Rebbachisaurus Garasbae
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''Rebbachisaurus'' (meaning " Aït Rebbach lizard") is a
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
sauropod Sauropoda (), whose members are known as sauropods (; from '' sauro-'' + '' -pod'', 'lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their b ...
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
of the superfamily
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, ...
, that lived during the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the more recent of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''cre ...
period in
Africa Africa is the world's second-largest and second-most populous continent after Asia. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 20% of Earth's land area and 6% of its total surfac ...
and possibly also
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...
about 99-97 million years ago. Remains attributed to ''Rebbachisaurus'' have been found in Morocco, Niger, Algeria, Tunisia and possibly also Argentina (if ''
Rayososaurus ''Rayososaurus'' is an extinct genus of sauropod dinosaur in the family Rebbachisauridae. ''Rayososaurus'' was named by Argentinian paleontologist José Bonaparte in 1996. Its type and only accepted species is ''Rayososaurus agrioensis''. The spe ...
'' is the same animal as ''Rebbachisaurus''), although only the Moroccan remains can be referred to the genus without doubt. The discovery of ''
Rayososaurus ''Rayososaurus'' is an extinct genus of sauropod dinosaur in the family Rebbachisauridae. ''Rayososaurus'' was named by Argentinian paleontologist José Bonaparte in 1996. Its type and only accepted species is ''Rayososaurus agrioensis''. The spe ...
'', a South American sauropod nearly identical to ''Rebbachisaurus'' which may have actually have been the same animal as ''Rebbachisaurus'', supports the theory that there was still a land connection between
Africa Africa is the world's second-largest and second-most populous continent after Asia. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 20% of Earth's land area and 6% of its total surfac ...
and
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...
during the Early Cretaceous, long after it was commonly thought the two continents had separated.Bonaparte, J. (1996). "Cretaceous tetrapods of Argentina". ''Münchner Geowissenschaften Abhandlungen'' 30: 73-130.


