| Alvarezsaurids | |
|---|---|
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| Mounted holotype ofMononykus | |
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| Skeletal mount ofAlvarezsaurus | |
Scientific classification | |
| Kingdom: | Animalia |
| Phylum: | Chordata |
| Class: | Reptilia |
| Clade: | Dinosauria |
| Clade: | Saurischia |
| Clade: | Theropoda |
| Superfamily: | †Alvarezsauroidea |
| Family: | †Alvarezsauridae Bonaparte, 1991 |
| Type species | |
| †Alvarezsaurus calvoi Bonaparte, 1991 | |
| Subgroups | |
| Synonyms | |
Alvarezsauridae is afamily of small, long-leggeddinosaurs. Although originally thought to represent the earliest known flightlessbirds, they are now thought to be an early diverging branch ofmaniraptorantheropods. Alvarezsaurids were highly specialized. They had tiny but stout forelimbs, with compact, bird-like hands. Their skeletons suggest that they had massive breast and arm muscles, possibly adapted for digging or tearing. They had long, tube-shaped snouts filled with tiny teeth. They have been interpreted asmyrmecophagous, adapted to prey on colonial insects such astermites, with the short arms acting as effective digging instruments to break into nests.
Alvarezsaurus, thetype genus of the family, was named for thehistorianGregorio Álvarez.[1]
Bonaparte (1991) described the first alvarezsaurid,Alvarezsaurus calvoi, from an incomplete skeleton found in Patagonia, Argentina. Bonaparte also named afamily, Alvarezsauridae, to contain it. He argued thatAlvarezsaurus might be most closely related to theornithomimosaurs.[2]
In 1993, Perleet al. described the next alvarezsaur to be discovered, naming itMononychus olecranus (meaning "one claw"). A month later they changed the genus name toMononykus, because the earlier spelling was already the genus name of an extant beetle.[3] Perleet al. mistakenly describedMononykus as a member ofAvialae, one more advanced thanArchaeopteryx. They argued that the family Alvarezsauridae was actually a group of Mesozoic flightless birds on the basis of several features that were supposedly unique to birds.[4]
In 1996, Novas described another member of the group calledPatagonykus puertai.[5] Karhu and Rautian (1996) described a Mongolian member of the family;Parvicursor remotus.[6] Chiappeet al.(1998) described another Mongolian member,Shuvuuia deserti, and found it to be a bird as in Perle et al.'s analysis.[7]
These mistaken assignments of alvarezsaurids to birds were caused primarily by features that are strikingly, or even uniquely, avian. The sternum, for example, is elongated and deeply keeled for an enlarged pectoralis muscle, as it is inneognathous birds andvolantratites. One bone in the skull ofShuvuuia appeared to be an ectethmoid fused to a prefrontal. The ectethmoid is an ossification known only inNeornithes. Other birdlike characters included the palatine, foramen magnum, cervical and caudal vertebrae, and many others.[8]
Several researchers disagreed with Perleet al. (1993) and Chiappeet al. (1998), Feduccia (1994), Ostrom (1994), Wellnhofer (1994), Kurochkin (1995), Zhou (1995), and Sereno (1997) considered it unlikely that alvarezsaurids were members ofAvialae. Martin (1997) performed a cladistic analysis but Sereno criticized it strongly, finding it flawed by incorrect codings, use of only select data, and results that did not support his conclusions. Sereno (1999) performed a new analysis, revising the anatomical interpretations and clarifying the characters. He found that alvarezsaurids were more parsimoniously related to theOrnithomimosauria.[8]
As the more primitive members of the Alvarezsauridae were better characterized, the monophyly of the clade was strongly supported,[5] but the more primitive members lacked the most birdlike traits. Some of these traits had been misinterpreted as well. The remaining similarities between birds and alvarezsaurs—like the keeled sterna—are another case ofhomoplasy, where the derived alvarezsaurids developed birdlike characters through convergent evolution, rather than inheriting them from a common ancestor with birds.[8]
Alvarezsaurids ranged from 50 centimetres (20 in) to 2 metres (6.6 ft) in length, although some possible members may have been larger, including the EuropeanHeptasteornis that may have reached 2.5 metres (8.2 ft) long. Fossils attributed to alvarezsaurids have also been found inNorth andSouth America andAsia, and range in age from about 86 to 66 million years ago.[9]
