Rahonavis

Rahonavis is a genus of bird-like theropods from the Late Cretaceous (Maastrichtian, about 70 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,[2] with the typical Velociraptor-like raised sickle claw on the second toe.

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.

Rahonavis
Temporal range: Late Cretaceous, 70 Ma
Maniraptoran ROM
Restored skeleton
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Order: Saurischia
Suborder: Theropoda
Clade: Paraves
Genus: Rahonavis
Forster et al., 1998b
Type species
Rahona ostromi
(Forster et al., 1998a) Forster et al., 1998b
Synonyms

Classification

Rahonavis
Skeletal restoration

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 primitive 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.[3]

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.[4] Norell and colleagues (2006) also found Rahonavis to lie within the Unenlagiinae, as the sister taxon to Unenlagia itself.[5] A 2007 study by Turner and colleagues again found it to be an unenlagiine dromaeosaurid, closely related to Unenlagia.[6]

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.[7] A cladistic analysis by Cau (2018) recovered Rahonavis as a probable relative of the long-tailed Early Cretaceous avialans Jeholornis and Jixiangornis.[8]

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.[9][10]

Discovery and species

The fossilized remains of Rahonavis were first recovered 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.[11]

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 mid-Cretaceous sediments (Albian/Cenomanian, c. 100 to 99 million years ago) of the Kem Kem region, 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.[12]

Paleobiology

Rahonavis NT
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.[3] 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.[11]

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."[11] Agnolín and Novas (2013) noted that, like Microraptor, a bat-like flightstroke using the deltoideus complexes seems to have been likely in R. ostromi.[7]

See also

References

  1. ^ Tudge, Colin (2009) The Bird:A Natural History of Who Birds Are, Where They Came From, and How They Live [1]
  2. ^ Holtz, Thomas R. Jr. (2008) Dinosaurs: The Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages Supplementary Information
  3. ^ a b Geist, Nicholas R.; Feduccia, Alan (2000). "Gravity-defying Behaviors: Identifying Models for Protoaves" (PDF). American Zoologist. 40 (4): 664–675. doi:10.1668/0003-1569(2000)040[0664:GDBIMF]2.0.CO;2.
  4. ^ Makovicky, Peter J.; Apesteguía, Sebastián; Agnolín, Federico L. (2005). "The earliest dromaeosaurid theropod from South America". Nature. 437 (7061): 1007–1011. Bibcode:2005Natur.437.1007M. doi:10.1038/nature03996. PMID 16222297. Supplementary information.
  5. ^ Norell, M.A.; Clark, J. M.; Turner, A.H.; Makovicky, P. J.; Barsbold, R.; Rowe, T. (2006). "A new dromaeosaurid theropod from Ukhaa Tolgod (Omnogov, Mongolia)". American Museum Novitates. 3545 (3545): 1–51. doi:10.1206/0003-0082(2006)3545[1:ANDTFU]2.0.CO;2.
  6. ^ Turner, Alan H.; Pol, Diego; Clarke, Julia A.; Erickson, Gregory M.; Norell, Mark (2007). "A basal dromaeosaurid and size evolution preceding avian flight" (PDF). Science. 317 (5843): 1378–1381. Bibcode:2007Sci...317.1378T. doi:10.1126/science.1144066. PMID 17823350.
  7. ^ a b Agnolín, F.L.; Novas, F.E. (2013). "Avian ancestors. A review of the phylogenetic relationships of the theropods Unenlagiidae, Microraptoria, Anchiornis and Scansoriopterygidae". SpringerBriefs in Earth System Sciences. pp. 1–96.
  8. ^ Andrea Cau (2018). "The assembly of the avian body plan: a 160-million-year long process" (PDF). Bollettino della Società Paleontologica Italiana. 57 (1): 1–25. doi:10.4435/BSPI.2018.01.
  9. ^ Forster, Catherine A.; Sampson, Scott D.; Chiappe, Luis M.; Krause, David W. (1998). "The Theropod ancestry of birds: New evidence from the late Cretaceous of Madagascar". Science. 279 (5358): 1915–1919. Bibcode:1998Sci...279.1915F. doi:10.1126/science.279.5358.1915. PMID 9506938.
  10. ^ Forster, Catherine A.; Sampson, Scott D.; Chiappe, Luis M.; Krause, David W. (1998). "Genus correction". Science. 280 (5361): 179. Bibcode:1998Sci...280..179F. doi:10.1126/science.280.5361.179g.
  11. ^ a b c Chiappe, L. M. (2007-02-02). Glorified Dinosaurs: The Origin and Early Evolution of Birds. Sydney: UNSW Press. ISBN 978-0-471-24723-4.
  12. ^ Riff, Douglas; Kellner, Alexander W. A.; Mader, Bryn; Russell, Dale (2002). "On the occurrence of an avian vertebra in Cretaceous strata of Morocco, Africa" (PDF). Anais da Academia Brasileira de Ciências. 2 (74): 367–368. doi:10.1590/S0001-37652002000200023.

