Rebbachisauridae

Rebbachisauridae is a family of sauropod dinosaurs known from fragmentary fossil remains from the Cretaceous of South America, Africa, North America, and Europe.

Rebbachisauridae
Temporal range: Late Jurassic to Late Cretaceous, 150–93 Ma
Limaysaurus
Limaysaurus tessonei skeleton restoration
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Order: Saurischia
Suborder: Sauropodomorpha
Clade: Sauropoda
Clade: Diplodocimorpha
Family: Rebbachisauridae
Bonaparte, 1997
Genera

Taxonomy

In 1990 sauropod specialist Jack McIntosh included the first known rebbachisaurid genus, the giant North African sauropod Rebbachisaurus, in the family Diplodocidae, subfamily Dicraeosaurinae, on the basis of skeletal details. With the discovery in subsequent years of a number of additional genera, it was realised that Rebbachisaurus and its relatives constituted a distinct group of dinosaurs. In 1997 the Argentine paleontologist José Bonaparte described the family Rebbachisauridae, and in 2011 Whitlock defined two new subfamilies within the group: Nigersaurinae and Limaysaurinae. Cladogram of the Rebbachisauridae after Fanti et al. (2013) which is based on Carballido et al. (2012):[2]

Rebbachisauridae

Amazonsaurus

Histriasaurus

Zapalasaurus

Comahuesaurus

Limaysaurinae

Rayososaurus

Rebbachisaurus

Cathartesaura

Limaysaurus

Nigersaurinae

Nigersaurus

Demandasaurus

Tataouinea

Cladogram after Fanti et al., 2015.[3]

Rebbachisauridae

Amazonsaurus

Zapalasaurus

Histriasaurus

Comahuesaurus

Khebbashia
Limaysaurinae

Cathartesaura

Limaysaurus

Rebbachisaurinae

Katepensaurus

Nigersaurus

Rebbachisaurus

Demandasaurus

Tataouinea

Evolutionary relationships and characteristics

Nigersaurus taqueti teeth
Nigersaurus taqueti teeth

Although all authorities agree that the rebbachisaurids are members of the superfamily Diplodocoidea, they lack the bifid (divided) cervical neural spines that characterise the diplodocids and dicraeosaurids, and for this reason are considered more primitive than the latter two groups. It is not yet known whether they share the distinctive whip-tail of the latter two taxa.

Rebbachisaurids are distinguished from other sauropods by their distinctive teeth, which have low angle, internal wear facets and asymmetrical enamel.

Unique among sauropods, at least some rebbachisaurids (such as Nigersaurus) are characterised by the presence of tooth batteries, similar to those of hadrosaur and ceratopsian dinosaurs. Such a feeding adaptation has thus developed independently three times among the dinosaurs.

So far, rebbachisaurids are known only from the middle and early part of the Late Cretaceous. Unless the nemegtosaurids are in fact diplodocoids (rather than titanosaurs), then the rebbachisaurids represent the last known representatives of this clade, and lived alongside the titanosaurs until fairly late in the Cretaceous. So far, no rebbachisaurids are known from the very end of the Cretaceous period.

