Tangasaurus
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''Tangasaurus'' is an
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
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 aquatic basal
tangasaurid Tangasauridae is a family of diapsids. Specimens have been found that are of Late Permian to Early Triassic in age from the Sakamena Group of western Madagascar. They lived alongside other taxa present from the Sakamena Group, including temnos ...
neodiapsid Neodiapsida is a clade, or major branch, of the reptilian family tree, typically defined as including all diapsids apart from some early primitive types known as the araeoscelidians. Modern reptiles and birds belong to the neodiapsid subclade ...
known from the
Late Permian Late may refer to: * LATE, an acronym which could stand for: ** Limbic-predominant age-related TDP-43 encephalopathy, a proposed form of dementia ** Local-authority trading enterprise, a New Zealand business law ** Local average treatment effect, ...
period Period may refer to: Common uses * Era, a length or span of time * Full stop (or period), a punctuation mark Arts, entertainment, and media * Period (music), a concept in musical composition * Periodic sentence (or rhetorical period), a concept ...
(late
Changhsingian In the geologic time scale, the Changhsingian or Changxingian is the latest age or uppermost stage of the Permian. It is also the upper or latest of two subdivisions of the Lopingian Epoch or Series. The Changhsingian lasted from to 251.902 mill ...
stage) of
Tanga Tanga may refer to: Places Burkina Faso * , a town in eastern Burkina Faso * Tanga, Sidéradougou, a village in western Burkina Faso * Tanga-Pela, a village in northern-central Burkina Faso Other places * Tanga, Tanzania, a city and port on th ...
, northeastern
Tanzania Tanzania (; ), officially the United Republic of Tanzania ( sw, Jamhuri ya Muungano wa Tanzania), is a country in East Africa within the African Great Lakes region. It borders Uganda to the north; Kenya to the northeast; Comoro Islands and ...
. It contains a single species, ''Tangasaurus mennelli''.


Discovery

''Tangasaurus'' is known only from three
syntype In biological nomenclature, a syntype is any one of two or more biological types that is listed in a description of a taxon where no holotype was designated. Precise definitions of this and related terms for types have been established as part of ...
s which represent two individuals. The small individual, which is well preserved in two specimens ( part and counterpart) and is known from a partial
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, the ...
and a nearly complete
postcranial Postcrania (postcranium, adjective: postcranial) in zoology and vertebrate paleontology is all or part of the skeleton apart from the skull. Frequently, fossil remains, e.g. of dinosaurs or other extinct tetrapods, consist of partial or isolated sk ...
skeleton A skeleton is the structural frame that supports the body of an animal. There are several types of skeletons, including the exoskeleton, which is the stable outer shell of an organism, the endoskeleton, which forms the support structure inside ...
, was designated as the
lectotype In biology, a type is a particular specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally attached. In other words, a type is an example that serves to anchor or centralizes the ...
of the genus. The part slab (showing the partial skull and postcranial) is hosted at the Bulawayo Museum,
Zimbabwe Zimbabwe (), officially the Republic of Zimbabwe, is a landlocked country located in Southeast Africa, between the Zambezi and Limpopo Rivers, bordered by South Africa to the south, Botswana to the south-west, Zambia to the north, and Mozam ...
, while the counterpart slab SAM 6231 (showing the other side of the postcranial) is hosted at the
South African Museum The Iziko South African Museum is a South African national museum located in Cape Town. The museum was founded in 1825, the first in the country. It has been on its present site in the Company's Garden since 1897. The museum houses important A ...
,
Cape Town Cape Town ( af, Kaapstad; , xh, iKapa) is one of South Africa's three capital cities, serving as the seat of the Parliament of South Africa. It is the legislative capital of the country, the oldest city in the country, and the second largest ...
. The third syntype, SAM 6232, represents a nearly complete postcranial skeleton from a larger individual, but it is poorly preserved. All specimens were collected by F. P. Mennell in 1922 at the Mizimbazi River near Tanga City, northeastern
Tanzania Tanzania (; ), officially the United Republic of Tanzania ( sw, Jamhuri ya Muungano wa Tanzania), is a country in East Africa within the African Great Lakes region. It borders Uganda to the north; Kenya to the northeast; Comoro Islands and ...