Further reading

  • Forster, Catherine A.; O'Conner (2000). "The avifauna of the Upper Cretaceous Maevarano Formation, Madagascar". Journal of Vertebrate Paleontology. 3 (20): 41A–42A.
  • Schweitzer, Mary H.; Watt, John A.; Avci, Recep; Forster, Catherine A.; Krause, David W.; Knapp, Loren; Rogers, Raymond R.; Beech, Iwona; Marshall, Mark (1999). "Keratin immunoreactivity in the Late Cretaceous bird Rahonavis ostromi". Journal of Vertebrate Paleontology. 19 (4): 712–722. doi:10.1080/02724634.1999.10011183. JSTOR 4524040.
Acheroraptor

Acheroraptor is an extinct genus of dromaeosaurid theropod dinosaur known from the latest Maastrichtian Hell Creek Formation of Montana, United States. It contains a single species, Acheroraptor temertyorum. A. temertyorum is one of the two geologically youngest known species of dromaeosaurids, the other being Dakotaraptor, which is also known from Hell Creek.

Austroraptor

Austroraptor ( AW-stroh-RAP-tər) is an extinct genus of dromaeosaurid dinosaur that lived about 70 million years ago during the Cretaceous Period in what is now modern Argentina. Austroraptor was a medium sized, moderately-built, ground-dwelling, bipedal carnivore, that could grow up to 5–6 m (16.4–19.7 ft) long. Its length makes Austroraptor one of the largest dromaeosaurids known, with only Achillobator, Dakotaraptor, and Utahraptor approaching or surpassing it in length. It is the largest dromaeosaur to be discovered in the Southern Hemisphere. Particularly notable about the taxon were its relatively short forearms, much shorter in proportion when compared to the majority of the members of its group.

Avialae

Avialae ("bird wings") is a clade of flying dinosaurs containing the only living dinosaurs, the birds. It is usually defined as all theropod dinosaurs more closely related to modern birds (Aves) than to deinonychosaurs, though alternative definitions are occasionally used (see below).

Archaeopteryx lithographica, from the late Jurassic Period Solnhofen Formation of Germany, is the earliest known avialan which may have had the capability of powered flight. However, several older avialans are known from the late Jurassic Tiaojishan Formation of China, dated to about 160 million years ago.

Buitreraptor

Buitreraptor is a predatory dromaeosaurid theropod dinosaur from the Cretaceous of Argentina.

Buitreraptor was described in 2005. The type species is Buitreraptor gonzalezorum. It was rooster-sized and had a very elongated head with many small teeth.

Dromaeosauridae

Dromaeosauridae is a family of feathered 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 δρομεῦς (dromeus) meaning 'runner' and σαῦρος (sauros) meaning 'lizard'. In informal usage they are often called raptors (after Velociraptor), a term popularized by the film Jurassic Park; a few types include the term "raptor" directly in their name and have come to emphasize their bird-like appearance and speculated bird-like behavior.