References

  • Bonaparte J.F. (1997). "Rayososaurus grioensis Bonaparte 1995". Ameghiniana. 34 (1): 116.
  • McIntosh, J. S., 1990, "Sauropoda" in The Dinosauria, Edited by David B. Weishampel, Peter Dodson, and Halszka Osmólska. University of California Press, pp. 345–401.
  • Upchurch, P., Barrett, P.M. and Dodson, P. 2004. "Sauropoda". In The Dinosauria, 2nd edition. Weishampel, Dodson, and Osmólska (eds.). University of California Press, Berkeley. pp. 259–322.
  • Wilson J.A. (2002). "Sauropod dinosaur phylogeny: critique and cladistic analysis" (PDF). Zoological Journal of the Linnean Society. 136 (2): 215–275. doi:10.1046/j.1096-3642.2002.00029.x.
  • ------ (2005) "Overview of Sauropod Phylogeny and Evolution", in The Sauropods: Evolution and Paleobiology
  • Wilson, J. A. and Sereno, P.C. (2005) "Structure and Evolution of a Sauropod Tooth Battery" in The Sauropods: Evolution and Paleobiology in Curry Rogers and Wilson, eds, 2005, The Sauropods: Evolution and Paleobiology, University of California Press, Berkeley, ISBN 0-520-24623-3
  1. ^ Paul C. Sereno, Jeffrey A. Wilson, Lawrence M. Witmer, John A. Whitlock, Abdoulaye Maga, Oumarou Ide, Timothy A. Rowe (2007). Kemp, Tom (ed.). "Structural Extremes in a Cretaceous Dinosaur". PLoS ONE. 2 (11): e1230. doi:10.1371/journal.pone.0001230. PMC 2077925. PMID 18030355.CS1 maint: Multiple names: authors list (link)
  2. ^ Carballido, José Luis; Salgado, Leonardo; Pol, Diego; Canudo, José Ignacio; Garrido, Alberto (2012). "A new basal rebbachisaurid (Sauropoda, Diplodocoidea) from the Early Cretaceous of the Neuquén Basin; evolution and biogeography of the group". Historical Biology. 24 (6): 631–654. doi:10.1080/08912963.2012.672416.
  3. ^ Fanti, F.; Cau, A.; Cantelli, L.; Hassine, M.; Auditore, M. (2015). "New Information on Tataouinea hannibalis from the Early Cretaceous of Tunisia and Implications for the Tempo and Mode of Rebbachisaurid Sauropod Evolution". PLOS ONE. 10 (4): e123475. doi:10.1371/journal.pone.0123475. PMC 4414570. PMID 25923211.
Amphicoelias

Amphicoelias (, meaning "biconcave", from the Greek ἀμφί, amphi: "on both sides", and κοῖλος, koilos: "hollow, concave") is a genus of herbivorous sauropod dinosaur that lived approximately 150 million years ago during the Tithonian (Late Jurassic Period) of what is now Colorado, United States. A herbivore, Amphicoelias was moderately sized at about 25 m (82 ft) long–roughly the same length as Diplodocus, to which it was related. Its hindlimbs were very long and thin, and its forelimbs were proportionally longer than in relatives.

The namesake fossil of the type species Amphicoelias altus, American Museum of Natural History 5764, is uncertain in included material. When described by Edward Drinker Cope shortly after its discovery in 1877, Amphicoelias was noted to include many back vertebrae, a single pubis, and a femur. However, after purchase and cataloging of the material by the AMNH, Henry Fairfield Osborn and Charles Mook described that the specimen had only two vertebrae, a pubis, femur, tooth, scapula, coracoid, ulna and a second femur. The additional material, not mentioned by Cope, was found close in proximity to the holotype and was similar to Diplodocus, a relative of Amphicoelias. Their assignment was questioned by subsequent authors Emanuel Tschopp et al. in an analysis of Diplodocidae.

During the description of Amphicoelias altus in 1877, Cope additionally named A. latus, for a femur and tail vertebrae. Following its description, Osborn and Mook in 1921 reidentified the material as a specimen of Camarasaurus, an assignment followed by other authors who reviewed the material. A year later 1878, Cope named the third species of Amphicoelias, A. fragillimus for a gigantic dorsal vertebra that was subsequently lost. Measuring approximately 2.7 m (8.9 ft) if reconstructed based on Diplodocus, early estimates for the length of the animal in life were between 40 and 60 m (130 and 200 ft) long. Due to the incomplete nature, such lengths–the longest of any known dinosaur and sauropod–were largely ignored. In 2018, Kenneth Carpenter renamed Amphicoelias fragillimus as the new genus Maraapunisaurus, and reclassified it from Diplodocidae to Rebbachisauridae.