. They were collected from the Ruhuhu Valley, in the middle part of the Tanga beds, dating to the late
Changhsingian In the geologic time scale, the Changhsingian or Changxingian is the latest age or uppermost stage of the Permian. It is also the upper or latest of two subdivisions of the Lopingian Epoch or Series. The Changhsingian lasted from to 251.902 mill ...
stage of the
Lopingian The Lopingian is the uppermost series/last epoch of the Permian. It is the last epoch of the Paleozoic. The Lopingian was preceded by the Guadalupian and followed by the Early Triassic. The Lopingian is often synonymous with the informal terms l ...
series Series may refer to: People with the name * Caroline Series (born 1951), English mathematician, daughter of George Series * George Series (1920–1995), English physicist Arts, entertainment, and media Music * Series, the ordered sets used in ...
, about 252.5–251 
million years ago The abbreviation Myr, "million years", is a unit of a quantity of (i.e. ) years, or 31.556926 teraseconds. Usage Myr (million years) is in common use in fields such as Earth science and cosmology. Myr is also used with Mya (million years ago). ...
. Although Mennell reported a presence of eight nearly complete individuals from that site, subsequent work couldn't find any evidence for the missing 6 individuals. Mennell's report, alongside some reptilian fragments that were discovered in 1913 and reported by Janensch (1927), are the only reptilian fossils currently known from the Tanga beds. Numerous well preserved specimens from various Late Permian localities of
Madagascar Madagascar (; mg, Madagasikara, ), officially the Republic of Madagascar ( mg, Repoblikan'i Madagasikara, links=no, ; french: République de Madagascar), is an island country in the Indian Ocean, approximately off the coast of East Africa ...
were initially identified as ''Tangasaurus'' by Piveteau (1926). During that time the
Mozambique Channel The Mozambique Channel (french: Canal du Mozambique, mg, Lakandranon'i Mozambika, pt, Canal de Moçambique) is an arm of the Indian Ocean located between the Southeast African countries of Madagascar and Mozambique. The channel is about long ...
had just started to open. However, a reexamination of these specimens by Carroll (1981) and Currie (1982) has shown that they represent a different genus, ''
Thadeosaurus ''Thadeosaurus'' is an extinct genus of diapsid reptile belonging to the family Younginidae. Fossils have been found in the Lower Sakamena Formation ( Sakamena Group) of the Morondava Basin, Madagascar in 1981, and date to the late Permian to ...
'', known only from Madagascar. Another specimen from Madagascar,
MNHN The French National Museum of Natural History, known in French as the ' (abbreviation MNHN), is the national natural history museum of France and a ' of higher education part of Sorbonne Universities. The main museum, with four galleries, is loca ...
 1908-32-57, was also tentatively referred to ''Tangasaurus''. However, Currie (1980) found it to represent a new and distinct genus which he named ''
Acerosodontosaurus ''Acerosodontosaurus'' is an extinct genus of neodiapsid reptiles that lived during the Upper Permian of Madagascar. The only species of ''Acerosodontosaurus'', ''A. piveteaui'', is known from a natural mold of a single partial skeleton including ...
''.


Description

''Tangasaurus'' was described and named by
Sidney H. Haughton Sidney Henry Haughton FRS (7 May 1888 – 24 May 1982) was an English-born South African paleontologist and geologist best known for his description of the sauropodomorph dinosaur '' Melanorosaurus'' in 1924, and his work on the geology of ...
in
1924 Events January * January 12 – Gopinath Saha shoots Ernest Day, whom he has mistaken for Sir Charles Tegart, the police commissioner of Calcutta, and is arrested soon after. * January 20– 30 – Kuomintang in China hol ...
who found it to be a probable
diapsid Diapsids ("two arches") are a clade of sauropsids, distinguished from more primitive eureptiles by the presence of two holes, known as temporal fenestrae, in each side of their skulls. The group first appeared about three hundred million years ago ...