History of discovery

Between 8 October 1948 and 3 January 1952, Rene Lavocat collected the holotype of ''Rebbachisaurus garasbae'' (MNHN-MRS 1958), which consists of ten ribs, the right shoulder blade, eleven vertebrae, the sacrum, a humerus, and two bones probably belonging to the pelvis, in the Aoufous Formation at Gara Sbaa,
Errachidia Errachidia (), formerly known as Ksar es-Souk, () is a city in Morocco, located in the Errachidia Province, and is the capital of the Drâa-Tafilalet region. Toponymy Formerly known as "Ksar es-Souk" (), the city was renamed Errachidia around ...
in
Morocco Morocco, officially the Kingdom of Morocco, is a country in the Maghreb region of North Africa. It has coastlines on the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria to Algeria–Morocc ...
. He named the type species ''R. garasbae'' during the same year but only the shoulder blade and a single vertebra from the string of eleven were initially described, very briefly and without illustrations. The fossils were left undescribed and neglected until 2015, when Jeffrey Wilson and Ronan Allain cleaned the holotype and described the remaining fossils that Lavocat never described in 1954. A second specimen, consisting of a partial vertebra measuring up to tall if complete (also found in Aoufous) has also been assigned to ''R. garasbae''. A second species was named by de Lapparent in 1960, ''Rebbachisaurus tamesnensis''. The type specimen, collected by Lapparent in the Continental intercalaire Formation (initially believed to have been found in the Gall locality,
Tiourarén Formation The Tiourarén Formation is a geological formation in the Agadez Region of Niger whose strata were originally thought to be Early Cretaceous. However, re-interpretation of the sediments showed that they are probably Middle or Late Jurassic (Bath ...
) of
Niger Niger, officially the Republic of the Niger, is a landlocked country in West Africa. It is a unitary state Geography of Niger#Political geography, bordered by Libya to the Libya–Niger border, north-east, Chad to the Chad–Niger border, east ...
, supposedly composed of two humeri and two femora, and a second specimen composed of four isolated teeth, a dentary fragment with three teeth, over 100 vertebrae, six chevrons, 12 ribs, 5 scapulae, an ilium, two ischia, and numerous limb elements, however, the material of this taxon was collected from multiple localities across the
Sahara The Sahara (, ) is a desert spanning across North Africa. With an area of , it is the largest hot desert in the world and the list of deserts by area, third-largest desert overall, smaller only than the deserts of Antarctica and the northern Ar ...
, such as several sites in the
Elrhaz Formation The Elrhaz Formation is a geological Formation (geology), formation in Niger, West Africa. Its strata date back to the Early Cretaceous, about 125 to 112 million years ago. Dinosaur remains are among the fossils that have been recovered from the ...
of Niger, and is not referrable to ''Rebbachisaurus''. A third species was named by Calvo and Salgado in 1995, '' Rebbachisaurus tessonei''. It was discovered in the
Candeleros Formation The Candeleros Formation is a geologic formation that outcrop, crops out in the Rio Negro Province, Río Negro, Neuquen Province, Neuquén, and Mendoza Province, Mendoza provinces of northern Patagonia, Argentina. It is the oldest formation in the ...
in
Argentina Argentina, officially the Argentine Republic, is a country in the southern half of South America. It covers an area of , making it the List of South American countries by area, second-largest country in South America after Brazil, the fourt ...
, near to where the holotype of ''
Giganotosaurus carolinii ''Giganotosaurus'' ( ) is a genus of large theropod dinosaur that lived in what is now Argentina, during the early Cenomanian age of the Late Cretaceous period (geology), period, approximately 99.6 to 95 million years ago. The holotype specim ...
'' was discovered. This species was later moved to the new genus ''
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 to 94 million years ago, in the Cenomanian, in what is now South America (n ...
'' in 2004. It is often believed that the rebbachisaurid ''
Rayososaurus agrioensis ''Rayososaurus'' is an extinct genus of sauropod dinosaur in the family Rebbachisauridae. ''Rayososaurus'' was named by Argentinian paleontologist José Bonaparte in 1996. Its type and only accepted species is ''Rayososaurus agrioensis''. The spe ...
'', named by
Jose Bonaparte Jose is the English transliteration of the Hebrew and Aramaic name ''Yose'', which is etymologically linked to ''Yosef'' or Joseph. Given name Mishnaic and Talmudic periods *Jose ben Abin *Jose ben Akabya *Jose the Galilean *Jose ben Halafta *Jo ...
in 1996, is synonymous with ''Rebbachisaurus garasbae''. ''Rayososaurus'' is extremely similar to ''Rebbachisaurus'' and there is some debate as to whether ''Rayososaurus'' is indeed a separate genus. However, morphological and temporal differences tend to support the distinction, as ''Rayososaurus'' is known from the
Candeleros Formation The Candeleros Formation is a geologic formation that outcrop, crops out in the Rio Negro Province, Río Negro, Neuquen Province, Neuquén, and Mendoza Province, Mendoza provinces of northern Patagonia, Argentina. It is the oldest formation in the ...
of
Argentina Argentina, officially the Argentine Republic, is a country in the southern half of South America. It covers an area of , making it the List of South American countries by area, second-largest country in South America after Brazil, the fourt ...
, while ''Rebbachisaurus'' is known from the Aoufous Formation of
Morocco Morocco, officially the Kingdom of Morocco, is a country in the Maghreb region of North Africa. It has coastlines on the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria to Algeria–Morocc ...
.


Description

''Rebbachisaurus'' is a medium-sized sauropod, measuring around long and weighing up to . It possessed a small head, a long, graceful neck and a whiplike tail. ''Rebbachisaurus'' is distinguished from other sauropods by its unusually tall, ridged back and the spine or sail that lay atop its back, distinguished by the tall ridges of the preserved dorsal vertebrae of the holotype and other specimens.Apesteguía S, Gallina PA and Haluza A (2010) "Not just a pretty face: anatomical peculiarities in the postcranium of Rebbachisaurids (Sauropoda: Diplodocoidea)". Historical Biology, Volume 22, Issue 1-3: Proceedings of the First International Congress on North African Vertebrate Palaeontology.


Classification

Below is a
cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...
following the 2013 analysis by Fanti and colleagues, which confirmed the placement of ''Rebbachisaurus'' as a basal rebbachisaurid. A 2015 cladistic study by Wilsona and the French palaeontologist Ronan Allain found ''Rebbachisaurus'' itself to group with the nigersaurines, and the authors suggested that Nigersaurinae was therefore a
junior synonym In taxonomy, the scientific classification of living organisms, a synonym is an alternative scientific name for the accepted scientific name of a taxon. The botanical and zoological codes of nomenclature treat the concept of synonymy differently. ...
of Rebbachisaurinae (since that name would have priority).