Cathartesaura

Cathartesaura is a genus of rebbachisaurid sauropod dinosaur hailing from the Late Cretaceous strata of the Huincul Formation, at the "La Buitrera" locality, in the Neuquén Basin of Río Negro Province, Argentina. The fossil remains, described by Gallina and Apesteguía in 2005, consist of a partial skeleton including vertebrae and limb bones. These were found at the base of the formation, which spans the Cenomanian and Coniacian epochs, in mudstone and sandstone levels.

Comahuesaurus

Comahuesaurus is a genus of sauropod dinosaur of the family Rebbachisauridae. It was found in the Lohan Cura Formation, in Argentina and lived during the Early Cretaceous, Aptian to Albian. The type species is C. windhauseni, named by Carballido and colleagues in 2012. It had originally been assigned to Limaysaurus by Salgado et al. (2004), but was later assigned its own genus based on the presence of diagnostic characters in the caudal centra, pubis and ischium.Comahuesaurus is known from abundant material compared to other rebbachisaurids; 37 caudal vertebrae, three fragmentary dorsal vertebrae and multiple appendicular elements, including a right humerus, pubis, ischium and a 113 cm long left femur. In their phylogenetic analysis, Carballido et al. (2012) placed Comahuesaurus in an intermediate position between basal rebbachisaurids such as Histriasaurus and the derived clade formed by subfamilies Rebbachisaurinae and Limaysaurinae.It shares with more derived rebbachisaurids a reduced hyposphene-hypantrum system, but hadn't yet completely lost said structure; that change would happen at some further point in the evolution of the clade, as it is so far only known to be fully absent in limaysaurines.

Demandasaurus

Demandasaurus (meaning "Demanda lizard") is a genus of rebbachisaurid sauropod dinosaur from early Cretaceous (late Barremian – early Aptian stage) deposits of Spain. Demandasaurus is known from an incomplete but associated skeleton that includes cranial and postcranial remains. It was collected from the Castrillo la Reina Formation in Burgos Province of Spain. It was first named by Fidel Torcida Fernández-Baldor, José Ignacio Canudo, Pedro Huerta, Diego Montero, Xabier Pereda Suberbiola and Leonardo Salgado in 2011 and the type species is Demandasaurus darwini.

Dicraeosaurus

Dicraeosaurus (Gr. δικραιος, dikraios "bifurcated, double-headed" + Gr. σαυρος, sauros "lizard") is a genus of small diplodocoid sauropod dinosaur that lived in what is now Tanzania during the late Jurassic. It was named for the spines on the back of the neck. The first fossil was described by paleontologist Werner Janensch in 1914.

Diplodocimorpha

Diplodocimorpha is a clade of extinct sauropod dinosaurs, existing from the Early Jurassic until the Late Cretaceous. The group includes three main families and some other genera, Rebbachisauridae, Dicraeosauridae and Diplodocidae, the latter two forming Flagellicaudata. The name was first used by Calvo & Salgado (1995), who defined it as "Rebbachisaurus tessonei sp. nov., Diplodocidae, and all descendants of their common ancestor." The group was not used often, and was synonymized with Diplodocoidea as the groups were often found to have the same content. In 2005, Mike P. Taylor and Darren Naish reviewed diplodocoid phylogeny and taxonomy, and realized that Diplodocimorpha could not be synonymized with Diplodocoidea. Whereas the former was defined node-based, the latter was branch-based. In 2015, Emanuel Tschopp, Octavio Mateus and Roger Benson published a specimen-based phylogeny on diplodocid interrelationships, and supported the separation of Diplodocimorpha. Haplocanthosaurus was found to be more basal than rebbachisaurids, and therefore outside Diplodocimorpha, but closer to Diplodocus than Saltasaurus, and therefore within Diplodocoidea. The below cladogram follows the findings of Tschopp et al.

