Rahonavis

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Rahonavis
Temporal range: Late Cretaceous (Maastrichtian) 72.1–66  Ma
Maniraptoran ROM.jpg
Reconstructed skeleton, Royal Ontario Museum
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Clade: Saurischia
Clade: Theropoda
Clade: Paraves
Genus: Rahonavis
Forster et al., 1998b
Type species
Rahonavis ostromi
(Forster et al., 1998a) Forster et al., 1998b
Synonyms
  • Rahona Forster et al. 1998 non Griveaud 1975
  • Rahona ostromiForster et al. 1998a

Rahonavis is a genus of bird-like theropod from the Late Cretaceous (Maastrichtian, from about 72.1 to 66 mya) of what is now northwestern Madagascar. It is known from a partial skeleton (UA 8656) found by Catherine Forster and colleagues in Maevarano Formation rocks at a quarry near Berivotra, Mahajanga Province. [1] Rahonavis was a small predator, at about 70 centimetres (2.3 ft) long and 0.45-2.27 kg (1-5 lbs), [2] with the typical dromaesaurid-like raised sickle claw on the second toe. It was originally the first African coelurosaur until the discovery of Nqwebasaurus in 2000. [3]

Contents

The name Rahonavis means, approximately, "cloud menace bird", from Malagasy rahona (RA-hoo-na, "cloud" or "menace") + Latin avis "bird". The specific name, R. ostromi, was coined in honor of John Ostrom.

Discovery and species

Skeletal diagram showing remains of the holotype specimen Rahonavis.jpg
Skeletal diagram showing remains of the holotype specimen

The fossilized remains of Rahonavis were first recovered from the Maevarano Formation in Madagascar in 1995 by a joint expedition of SUNY and the University of Antananarivo, near the village of Berivotra. Most geological formations in this area are covered in dense grass, making identification of fossils difficult. However, when a portion of hillside was exposed by fire, the remains of a giant titanosaur were revealed. It was during the excavation of this find that paleontologists discovered the bones of Rahonavis among the bones of the much larger dinosaur. Rahonavis is known from this single specimen, consisting of the hind limbs, trunk, portions of the tail (all of which were found articulated), as well as portions of the wing and shoulder bones. Rahonavis was one-fifth larger than the closely related Archaeopteryx, about the size of a modern raven. [4]

The discoverers of Rahonavis initially named it Rahona but changed the name after discovering that the name Rahona was already assigned to a genus of lymantriid moths. [5] [6]

The lack of well-documented relatives of this species nonwithstanding, a single thoracic vertebra (NMC 50852) most similar to those of R. ostromi was found in the Albian to Cenomanian Kem Kem Beds in Morocco. Lacking the pleurocoels found in Rahonavis and having a larger neural canal, it appears to belong to a different genus. Although former character can vary in species of the same genus, in individual vertebrae of the same animal, and ontogenetically, the distance in space and time suggests that whatever this specimen may be, it does not belong into Rahonavis. [7]

A dentary has been found in association with the holotype, though it is seldom described. [8]

Classification

Size compared to a human hand Rahonavis Scale.svg
Size compared to a human hand

Rahonavis has historically been the subject of some uncertainty as to its proper taxonomic position – whether it is a member of the clade Avialae (birds) or a closely related dromaeosaurid. The presence of quill knobs on its ulna (forearm bone) led initially to its inclusion as an avialan; however, the rest of the skeleton is rather typically dromaeosaurid in its attributes. Given the extremely close affinities between basal birds and their dromaeosaurid cousins, along with the possibility that flight may have developed and been lost multiple times among these groups, it has been difficult to place Rahonavis firmly among or outside the birds.