Dromaeosaurid fossils have been found across the globe in North America, Europe, Africa, Asia, South America and Antarctica, with fossilized teeth giving credence to the possibility that they inhabited Australia as well. They first appeared in the mid-Jurassic Period (late Bathonian stage, about 167 million years ago) and survived until the end of the Cretaceous (Maastrichtian stage, 66 ma), existing until the Cretaceous–Paleogene extinction event. The presence of dromaeosaurids as early as the Middle Jurassic has been suggested by the discovery of isolated fossil teeth, though no dromaeosaurid body fossils have been found from this period.

Hyposphene-hypantrum articulation

The hyposphene-hypantrum articulation is an accessory joint found in the vertebrae of several fossil reptiles of the group Archosauromorpha. It consists of a process on the backside of the vertebrae, the hyposphene, that fits in a depression in the front side of the next vertebrae, the hypantrum. Hyposphene-hypantrum articulations occur in the dorsal vertebrae and sometimes also in the posteriormost cervical and anteriormost caudal vertebrae.In most tetrapods including the human, the vertebrae are connected with each other only via the centrum and the zygapophysis joints. Additional joints like the hyposphene-hypantrum articulations, which add rigidity to the vertebral column, are found in several different reptile lineages; a known example are the zygosphene-zygantrum articulations found in snakes.Hyposphene-hypantrum articulations are found in several unrelated groups within the Archosauromorpha. They occur especially in large forms, for example in rauisuchids and in silesaurids and – within the Dinosauria – in saurischians. They evolved to make the vertebral column more rigid and stable and probably had supported the gigantism in sauropod dinosaurs.Early Dinosauromorphs (early ancestors of dinosaurs) like Marasuchus, Lagosuchus and Euparkeria as well as ornithischian dinosaurs lack hyposphene-hypantrum articulations. Because these articulations are absent in both saurischian ancestors and all non-saurischian dinosaurs, they are considered a synapomorphy (a distinctive feature) of the Saurischia, as proposed by Gauthier (1986). Hyposphene-hypantrum articulations are found in all the basal members of the Saurischia. However, they became lost in several saurischian lineages. They were present in the derived and birdlike dromaeosaurid Rahonavis, but are lost in modern day's birds, probably due to their highly modified vertebrae. Within the Sauropodomorpha, they were present in prosauropods and most sauropods, but became independently lost in two cretaceous sauropod lineages, the Titanosauria and the Rebbachisauridae.The hyposphene usually consists of a vertical ridge and is situated below the postzygapophysis, whereas the hypantrum is situated between the prezygapophysis. In sauropods, this joint is extremely variable.

Itemirus

Itemirus is a genus of theropod dinosaur from the Turonian age of the Late Cretaceous period of Uzbekistan.

List of Indian and Madagascan dinosaurs

This is a list of dinosaurs whose remains have been recovered from India or Madagascar. Though widely separated today, India and Madagascar were connected throughout much of the Mesozoic and shared similar dinosaur faunas, distinct from what has been found on other modern African and Asian landmasses.

The Indian fossil record of dinosaurs is good, with fossils coming from the entire Mesozoic era – starting with the Triassic period (a geological period that started 251.9 million years ago and continued till 201.3 million years ago), to the Jurassic period (201 million years ago to 145 million years ago) and Cretaceous period (from 145 million years ago to 66 million years ago), when globally all non-avian dinosaurs and 65 per cent of all life became extinct. Madagascar also preserves various unique dinosaurs from the Jurassic and Cretaceous.

Luanchuanraptor

Luanchuanraptor (meaning "Luanchuan thief") is a genus of dromaeosaurid theropod dinosaur from the Late Cretaceous of China. It is based on a partial skeleton from the Qiupa Formation in Luanchuan, Henan. A medium-sized dromaeosaurid, it is the first Asian dromaeosaurid described from outside the Gobi Desert or northeastern China. The fossil material is cataloged as 4HIII-0100 in the Henan Geological Museum and includes four teeth, one frontal, a neck vertebra, one or two back vertebrae, seventeen tail vertebrae, ribs, chevrons, a humerus (upper arm bone), claw and finger bones, partial shoulder and pelvic girdles, and other fragmentary bones from a moderately sized dromaeosaurid. The type species is L. henanensis, described by Lü et al. in 2007.

