Rebbachisaurus Garasbae
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''Rebbachisaurus'' (meaning " Aït Rebbach lizard") is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
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 bo ...
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
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 younger 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 ''creta'', the ...
period in
Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...
and possibly also
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...
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 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 Fo ...
'' 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 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 Fo ...
'', 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 in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...
and
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...
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 The Aoufous Formation is a geological formation that contains some of the vertebrate assemblage of the Kem Kem Group, of Late Cretaceous date.Cavin et al., 2010, p.391Dal Sasso, C., Maganuco, S., Buffetaut, E., Mendez, M.A., 2005. New information o ...
at
Gara Sbaa ''Gara'' (Basque language, Basque: ''We Are'') is a bilingual (Basque/Spanish language, Spanish) newspaper published in the city of San Sebastián, Donostia-San Sebastián in the Basque Country (autonomous community), Basque Autonomous Commun ...
,
Errachidia Errachidia ( ar, الرشيدية, Berber: ⵉⵎⴻⵜⵖⴰⵔⵏ Imetɣaren) is a city in Morocco, located in the Errachidia Province, and is the capital of the Drâa-Tafilalet region. The city's residents speak Berber and Moroccan Arabic. ...
in
Morocco Morocco (),, ) officially the Kingdom of Morocco, is the westernmost country in the Maghreb region of North Africa. It overlooks the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria to ...
. 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 (geology), 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 Jur ...
) of
Niger ) , official_languages = , languages_type = National languagesSahara , photo = Sahara real color.jpg , photo_caption = The Sahara taken by Apollo 17 astronauts, 1972 , map = , map_image = , location = , country = , country1 = , ...
, such as several sites in the
Elrhaz Formation The Elrhaz Formation is a geological formation in Niger, central 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 formation, alongsi ...
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 (formerly known as the Candeleros Member of the "Río Limay Formation") is a geologic formation that crops out in the Río Negro, Neuquén, and Mendoza provinces of northern Patagonia, Argentina. It is the oldest formati ...
in
Argentina Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the second-largest country in South America after Brazil, th ...
, near to where the holotype of ''
Giganotosaurus carolinii ''Giganotosaurus'' ( ) is a genus of theropod dinosaur that lived in what is now Argentina, during the early Cenomanian age of the Late Cretaceous period, approximately 99.6 to 95 million years ago. The holotype specimen was discovered in th ...
'' 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 97 million years ago, in the Cenomanian, in what is now South America ( ...
'' in 2004. It is often believed that the rebbachisaurid '' Rayososaurus agrioensis'', named by
Jose Bonaparte Jose is the English transliteration of the Hebrew and Aramaic name ''Yose'', which is etymologically linked to ''Yosef'' or Joseph. The name was popular during the Mishnaic and Talmudic periods. * Jose ben Abin * Jose ben Akabya *Jose the Galile ...
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 (formerly known as the Candeleros Member of the "Río Limay Formation") is a geologic formation that crops out in the Río Negro, Neuquén, and Mendoza provinces of northern Patagonia, Argentina. It is the oldest formati ...
of
Argentina Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the second-largest country in South America after Brazil, th ...
, while ''Rebbachisaurus'' is known from the
Aoufous Formation The Aoufous Formation is a geological formation that contains some of the vertebrate assemblage of the Kem Kem Group, of Late Cretaceous date.Cavin et al., 2010, p.391Dal Sasso, C., Maganuco, S., Buffetaut, E., Mendez, M.A., 2005. New information o ...
of
Morocco Morocco (),, ) officially the Kingdom of Morocco, is the westernmost country in the Maghreb region of North Africa. It overlooks the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria to ...
.


Description

In 2010,
Gregory S. Paul Gregory Scott Paul (born December 24, 1954) is an American freelance researcher, author and illustrator who works in paleontology, and more recently has examined sociology and theology. He is best known for his work and research on theropod dino ...
estimated ''Rebbachisaurus'' at and .Paul, G.S., 2010, ''The Princeton Field Guide to Dinosaurs'', Princeton University Press p. 186
Holtz Holtz is the surname of: * Alexander Holtz (born 2002), Swedish ice hockey player * Andrew Holtz, American journalist * Carl Holtz (1920–2006), American oarsman and farmer * Daniel Holtz, a fictional character on the TV series ''Angel'' * Eric H ...
gave a bigger length of .Holtz, Thomas R. Jr. (2011) ''Dinosaurs: The Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages,'
Winter 2010 Appendix.
/ref> In 2020 Molina-Pérez and Larramendi gave a larger estimation of 26 meters (85.3 ft) and 40 tonnes (44 short tons). 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 ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...
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 The Botanical and Zoological Codes of nomenclature treat the concept of synonymy differently. * In botanical nomenclature, a synonym is a scientific name that applies to a taxon that (now) goes by a different scientific name. For example, Linna ...