Paleoecology

The most common vertebrates which coexisted alongside ''R. garasbae'' in the Aoufous belong to the
elasmobranch Elasmobranchii () is a subclass of Chondrichthyes or cartilaginous fish, including modern sharks ( division Selachii), and batomorphs (division Batomorphi, including rays, skates, and sawfish). Members of this subclass are characterised by h ...
fish '' Onchopristis numidus''. Another seven elasmobranchs are reported: '' Asteracanthus aegyptiacus'', '' Distobatus nutiae'', '' Tribodus'' sp., ''
Lissodus ''Lissodus'' is an extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to ...
'' sp., '' Haimirichia amonensis'', Cretoxyrhinidae indet., and '' Marckgrafia lybica''. Lungfish fossils referred to '' Ceratodus humei'' and '' Neoceratodus africanus'' are known to have coexisted alongside ''Rebbachisaurus'' and the
coelacanth Coelacanths ( ) are an ancient group of lobe-finned fish (Sarcopterygii) in the class Actinistia. As sarcopterygians, they are more closely related to lungfish and tetrapods (the terrestrial vertebrates including living amphibians, reptiles, bi ...
s '' Mawsonia lavocati'' and ''
Axelrodichthys ''Axelrodichthys'' is an extinct genus of mawsoniidae, mawsoniid coelacanth from the Cretaceous of Africa, North America, North and South America, and Europe. Several species are known, the remains of which were discovered in the Lower Cretaceous ...
'' also coexisted with ''Rebbachisaurus''. Several taxa of
Cladistia Cladistia is a subclass of bony fishes whose only living members are the bichirs of tropical Africa. Their major synapomorphies are a heterocercal tail in which the dorsal fin has independent rays, and a posteriorly elongated parasphenoid. ...
and two genera of seminiomorphs (an unnamed ''
Lepidotes ''Lepidotes'' (from , 'covered with scales') (previously known as ''Lepidotus'') is an extinct genus of Mesozoic ray-finned fish. It has long been considered a wastebasket taxon, characterised by "general features, such as thick rhomboid scales ...
''-like species and '' Oniichthys falipoui'') are known to have coexisted with ''Rebbachisaurus'' in the Aoufous Formation.
Teleostean Teleostei (; Greek ''teleios'' "complete" + ''osteon'' "bone"), members of which are known as teleosts (), is, by far, the largest group of ray-finned fishes (class Actinopterygii), with 96% of all extant species of fish. The Teleostei, which i ...
s are represented by '' Cladocyclus pankowskii'', '' Palaeonotopterus greenwoodi'', '' Erfoudichthys rosae'' and '' Concavotectum moroccensis''.
Amphibians Amphibians are ectothermic, anamniote, anamniotic, tetrapod, four-limbed vertebrate animals that constitute the class (biology), class Amphibia. In its broadest sense, it is a paraphyletic group encompassing all Tetrapod, tetrapods, but excl ...
are also present: '' Kababisha'' sp., the pipid frog '' Oumtkoutia anae'' and non-pipids frogs.
Turtle Turtles are reptiles of the order (biology), order Testudines, characterized by a special turtle shell, shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Crypt ...
s are represented by several species: '' Dirqadim schaefferi'', the podocnemidids '' Hamadachelys escuilliei'', the bothremydids '' Galianemys whitei'' and ''G. emringeri'', and the araripemydids ''
Araripemys ''Araripemys'' is an extinct marine turtle genus from 112 to 109 million years ago, in the Early Cretaceous Crato and Romualdo Formations of the Araripe Basin in northeastern Brazil Brazil, officially the Federative Republic of Brazil, ...
'' sp.
Crocodilian Crocodilia () is an Order (biology), order of semiaquatic, predatory reptiles that are known as crocodilians. They first appeared during the Late Cretaceous and are the closest living relatives of birds. Crocodilians are a type of crocodylomorp ...
s are commonly found; four species are present: '' Elosuchus cherifiensis'', a genus that belonged to the
Trematochampsidae Trematochampsidae is an extinct family of mesoeucrocodylian crocodyliforms. Fossils are present from Madagascar, Morocco, Niger, Argentina, and Brazil (in the case of '' Caririsuchus'', where some specimens have been found in the Romualdo Membe ...
, '' Araripesuchus rattoides'' and ''
Laganosuchus maghrebensis ''Laganosuchus'' is an extinct genus of Stomatosuchidae, stomatosuchid crocodyliform. Fossils have been found from Niger and Morocco and date back to the Upper Cretaceous. Discovery The name means "pancake crocodile" from the Ancient Greek, Gre ...