Maevarano Formation

The Maevarano Formation is an Upper Cretaceous sedimentary rock formation found in the Mahajanga Province of northwestern Madagascar. It is most likely Maastrichtian in age, and records a seasonal, semiarid environment with rivers that had greatly varying discharges. Notable animal fossils recovered include the theropod dinosaur Majungasaurus, the early bird Vorona, the flying dromaeosaur Rahonavis, the titanosaurian sauropod Rapetosaurus, and the giant frog Beelzebufo.

Masiakasaurus

Masiakasaurus is a genus of small predatory theropod dinosaurs from the Late Cretaceous of Madagascar. In Malagasy, masiaka means "vicious"; thus, the genus name means "vicious lizard". The type species, Masiakasaurus knopfleri, was named after the musician Mark Knopfler, whose music inspired the expedition crew. It was named in 2001 by Scott D. Sampson, Matthew Carrano, and Catherine A. Forster. Unlike most theropods, the front teeth of M. knopfleri projected forward instead of straight down. This unique dentition suggests that they had a specialized diet, perhaps including fish and other small prey. Other bones of the skeleton indicate that Masiakasaurus were bipedal, with much shorter forelimbs than hindlimbs. M. knopfleri reached an estimated adult body length of around 2 metres (6 ft 7 in).Masiakasaurus lived around 70 million years ago, along with animals such as Majungasaurus, Rapetosaurus, and Rahonavis. Masiakasaurus was a member of the group Noasauridae, small predatory ceratosaurs found primarily in South America.

Rahona

Rahona is a genus of moths in the subfamily Lymantriinae. It was named by Paul Griveaud in 1975.Most of the species of this genus occur in central Africa or Madagascar.

The name was inadvertently used again in 1998 for a fossil species of Avian theropod Rahona ostromi, by Catherine Forster and colleagues. When they discovered that the name had already been used by Griveaud, they renamed the fossil Rahonavis.

Saurornitholestinae

Saurornitholestinae is a subfamily of dromaeosaurid dinosaurs. The saurornitholestines currently include three monotypic genera: Atrociraptor marshalli, Bambiraptor feinbergorum, and Saurornitholestes langstoni. All are medium-sized dromaeosaurs from the Late Cretaceous of western North America. The group was originally recognized by Longrich and Currie as the sister taxon to a clade formed by the Dromaeosaurinae and Velociraptorinae. However, not all phylogenetic analyses recover this group.

Tianyuraptor

Tianyuraptor is a genus of short-armed dromaeosaurid dinosaur ('running lizard'; a type of small dinosaur considered to be closely related to birds) 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.

Unenlagia

Unenlagia (meaning "half-bird" in latinized mapudungun) is a genus of theropod dinosaur from the Late Cretaceous of Argentina.

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.

Unenlagiinae

Unenlagiinae is a subfamily of dromaeosaurid theropods. Unenlagiines are known from South America and Antarctica.

Unquillosaurus

Unquillosaurus (meaning "Unquillo river lizard") is a genus of maniraptoran dinosaur from the Late Cretaceous Period, discovered in Argentina. Known only from a single fossilized pubis (a pelvic bone), its total body length may have reached 2 to 3 metres (6.6 to 9.8 ft).

Vorona

For the commune in Botoşani County, Romania, see Vorona, Botoșani.

Vorona ( VOOR-oo-nə; Malagasy for "bird", V. berivotrensis, "from Berivotra") 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 is Late Cretaceous, probably Campanian (70.6–83.5 mya). V. berivotrensis is known from scattered remains, possibly from a single individual (UA 8651 and FMNH PA715).

The phylogenic affiliation of Vorona is hard to determine due to the fragmentary nature of the remains, mainly because the fossil shows a mix of primitive avian features as well as some that seem very modern. Vorona might be a primitive ornithuromorph.

Vorona is sometimes confused with the dromaeosaur Rahonavis ostromi, a fossil of which was found in the same location. This confusion has led to the common misconception that Vorona had a deinonychosaur-like sickle claw on each foot.

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