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 sharks (superorder Selachii), rays, skates, and sawfish (superorder Batoidea). Members of this subclass are characterised by having five to seven pairs of gil ...
fish ''
Onchopristis numidus ''Onchopristis'' is an extinct genus of Sclerorhynchoidei, sclerorhynchoid from the Cretaceous of North Africa, Europe, and North America. Its name is derived from the Ancient Greek ''ónkos'' (wiktionary: ὄγκος, ὄγκος, 'barb') and ' ...
''. Another seven elasmobranchs are reported: ''
Asteracanthus aegyptiacus ''Asteracanthus'' (from el, ἀστήρ , 'star' and el, ἄκανθα , 'spine') is an extinct genus of hybodontiform, known from the Middle Jurassic (Bathonian) to the Early Cretaceous (Valanginian). Description Astercanthus was among the ...
'', '' Distobatus nutiae'', ''
Tribodus ''Tribodus'' is an extinct genus of hybodont shark from the Romualdo Formation of northeastern Brazil. It lived during the Early Cretaceous period (Aptian to Albian The Albian is both an age of the geologic timescale and a stage in the stratig ...
'' sp., ''
Lissodus ''Lissodus'' is an extinct genus of freshwater shark. It lived from the Early Carboniferous stage to the Albian age of the Cretaceous. It was about long and had flat teeth that it used for eating clams Clam is a common name for several ...
'' sp., ''
Haimirichia amonensis ''Haimirichia'' is an extinct genus of mackerel shark from the late Cenomanian, Cenomanian epoch of the Cretaceous period. It currently contains a single species: ''H. amonensis''. Teeth now assigned to this genus were originally attributed to t ...
'',
Cretoxyrhinidae ''Cretoxyrhina'' (; meaning 'Cretaceous sharp-nose') is an extinct genus of large mackerel shark that lived about 107 to 73 million years ago during the late Albian to late Campanian of the Late Cretaceous. The type species, ''C. mantelli'', is m ...
indet., and '' Marckgrafia lybica''. Lungfish fossils referred to ''
Ceratodus humei ''Ceratodus'' (from el, κέρας , 'horn' and el, ὀδούς 'tooth') was a wide-ranging genus of extinct lungfish. Fossil evidence dates back to the Early Triassic. A wide range of fossil species from different time periods have been found ...
'' and ''
Neoceratodus africanus ''Neoceratodus'' is a genus of lungfish in the family Neoceratodontidae. The extant Australian lungfish (''Neoceratodus forsteri'') is the only surviving member of this genus, but it was formerly much more widespread, being distributed throughout ...
'' are known to have coexisted alongside ''Rebbachisaurus'' and the
coelacanth The coelacanths ( ) are fish belonging to the order Actinistia that includes two extant species in the genus ''Latimeria'': the West Indian Ocean coelacanth (''Latimeria chalumnae''), primarily found near the Comoro Islands off the east coast ...
s ''
Mawsonia lavocati ''Mawsonia'' is an extinct genus of prehistoric coelacanth fish. It is the amongst the largest of all coelacanths, with one quadrate specimen (DGM 1.048-P) possibly belonging to an individual measuring in length. It lived in freshwater and brac ...
'' ''
Axelrodichthys ''Axelrodichthys'' is an extinct genus of mawsoniid coelacanth from the Cretaceous of Africa, North and South America, and Europe. Several species are known, the remains of which were discovered in the Lower Cretaceous (Aptian-Albian) of Brazil ...
'' also coexisted with ''Rebbachisaurus''. Several taxa of
Cladistia Cladistia is a clade of bony fishes whose only living members are the bichirs. Their major synapomorphies are a heterocercal tail in which the dorsal fin has independent rays, and a posteriorly elongated parasphenoid. Cladistia are the earlie ...
and two genera of seminiomorphs (an unnamed ''
Lepidotes ''Lepidotes'' (from el, λεπιδωτός , 'covered with scales') (previously known as ''Lepidotus'') is an extinct genus of Mesozoic ray-finned fish. It has been considered a wastebasket taxon, characterised by "general features, such as thic ...
''-like species and '' Oniichthys falipoui'') are known to have coexisted with ''Rebbachisaurus'' in the Aoufous Formation.
Teleostean Actinopterygii (; ), members of which are known as ray-finned fishes, is a class of bony fish. They comprise over 50% of living vertebrate species. The ray-finned fishes are so called because their fins are webs of skin supported by bony or ho ...
s are represented by '' Cladocyclus pankowskii'', '' Palaeonotopterus greenwoodi'', '' Erfoudichthys rosae'' and '' Concavotectum moroccensis''.