Rahonavis could be a close relative to Archaeopteryx , as originally suggested by the describers, and thus a member of the clade Avialae, but while the pelvis shows adaptations to flight similar in function to those of Archaeopteryx, they seem to be independently derived. [9]

Beginning in the early 2000s, a consensus emerged among most theropod researchers that Rahonavis was more closely related to deinonychosaurs than to avialans, and specifically was a member of the South American dromaeosaurid clade Unenlagiinae. A 2005 analysis by Makovicky and colleagues found Rahonavis to be closely related to the unenlagiines Unenlagia and Buitreraptor . [10] Norell and colleagues (2006) also found Rahonavis to lie within the Unenlagiinae, as the sister taxon to Unenlagia itself. [11] A 2007 study by Turner and colleagues again found it to be an unenlagiine dromaeosaurid, closely related to Unenlagia. [12]

This consensus has been challenged, however, by a few studies published since 2009 that have found many traditional "dromaeosaurids", including the unenlagiines, closer to Avialae than to dromaeosaurines. A large analysis published by Agnolín and Novas (2013) recovered Rahonavis as closer to Avialae than to Dromaeosauridae. [13] A cladistic analysis by Cau (2018) recovered Rahonavis as a probable relative of the long-tailed Early Cretaceous avialans Jeholornis and Jixiangornis . [14] The analysis of Hartman et al. (2019) "strongly rejected" the supposed avialan position of Rahonavis, finding its placement in Unenlagiinae better supported as it takes 10 less steps. This placement suggests Rahonavis is one among multiple different paravian lineages that evolved flight independently. [15] In 2020, Rahonavis and the South American Overoraptor were found to be sister taxa in a clade sister to the Avialae with a dataset based on that of the 2013 study. [16] As of 2020, it is undecided among paleontologists whether the paravian Rahonavis is an unenlagiine, a dromaeosaurid or an avialan. [8]

Paleobiology

Hypothetical life restoration Rahonavis NT.jpg
Hypothetical life restoration

Although numerous artists' reconstructions of Rahonavis show it in flight, it is not clear that it could fly; there has even been some doubt that the forearm material, which includes the quill knobs, belongs with the rest of the skeleton. Some researchers have suggested that Rahonavis represents a chimera consisting of the forelimb of a bird conflated with the skeleton of a dromaeosaurid, and consider Rahona as described a nomen dubium . [9] The nearby discovery of the primitive bird Vorona berivotrensis at least shows that the possibility of a mix-up cannot be fully excluded. However, many other scientists, including the original describers of Rahonavis, maintain that its remains belong to a single animal, citing the close proximity of the wing bones to the rest of the skeleton. All the bones attributed to Rahonavis were buried in an area "smaller than a letter-sized page", according to co-describer Luis M. Chiappe in his 2007 book Glorified Dinosaurs. Additionally, Chiappe argued that suggestions of a chimera by paleornithologist Larry Martin were based on Martin's misinterpretation of the wing and shoulder bones as being more advanced than they really are. [4]

A speculative restoration of two Rahonavis (hypothetically-depicted male & female) on a charred branch. Birds of Old (Rahonavis).jpg
A speculative restoration of two Rahonavis (hypothetically-depicted male & female) on a charred branch.

Chiappe maintained that Rahonavis could probably fly, noting that its ulna was large and robust compared to Archaeopteryx, and that this fact, coupled with the prominent quill knobs, suggest that Rahonavis had larger and more powerful wings than that earlier bird. Additionally, Rahonavis shoulder bones show evidence of ligament attachments allowing the independent mobility needed for flapping flight. Chiappe concluded that Rahonavis was capable of flight, though it would have been more "clumsy in the air than modern birds." [4] Agnolín and Novas (2013) noted that, like Microraptor , a bat-like flightstroke using the deltoideus complexes seems to have been likely in R. ostromi. [13]

See also

Related Research Articles

<span class="mw-page-title-main">Dromaeosauridae</span> Family of theropod dinosaurs

Dromaeosauridae is a family of feathered coelurosaurian theropod dinosaurs. They were generally small to medium-sized feathered carnivores that flourished in the Cretaceous Period. The name Dromaeosauridae means 'running lizards', from Greek δρομαῖος (dromaîos), meaning 'running at full speed', 'swift', and σαῦρος (saûros), meaning 'lizard'. In informal usage, they are often called raptors, a term popularized by the film Jurassic Park; several genera include the term "raptor" directly in their name, and popular culture has come to emphasize their bird-like appearance and speculated bird-like behavior.

Vorona is a monotypic genus of prehistoric birds. It was described from fossils found in a Maevarano Formation quarry near the village of Berivotra, Mahajanga Province, Madagascar. The age of the fossilised specimen is Late Cretaceous, probably Maastrichtian. V. berivotrensis is known from scattered remains, possibly from a single individual.