Diplodocoidea

Diplodocoidea is a superfamily of sauropod dinosaurs, which included some of the longest animals of all time, including slender giants like Supersaurus, Diplodocus, Apatosaurus, and Amphicoelias. Most had very long necks and long, whip-like tails; however, one family (the dicraeosaurids) are the only known sauropods to have re-evolved a short neck, presumably an adaptation for feeding low to the ground. This adaptation was taken to the extreme in the highly specialized sauropod Brachytrachelopan. A study of snout shape and dental microwear in diplodocoids showed that the square snouts, large proportion of pits, and fine subparallel scratches in Apatosaurus, Diplodocus, Nigersaurus, and Rebbachisaurus suggest ground-height nonselective browsing; the narrow snouts of Dicraeosaurus, Suuwassea, and Tornieria and the coarse scratches and gouges on the teeth of Dicraeosaurus suggest mid-height selective browsing in those taxa. This taxon is also noteworthy because diplodocoid sauropods had the highest tooth replacement rates of any vertebrates, as exemplified by Nigersaurus, which had new teeth erupting every 30 days.

Flagellicaudata

Flagellicaudata is a clade of Dinosauria. It belongs to Sauropoda and includes two families, the Dicraeosauridae and the Diplodocidae.

Histriasaurus

Histriasaurus (HIS-tree-ah-SAWR-us) (meaning "Istria lizard") was a genus of dinosaur from the Early Cretaceous (Hauterivian to Barremian stages, around 130 million years ago). Its fossils, holotype WN V-6, were found near the town of Bale on the Istrian peninsula in Croatia, and described in 1998 by Dalla Vecchia. It was a diplodocoid sauropod, related to, but more primitive than, Rebbachisaurus. Phylogenetic analyses published in 2007 and 2011 placed Histriasaurus as the most basal member of Rebbachisauridae.The type species, H. boscarollii, was described by Dalla Vecchia in 1998. The specific name honours the discoverer of the site, Darío Boscarolli. Although some authors consider Histriasaurus a dubious taxon, more recent papers support the original classification.

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.

Katepensaurus

Katepensaurus is an extinct genus of rebbachisaurid sauropod dinosaur known from the Late Cretaceous of south-central Chubut Province of central Patagonia, Argentina. It contains a single species, Katepensaurus goicoecheai.

Lavocatisaurus

Lavocatisaurus is a genus of sauropod in the family Rebbachisauridae from the Early Cretaceous (Aptian to Albian) Rayoso Formation of the Neuquén Basin, northern Patagonia, Argentina.

Limaysaurus

Limaysaurus (“Limay lizard”) is a genus represented by a single species of rebbachisaurid sauropod dinosaurs, which lived during the mid-Cretaceous period, about 99.6 and 97 million years ago, in the Cenomanian, in what is now South America (northwestern Patagonia).

Nigersaurus

Nigersaurus is a genus of rebbachisaurid sauropod dinosaur that lived during the middle Cretaceous period, about 115 to 105 million years ago. It was discovered in the Elrhaz Formation in an area called Gadoufaoua, in the Republic of Niger. Fossils of this dinosaur were first described in 1976, but it was only named Nigersaurus taqueti in 1999, after further and more complete remains were found and described. The genus name means "Niger reptile", and the specific name honours the French palaeontologist Philippe Taquet, who discovered the first remains.

Small for a sauropod, Nigersaurus was about 9 metres (30 feet) long, and had a short neck. It weighed around four tonnes, comparable to a modern elephant. Its skeleton was highly pneumatised (filled with air spaces connected to air sacs), but the limbs were robustly built. Its skull was very specialised for feeding, with large fenestrae and thin bones. It had a wide muzzle filled with more than 500 teeth, which were replaced at a rapid rate: around every 14 days. The jaws may have borne a keratinous sheath. Unlike other tetrapods, the tooth-bearing bones of its jaws were rotated transversely relative to the rest of the skull, so that all of its teeth were located far to the front.

