantiornithines". Nature Communications 3 (1): 1116. doi:10.1038/ncomms2104. PMID 23047674. Bibcode: 2012NatCo...3.1116Z. 
  79. ↑ 79.0 79.1 O'Connor, Jingmai; Dyke, Gareth (2010). "A Reassessment of Sinornis santensis and Cathayornis yandica (Aves: Enantiornithes)". Records of the Australian Museum 62: 7–20. doi:10.3853/j.0067-1975.62.2010.1540. https://zenodo.org/record/4676605. 
  80. ↑ Clarke, Julia A.; Norell, Mark A. (2002). "The Morphology and Phylogenetic Position of Apsaravis ukhaana from the Late Cretaceous of Mongolia". American Museum Novitates (3387): 1–46. doi:10.1206/0003-0082(2002)387<0001:TMAPPO>2.0.CO;2. 
  81. ↑ 81.0 81.1 Sereno, P.C. (2005) TaxonSearch: Stem Archosauria . Version 1.0, 2005-NOV- 7. Retrieved 2006-OCT-02.
  82. ↑ Wang, Xuri; Cau, Andrea; Luo, Xiaoling; Kundrát, Martin; Wu, Wensheng; Ju, Shubin; Guo, Zhen; Liu, Yichuan et al. (2022-02-11). "A new bohaiornithid-like bird from the Lower Cretaceous of China fills a gap in enantiornithine disparity" (in en). Journal of Paleontology 96 (4): 961–976. doi:10.1017/jpa.2022.12. ISSN 0022-3360. Bibcode: 2022JPal...96..961W. https://www.cambridge.org/core/journals/journal-of-paleontology/article/new-bohaiornithidlike-bird-from-the-lower-cretaceous-of-china-fills-a-gap-in-enantiornithine-disparity/904F4A5175D5E6A5A641116267945BA5. 
  83. ↑ Carvalho; Novas; Agnolín; Isasi; Freitas; Andrade (2015). "A new genus and species of enantiornithine bird from the Early Cretaceous of Brazil". Brazilian Journal of Geology 45 (2): 161–171. doi:10.1590/23174889201500020001. 
  84. ↑ Wang, Li; O'Connor, Zhou; You (2015). "Second species of enantiornithine bird from the Lower Cretaceous Changma Basin, northwestern China with implications for the taxonomic diversity of the Changma avifauna". Cretaceous Research 55: 56–65. doi:10.1016/j.cretres.2015.01.008. Bibcode: 2015CrRes..55...56W. 
  85. ↑ Wang, M.; O'Connor, J. K.; Zhou, Z. (2014). "A new robust enantiornithine bird from the Lower Cretaceous of China with scansorial adaptations". Journal of Vertebrate Paleontology 34 (3): 657–671. doi:10.1080/02724634.2013.812101. Bibcode: 2014JVPal..34..657W. 
  86. ↑ Dalsätt, J.; Ericson, P. G.; Zhou, Z. (2015). "A New Enantiornithes (Aves) from the Early Cretaceous of China". Acta Geologica Sinica 86 (2): 801–807. doi:10.1111/1755-6724.12270. 
  87. ↑ Bell, Alyssa K.; Chiappe, Luis M.; Erickson, Gregory M.; Suzuki, Shigeru; Watabe, Mahito; Barsbold, Rinchen; Tsogtbaatar, K. (February 2010). "Description and ecologic analysis of Hollanda luceria, a Late Cretaceous bird from the Gobi Desert (Mongolia)". Cretaceous Research 31 (1): 16–26. doi:10.1016/j.cretres.2009.09.001. Bibcode: 2010CrRes..31...16B. 
  88. ↑ Hartman, Scott; Mortimer, Mickey; Wahl, William R.; Lomax, Dean R.; Lippincott, Jessica; Lovelace, David M. (10 July 2019). "A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight". PeerJ 7: e7247. doi:10.7717/peerj.7247. PMID 31333906. 
  89. ↑ Zelenkov, Nikita V.; Averianov, Alexander O. (13 June 2015). "A historical specimen of enantiornithine bird from the Early Cretaceous of Mongolia representing a new taxon with a specialized neck morphology". Journal of Systematic Palaeontology 14 (4): 319–338. doi:10.1080/14772019.2015.1051146. 
  90. ↑ Wang, M.; Liu, D. (2015). "Taxonomical reappraisal of Cathayornithidae (Aves: Enantiornithes)". Journal of Systematic Palaeontology 14: 1–19. doi:10.1080/14772019.2014.994087. 