At least one specimen of alvarezsaurid, from thespeciesShuvuuia deserti, preserved down-like,feathery,integumental structures covering the fossil. Schweitzeret al. (1999) subjected these filaments to microscopic, morphological, mass spectrometric, and immunohistochemical studies and found that they consisted ofbeta-keratin, which is the primary protein infeathers.[10]
The lifestyle of alvarezsaurids has been debated since the nature of these dinosaurs was established. It has been suggested by numerous palaeontologists that they used their claws to break into ant and termite colonies, though the arm anatomy of an alvarezsaurid would require the animal to lie on its chest against a termite nest. It is also possible that the alvarezsaurids filled some ecological niche that has not yet been considered.[11] Studies of the tails in various alvarezsaur genera also suggest they were possessed of an incredible ability to change their rotational inertia, and combined with their forelimbs, this suggests their ecological niches were similar to those of aardvarks, pangolins, and anteaters.[12]
Additionally, it is known that alvarezsaurids, with their long legs, appear to be built for speed. What implications this has on possible lifestyle is unknown.[11] The discovery ofQiupanykus in association withoviraptorid eggs, indicates that the advanced alvarezsaurids may also have been specialists in nest raiding, using their robust thumb claws to crack open eggshells.[13]
Turneret al. (2007) place the alvarezsaurs as the most basal group in theManiraptora, one step more derived thanOrnitholestes and two more derived than theOrnithomimosauria. The alvarezsaurs are more primitive than theOviraptorosauria.[14]
Novas' 1996 description ofPatagonykus, demonstrated that it was a link between the more primitive (basal)Alvarezsaurus and the more advanced (derived)Mononykus, and reinforced their monophyly.Parvicursor was discovered shortly after, and placed in its own family Parvicursoridae, and thenShuvuuia in 1998. Everything has since been lumped into Alvarezsauridae, with Mononykinae surviving as asubfamily.
There may be a relationship between the alvarezsaurids and theOrnithomimosauria as sister clades within eitherThomas Holtz'sArctometatarsalia orPaul Sereno'sOrnithomimiformes. The discovery ofHaplocheirus, which exhibits transitional features between the more derived alvarezsaurs and othermaniraptorans, particularly in relation to the skull structure and development of the hand, has provided further support for that relationship.[15]
The taxonomy of the alvarezsaurs has been somewhat confused, due to different authors using different names for groups with the same definition. Thefamily Alvarezsauridae was first coined by Jose Bonaparte in 1991, but given no specific phylogenetic definition. Novas later defined the group as the most recent common ancestor ofAlvarezsaurus andMononykus plus all its descendants, though others, such asPaul Sereno, used a more inclusive definition, such as all dinosaurs closer toShuvuuia than to modern birds. In 2009, Livezey and Zusi used the name Alvarezsauroidea for the total group of all alvarezsaurs, restricting the name Alvarezsauridae to the clade defined byAlvarezsaurus +Mononykus. This was followed by Choiniere and colleagues in 2010, who described the first non-alvarezsaurid alvarezsauroid,Haplocheirus.[16] Some authors have used the name Mononykinae for the sub-group of alvarezsaurs including the advanced Mongolian species. However, Choiniere and colleagues argued that Parvicursorinae has priority, since its coordinate name under theICZN Code, Parvicursoridae, was named earlier.[16] Another subfamily, Patagonykinae, has been named to include the South AmericanPatagonykus andBonapartenykus, but a few recent studies have placed them just outside Alvarezsauridae, some of which do not even recover them in a single clade, making Patagonykinae turn out to be paraphyletic.[17][18][19][20][21][22]
The cladogram below is the latest and most comprehensive analysis of alvarezsaurid evolution, following Meso et al. (2024).[23]
| Alvarezsauridae |
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