''. ''Rebbachisaurus garasbae'' coexisted with many dinosaurs, including an
ornithischia Ornithischia () is an extinct clade of mainly herbivorous dinosaurs characterized by a pelvic structure superficially similar to that of birds. The name ''Ornithischia'', or "bird-hipped", reflects this similarity and is derived from the Greek ...
n and an unnamed basal
theropod Theropoda (; from ancient Greek , (''therion'') "wild beast"; , (''pous, podos'') "foot"">wiktionary:ποδός"> (''pous, podos'') "foot" is one of the three major groups (clades) of dinosaurs, alongside Ornithischia and Sauropodom ...
, both only known from footprints. Among theropods, two
carcharodontosaurid Carcharodontosauridae (carcharodontosaurids; from the Greek καρχαροδοντόσαυρος, ''carcharodontósauros'': "shark-toothed lizards") is a group of carnivorous theropod dinosaurs. In 1931, Ernst Stromer named Carcharodontosauridae ...
s ('' Carcharodontosaurus saharicus'' and '' Sauroniops pachytholus''), one
spinosaur Spinosauridae (or spinosaurids) is a clade or family of tetanuran theropod dinosaurs comprising ten to seventeen known genera. Spinosaurid fossils have been recovered worldwide, including Africa, Europe, South America, and Asia. Their remains have ...
id (''
Spinosaurus aegyptiacus ''Spinosaurus'' (; ) is a genus of large spinosaurid theropod dinosaurs that lived in what now is North Africa during the Cenomanian stage of the Late Cretaceous period, about 100 to 94 million years ago. The genus was known first from ...
''), one
coelurosaur Coelurosauria (; from Greek, meaning "hollow-tailed lizards") is the clade containing all theropod dinosaurs more closely related to birds than to carnosaurs. Coelurosauria is a subgroup of theropod dinosaurs that includes compsognathids, ty ...
(''
Deltadromeus agilis ''Deltadromeus'' (meaning "delta runner") is an extinct genus of controversial theropod dinosaurs from the Aoufous Formation (Kem Kem Beds) of Morocco. The genus contains a single species, ''Deltadromeus agilis'', known from multiple partial ...
''), an unnamed
abelisaur Ceratosaurs are members of the clade Ceratosauria, a group of dinosaurs defined as all theropods sharing a more recent common ancestor with '' Ceratosaurus'' than with birds. The oldest known ceratosaur, '' Saltriovenator'', dates to the earlie ...
and an unnamed
dromaeosaur 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 Gr ...
also coexisted with ''Rebbachisaurus''.
Pterosaur Pterosaurs are an extinct clade of flying reptiles in the order Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 million to 66 million years ago). Pterosaurs are the earli ...
s are also present, although their
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserve ...
s are extremely rare and enigmatic: an
azhdarchid Azhdarchidae (from the Persian word , , a dragon-like creature in Persian mythology) is a family of pterosaurs known primarily from the Late Cretaceous Period, though an isolated vertebra apparently from an azhdarchid is known from the Early Cre ...
, an
ornithocheirid Anhangueridae (alternatively called Ornithocheiridae, meaning "bird hands") is a group of pterosaurs within the suborder Pterodactyloidea. These pterosaurs were among the last to possess teeth. Members that belong to this group lived from the ...
, a
tapejarid Tapejaridae (from a Tupi word meaning 'the lord of the ways') is a family of azhdarchoid pterosaurs from the Cretaceous period. Members are currently known from Brazil, England, Hungary, Morocco, Spain, the United States, and China. The most p ...
and a
pteranodontid The Pteranodontidae are a family of large pterosaurs from the Late Cretaceous of North America and possibly other continents including Europe and Africa. The family was named in 1876 by Othniel Charles Marsh. Pteranodontids had a distinctive, elon ...
are known to have coexisted with ''Rebbachisaurus''.L. Cavin, H. Tong, L. Boudad, C. Meister, A. Piuz, J. Tabouelle, M. Aarab, R. Amiot, E. Buffetaut, G. Dyke, S. Hua and J. Le Loeuff. (2010). Vertebrate assemblages from the early Late Cretaceous of southeastern Morocco: An overview. ''Journal of African Earth Sciences'' 57:391-412


References

{{Taxonbar, from=Q133675 Rebbachisauridae Dinosaur genera Cenomanian dinosaurs Taxa named by René Lavocat Fossil taxa described in 1954 Dinosaurs of Morocco