  91. ↑ Liu, Di; Chiappe, Luis M.; Serrano, Francisco; Habib, Michael; Zhang, Yuguang; Meng, Qinjing; Shawkey, Matthew (11 October 2017). "Flight aerodynamics in enantiornithines: Information from a new Chinese Early Cretaceous bird". PLOS ONE 12 (10): e0184637. doi:10.1371/journal.pone.0184637. PMID 29020077. Bibcode: 2017PLoSO..1284637L. 
  92. ↑ Hu, Han; O'Connor, Jingmai K. (14 November 2016). "First species of Enantiornithes from Sihedang elucidates skeletal development in Early Cretaceous enantiornithines". Journal of Systematic Palaeontology 15 (11): 909–926. doi:10.1080/14772019.2016.1246111. 
  93. ↑ Wang, Min; Zhou, Zhonghe; Xu, Guanghui (7 January 2014). "The first enantiornithine bird from the Upper Cretaceous of China". Journal of Vertebrate Paleontology 34 (1): 135–145. doi:10.1080/02724634.2013.794814. Bibcode: 2014JVPal..34..135W. 
  94. ↑ Wang, Min; Zhou, Zhonghe (12 April 2017). "A morphological study of the first known piscivorous enantiornithine bird from the Early Cretaceous of China". Journal of Vertebrate Paleontology 37 (2): e1278702. doi:10.1080/02724634.2017.1278702. Bibcode: 2017JVPal..37E8702W. 
  95. ↑ Herrera, Gerardo Álvarez; Agnolín, Federico; Rozadilla, Sebastián; Lo Coco, Gastón E.; Manabe, Makoto; Tsuihiji, Takanobu; Novas, Fernando E. (2022-12-16). "New enantiornithine bird from the uppermost Cretaceous (Maastrichtian) of southern Patagonia, Argentina" (in en). Cretaceous Research 144: 105452. doi:10.1016/j.cretres.2022.105452. ISSN 0195-6671. https://www.sciencedirect.com/science/article/pii/S0195667122003160. 
  96. ↑ Hu, Dongyu; Liu, Ying; Li, Jinhua; Xu, Xing; Hou, Lianhai (July 2015). "Yuanjiawaornis viriosus, gen. et sp. nov., a large enantiornithine bird from the Lower Cretaceous of western Liaoning, China". Cretaceous Research 55: 210–219. doi:10.1016/j.cretres.2015.02.013. Bibcode: 2015CrRes..55..210H. 
  97. ↑ O'Connor, Patrick M.; Turner, Alan H.; Groenke, Joseph R.; Felice, Ryan N.; Rogers, Raymond R.; Krause, David W.; Rahantarisoa, Lydia J. (10 December 2020). "Late Cretaceous bird from Madagascar reveals unique development of beaks". Nature 588 (7837): 272–276. doi:10.1038/s41586-020-2945-x. PMID 33239782. Bibcode: 2020Natur.588..272O. 
  98. ↑ Hu, Han; O'Connor, Jingmai K.; Zhou, Zhonghe; Farke, Andrew A. (3 June 2015). "A New Species of Pengornithidae (Aves: Enantiornithes) from the Lower Cretaceous of China Suggests a Specialized Scansorial Habitat Previously Unknown in Early Birds". PLOS ONE 10 (6): e0126791. doi:10.1371/journal.pone.0126791. PMID 26039693. Bibcode: 2015PLoSO..1026791H. 
  99. ↑ Chiappe, Luis M.; Qingjin, Meng; Serrano, Francisco; Sigurdsen, Trond; Min, Wang; Bell, Alyssa; Di, Liu (25 October 2019). "New Bohaiornis-like bird from the Early Cretaceous of China: enantiornithine interrelationships and flight performance". PeerJ 7: e7846. doi:10.7717/peerj.7846. PMID 31667014. 
  100. ↑ Wang, Yan; Wang, Min; O'Connor, Jingmai K.; Wang, Xiaoli; Zheng, Xiaoting; Zhang, Xiaomei (11 January 2016). "A new Jehol enantiornithine bird with three-dimensional preservation and ovarian follicles". Journal of Vertebrate Paleontology 36 (2): e1054496. doi:10.1080/02724634.2015.1054496. Bibcode: 2016JVPal..36E4496W. 
  101. ↑ Kurochkin, E. N.; Chatterjee, S.; Mikhailov, K. E. (19 December 2013). "An embryonic enantiornithine bird and associated eggs from the cretaceous of Mongolia". Paleontological Journal 47 (11): 1252–1269. doi:10.1134/s0031030113110087. Bibcode: 2013PalJ...47.1252K. 

External links

Wikidata ☰ Q132268 entry



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