<span class="mw-page-title-main">Maniraptora</span> Clade of dinosaurs

Maniraptora is a clade of coelurosaurian dinosaurs which includes the birds and the non-avian dinosaurs that were more closely related to them than to Ornithomimus velox. It contains the major subgroups Avialae, Dromaeosauridae, Troodontidae, Oviraptorosauria, and Therizinosauria. Ornitholestes and the Alvarezsauroidea are also often included. Together with the next closest sister group, the Ornithomimosauria, Maniraptora comprises the more inclusive clade Maniraptoriformes. Maniraptorans first appear in the fossil record during the Jurassic Period, and survive today as living birds.

<i>Buitreraptor</i> Dromaeosaurid dinosaur genus from the Late Cretaceous

Buitreraptor is a genus of dromaeosaurid dinosaurs that lived during the Late Cretaceous of Argentina at the Candeleros Formation. Buitreraptor was described in 2005 and the type species is Buitreraptor gonzalezorum. It was rooster-sized and had a very elongated head with many small teeth.

<i>Jinfengopteryx</i> Theropod dinosaur genus

Jinfengopteryx is a genus of maniraptoran dinosaur. It was found in the Qiaotou Member of the Huajiying Formation of Hebei Province, China, and is therefore of uncertain age. The Qiaotou Member may correlate with the more well-known Early Cretaceous Yixian Formation, and so probably dates to around 122 Ma ago.

<i>Pyroraptor</i> Extinct genus of dinosaurs

Pyroraptor is an extinct genus of paravian dinosaur, probably a dromaeosaurid or unenlagiid, from the Late Cretaceous Ibero-Armorican island, of what is now southern France. It lived during the late Campanian and early Maastrichtian stages, approximately 72 million years ago. It is known from a single partial specimen that was found in Provence in 1992, after a forest fire. The animal was named Pyroraptor olympius by Allain and Taquet in 2000.

Unenlagia is a genus of dromaeosaurid theropod dinosaur that lived in South America during the Late Cretaceous period. The genus Unenlagia has been assigned two species: U. comahuensis, the type species described by Novas and Puerta in 1997, and U. paynemili, described by Calvo et al. in 2004.

<span class="mw-page-title-main">Scansoriopterygidae</span> Extinct family of dinosaurs

Scansoriopterygidae is an extinct family of climbing and gliding maniraptoran dinosaurs. Scansoriopterygids are known from five well-preserved fossils, representing four species, unearthed in the Tiaojishan Formation fossil beds of Liaoning and Hebei, China.

<span class="mw-page-title-main">Archaeopterygidae</span> Family of dinosaurs

Archaeopterygidae is a group of paravian dinosaurs, known from the latest Jurassic and earliest Cretaceous of Europe. In most current classifications, it contains only the genera Archaeopteryx and Wellnhoferia. As its name suggests, Protarchaeopteryx was also once referred to this group, but most paleontologists now consider it an oviraptorosaur. Other referred genera, like Jurapteryx, Wellnhoferia, and "Proornis", are probably synonymous with Archaeopteryx or do not belong into this group. Jinfengopteryx was originally described as an archaeopterygid, though it was later shown to be a troodontid. A few studies have recovered Anchiornis and Xiaotingia to also be members of the Archaeopterygidae, though most subsequent analyses have failed to arrive at the same result. Uncertainties still exist, however, and it may not be possible to confidently state whether archaeopterygids are more closely related to modern birds or to deinonychosaurs barring new and better specimens of relevant species. Teeth attributable to archaeopterygids are known from the earliest Cretaceous (Berriasian) Cherves-de-Cognac locality and the Angeac-Charente bonebed of France.

<span class="mw-page-title-main">Unenlagiinae</span> Extinct subfamily of dinosaurs

Unenlagiinae is a subfamily of long-snouted paravian theropods. They are traditionally considered to be members of Dromaeosauridae, though some authors place them into their own family, Unenlagiidae, sometimes alongside the subfamily Halszkaraptorinae. Definitive members are known from South America, though some researchers include taxa from other continents within this subfamily based on phylogenetic analyses.

<i>Shanag</i> Extinct genus of dinosaurs

Shanag is a genus of paravian theropod dinosaur from the Early Cretaceous Period of Mongolia. It may be a dromaeosaurid, but some researchers are skeptical of this classification. The type species is S. ashile.