Nigersaurus and its closest relatives are grouped within the subfamily Rebbachisaurinae (formerly thought to be grouped in the eponymous Nigersaurinae) of the family Rebbachisauridae, which is part of the sauropod superfamily Diplodocoidea. Nigersaurus was probably a browser, and fed with its head close to the ground. The region of its brain that detected smell was underdeveloped, although its brain size was comparable to that of other dinosaurs. There has been debate on whether its head was habitually held downwards, or horizontally like other sauropods. It lived in a riparian habitat, and its diet probably consisted of soft plants, such as ferns, horsetails, and angiosperms. It is one of the most common fossil vertebrates found in the area, and shared its habitat with other dinosaurian megaherbivores, as well as large theropods and crocodylomorphs.

Nopcsaspondylus

Nopcsaspondylus (meaning "Nopsca's vertebra", in reference to the original describer) is a genus of rebbachisaurid sauropod dinosaur (a type of large, long-necked quadrupedal herbivorous dinosaur) from the Cenomanian-age (Upper Cretaceous) Candeleros Formation of Neuquén, Argentina. It is based on a now-lost back vertebra described by Nopcsa in 1902 but not named at the time. The specimen had a small vertebral body and large hollows, now known to be typical of rebbachisaurids.

Rayososaurus

Rayososaurus is a genus of plant-eating sauropod dinosaur of the superfamily Diplodocoidea. It was found in the Candeleros Formation, but was named Rayososaurus after the Rayoso Member, which later has been elevated to the older Rayoso Formation. The formations are located in the Neuquén Basin of northern Patagonia, Argentina. Rayososaurus lived during the Cenomanian epoch of the Late Cretaceous, about 99 to 96 million years ago. The type species is R. agrioensis, named by Argentinian paleontologist José Bonaparte in 1996. The species epithet agrioensis refers to the Agrio del Medio locality.

Rebbachisaurinae

Rebbachisaurinae is a subfamily within the family Rebbachisauridae, defined to include Rebbachisaurus garasbae and exclude Limaysaurus tessonei. It was first proposed as a rank by Jose Bonaparte in 1995, to include Rebbachisaurus. Some phylogenies however, include Rebbachisaurus in a clade with Limaysaurus, and thus the subfamily was not used. In 2015, a phylogenetic analysis was conducted, and it found Rebbachisaurus instead to be closer to Nigersaurus and related genera than Limaysaurus, and thus was used to replace Nigersaurinae as Rebbachisaurinae is the older term and is named after the genus used for the formation of the family Rebbachisauridae. The 2015 cladogram of Fanti et al. is shown below.

Tataouinea

Tataouinea is a genus of sauropod dinosaur (with a single species, Tataouinea hannibalis) in the subfamily Rebbachisaurinae of Rebbachisauridae which lived in the Early Cretaceous Tunisia. Its bones were extensively pneumatic, providing strong support for the theory that sauropods had birdlike respiratory systems. Key characteristics of its vertebral morphology show that Tatouinea was a rebbachisaurid, closely related to the nigersaurines of Europe. In 2015, more material of the holotype specimen uncovered after the initial description were analysed. These included additional tail vertebrae. A phylogenetic analysis was published alongside the paper, finding a clade of nigersaurines to include Rebbachisaurus, thus forcing the subfamily to be renamed Rebbachisaurinae.It bears the name of the region where it was discovered, Tataouine, and a punic military commander Hannibal.

Xenoposeidon

Xenoposeidon (meaning "strange or alien Poseidon", in allusion to Sauroposeidon) is a genus of rebbachisaurid sauropod dinosaur from the Early Cretaceous of England, living about 140 million years ago. It is known from a single partial vertebra with unusual features, unlike those of other sauropods. This bone was first discovered in the early 1890s but received little attention until it was found by University of Portsmouth student Mike Taylor, who formally described and named it in 2007 with Darren Naish.

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