Amphibians Amphibians are four-limbed and ectothermic vertebrates of the class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terrestrial, fossorial, arbore ...
are also present: '' Kababisha'' sp., the pipid frog '' Oumtkoutia anae'' and non-pipids frogs.
Turtle Turtles are an order of reptiles known as Testudines, characterized by a special shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Cryptodira (hidden necked tu ...
s are represented by several species: '' Dirqadim schaefferi'', the podocnemidids '' Hamadachelys escuilliei'', the bothremydids ''
Galianemys whitei ''Galianemys'' is an extinct genus of turtle in the family Bothremydidae, discovered in the Kem Kem Beds. References Further reading

* Gaffney E. S., Tong H., et al (2002) Galianemys, a new side-necked turtle (Pelomedusoides: Bothremydid ...
'' 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. It is one of the oldest known pleurodire The Pleurodira ar ...
'' sp.
Crocodilian Crocodilia (or Crocodylia, both ) is an order of mostly large, predatory, semiaquatic reptiles, known as crocodilians. They first appeared 95 million years ago in the Late Cretaceous period ( Cenomanian stage) and are the closest living ...
s are commonly found; four species are present: ''
Elosuchus cherifiensis ''Elosuchus'' is an extinct genus of neosuchian crocodyliform that lived during the Middle Cretaceous of what is now Africa (Niger, Morocco Morocco (),, ) officially the Kingdom of Morocco, is the westernmost country in the Maghreb reg ...
'', a genus that belonged to the
Trematochampsidae Trematochampsidae is an extinct family of mesoeucrocodylian crocodylomorphs. Fossils are present from Madagascar, Morocco, Niger, Argentina, and Brazil (in the case of '' Caririsuchus'', where some specimens have been found in the Romualdo Memb ...
, ''
Araripesuchus rattoides ''Araripesuchus'' is a genus of extinct crocodyliform that existed during the Cretaceous period of the late Mesozoic era some 125 to 66 million years ago. Six species of ''Araripesuchus'' are currently known. They are generally considered to be n ...
'' and '' Laganosuchus maghrebensis''. ''Rebbachisaurus garasbae'' coexisted with many dinosaurs, including an
ornithischia Ornithischia () is an extinct order 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 s ...
n and an unnamed basal
theropod Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally c ...
, 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 ''Carcharodontosaurus'' (; ) is a genus of large Carcharodontosauridae, carcharodontosaurid Theropoda, theropod dinosaur that existed during the Cenomanian age of the Late Cretaceous in Northern Africa. The genus ''Carcharodontosaurus'' is na ...
'' and ''
Sauroniops pachytholus ''Sauroniops'' is a genus of carnivorous basal carcharodontosaurid theropod dinosaur known from the Late Cretaceous (Cenomanian stage) of Morocco. Discovery In the early twenty-first century a collector donated a dinosaur skull bone to the Ital ...
''), one
spinosaur The Spinosauridae (or spinosaurids) are a clade or family of tetanuran theropod dinosaurs comprising ten to seventeen known genera. They came into prominence during the Cretaceous period. Spinosaurid fossils have been recovered worldwide, includi ...
id (''
Spinosaurus aegyptiacus ''Spinosaurus'' (; ) is a genus of spinosaurid dinosaur that lived in what now is North Africa during the Cenomanian to upper Turonian stages of the Late Cretaceous period, about 99 to 93.5 million years ago. The genus was known first fro ...
''), 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, tyrann ...
('' Deltadromeus agilis''), an unnamed
abelisaur Abelisauroidea is typically regarded as a Cretaceous group, though the earliest abelisauridae remains are known from the Middle Jurassic of Argentina (classified as the species Eoabelisaurus mefi) and possibly Madagascar (fragmentary remains o ...
and an unnamed
dromaeosaur 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 ('), meaning ...
also coexisted with ''Rebbachisaurus''.
Pterosaur Pterosaurs (; from Greek ''pteron'' and ''sauros'', meaning "wing lizard") is 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 to ...
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 preserved ...
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 Cret ...
, an
ornithocheirid Ornithocheiridae (or ornithocheirids, 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 Early to Late Cr ...
, a
tapejarid Tapejaridae (from a Tupi word meaning "the old being") are a family of pterodactyloid pterosaurs from the Cretaceous period. Members are currently known from Brazil, England, Hungary, Morocco, Spain, the United States, and China. The most primiti ...
and a
pteranodontid The Pteranodontidae are a Family (biology), family of large pterosaurs of the Cretaceous Period of North America and Africa. The family was named in 1876 by Othniel Charles Marsh. Pteranodontids had a distinctive, elongated crest jutting from the ...
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 Rebbachisaurids Cenomanian life Late Cretaceous dinosaurs of Africa Cretaceous Morocco Fossils of Morocco Fossil taxa described in 1954 Taxa named by René Lavocat