<span class="mw-page-title-main">Avialae</span> Clade including all birds and their ancestors

Avialae is a clade containing the only living dinosaurs, the birds, and their closest relatives. It is usually defined as all theropod dinosaurs more closely related to birds (Aves) than to deinonychosaurs, though alternative definitions are occasionally used.

<span class="mw-page-title-main">Paraves</span> Clade of all dinosaurs that are more closely related to birds than to oviraptorosaurs

Paraves are a widespread group of theropod dinosaurs that originated in the Middle Jurassic period. In addition to the extinct dromaeosaurids, troodontids, anchiornithids, and possibly the scansoriopterygids, the group also contains the avialans, which include diverse extinct taxa as well as the over 10,000 species of living birds.

<span class="mw-page-title-main">Mahakala omnogovae</span> Extinct species of dinosaur

Mahakala is a genus of halszkaraptorine theropod dinosaur from the Campanian-age Upper Cretaceous Djadokhta Formation of Ömnögovi, Mongolia. It is based on a partial skeleton found in the Gobi Desert. Mahakala was a small dromaeosaurid, and its skeleton shows features that are also found in early troodontids and avialans. Despite its late appearance, it is among the most basal dromaeosaurids. Its small size, and the small size of other basal deinonychosaurians, suggests that small size appeared before flight capability in birds. The genus is named for Mahakala, one of eight protector deities (dharmapalas) in Tibetan Buddhism.

The proximodorsal process is a feature of the skeleton of archosaurs. It may be a pair of tabs or blade - shaped flanges on the pelvis, and serves as an anchor point for the attachment of leg muscles. This process is of particular importance in the anatomy and comparative morphology of Mesozoic birds and advanced maniraptoran dinosaurs. The pelvis is made up of three paired bones and a sacrum. The three paired bones are called the ilium, the ischium, and the pubis. On the ischium there may be an obturator process and/or a proximodorsal process. The more primitive condition is for there to be no proximodorsal process, but a large obturator process. In primitive birds the ischia are complex, usually with a small or even absent obturator process and a large, rectangular, proximodorsal process extending up toward the ilium. This is the condition in Archaeopteryx, Confuciusornis, and enantiornithines. The South American dromaeosaurs called the unenlagiinae have an intermediate condition between the two, with both a large obturator process and a proximodorsal process.

<i>Tianyuraptor</i> Extinct genus of dinosaurs

Tianyuraptor is a genus of short-armed dromaeosaurid dinosaur that lived during the Early Cretaceous, about 122 million years ago. Its remains have been found in western Liaoning, China. It was similar to other dromaeosaurids found in Liaoning, with the exception of being somewhat more primitive. The type specimen, formally named in 2009, shows features not seen in previously known Northern Hemisphere (Laurasian) dromaeosaurids, but present in Southern Hemisphere (Gondwanan) species and early birds. Because of this, the scientists who first studied Tianyuraptor described it as a "transitional species", bridging the gap between northern and southern types of dromaeosaurid. Tianyuraptor also differs from previously known dromaeosaurids in that it possesses a relatively small furcula ("wishbone"), and unusually short forelimbs.

<span class="mw-page-title-main">Jinfengopteryginae</span> Extinct subfamily of dinosaurs

Jinfengopteryginae is a subfamily of bird-like theropod dinosaurs known from the Cretaceous of Eurasia. This group includes relatively few genera, with members discovered in 2005 but the name erected in 2012. Like other troodontids, this group of dinosaurs resided in the Paraves potentially close to the Avialae.

<span class="mw-page-title-main">Anchiornithidae</span> Extinct family of dinosaurs

Anchiornithidae is a family of small paravian dinosaurs. Anchiornithids have been classified at varying positions in the paravian tree, with some scientists classifying them as a distinct family, a basal subfamily of Troodontidae, members of Archaeopterygidae, or an assemblage of dinosaurs that are an evolutionary grade within Avialae or Paraves.

<i>Overoraptor</i> Extinct genus of theropod dinosaurs

Overoraptor is an extinct genus of paravian theropod of uncertain affinities from the Late Cretaceous Huincul Formation of Argentinian Patagonia. The genus contains a single species, O. chimentoi, known from several bones of the hands, feet, and hips alongside some vertebrae.

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Further reading