reptile that, because of the long, powerful, flattened tail, had become adapted for an aquatic existence. Contrary to Haughton, Piveteau (1926) considered ''Tangasaurus'' to be a primarily terrestrial animal. Following Nopcsa (1924), he considered it to be related to the ''
Araeoscelis ''Araeoscelis'' (from el, αραιά , 'thin' and el, σκελίς , 'ribs of beef') is an extinction, extinct genus of reptile, and one of the earliest diapsids. Fossils have been found in the Nocona Formation, Nocona, Arroyo Formation, Arroyo ...
'', '' Kadaliosaurus'', '' Broomia'' and the "eosuchians" (a defunct clade that used to unite all diapsids more advanced than ''
Araeoscelis ''Araeoscelis'' (from el, αραιά , 'thin' and el, σκελίς , 'ribs of beef') is an extinction, extinct genus of reptile, and one of the earliest diapsids. Fossils have been found in the Nocona Formation, Nocona, Arroyo Formation, Arroyo ...
'') ''
Saurosternon ''Saurosternon'' is an extinct genus of neodiapsid reptile from the Late Permian of South Africa. It is based on a partial skeleton split between two slabs of sandstone from the ''Daptocephalus'' Assemblage Zone. ''Saurosternon'' was one of th ...
'' and ''
Pleurosaurus ''Pleurosaurus'' (meaning "side lizard") is an extinct genus of diapsid reptiles belonging to the group Sphenodontia, extinct relatives of the modern tuatara. ''Pleurosaurus'' fossils were discovered in the Solnhofen Limestone of Bavaria, Germ ...
''. Described by Piveteau (1926), '' Hovasaurus boulei'' from Madagascar was considered to be related to ''
Mesosaurus ''Mesosaurus'' (meaning "middle lizard") is an extinct genus of reptile from the Early Permian of southern Africa and South America. Along with it, the genera '' Brazilosaurus'' and ''Stereosternum'', it is a member of the family Mesosauridae a ...
''. Although not as specialized as ''Mesosaurus'' for living in the water, Piveteau noted its short neck, short manus, well developed haemal spines and slight pachyostosis of the ribs. Haughton (1930) restudied Piveteau's specimens from Madagascar, concluding that ''Tangasaurus'' (then included the Malagasy specimens) and ''Hovasaurus'' were allied and that both were diapsids. ''Tangasaurus'' was considered to be morphologically intermediate between ''
Youngina ''Youngina'' is an extinct genus of diapsid reptile from the Late Permian Beaufort Group ('' Tropidostoma''-''Dicynodon'' zones) of the Karoo Red Beds of South Africa. This, and a few related forms, make up the family Younginidae, within the Ord ...
'' and ''Hovasaurus'' which was recognized as an aquatic reptile due to its short
forelimb A forelimb or front limb is one of the paired articulated appendages (limbs) attached on the cranial ( anterior) end of a terrestrial tetrapod vertebrate's torso. With reference to quadrupeds, the term foreleg or front leg is often used instead. ...
and
coracoid A coracoid (from Greek κόραξ, ''koraks'', raven) is a paired bone which is part of the shoulder assembly in all vertebrates except therian mammals (marsupials and placentals). In therian mammals (including humans), a coracoid process is prese ...
, small
ossification Ossification (also called osteogenesis or bone mineralization) in bone remodeling is the process of laying down new bone material by cells named osteoblasts. It is synonymous with bone tissue formation. There are two processes resulting in t ...
and elongated body. Piveteau (1926) included ''Broomia'', ''Saurosternon'' and ''Tangasaurus'' in the Tangasaurinae. After Haughton's paper (1930), ''Tangasaurus'' and ''Hovasaurus'' were usually included as the only representatives of the family
Tangasauridae Tangasauridae is a family of diapsids. Specimens have been found that are of Late Permian to Early Triassic in age from the Sakamena Group of western Madagascar. They lived alongside other taxa present from the Sakamena Group, including temnosp ...
. The known specimens of ''Tangasaurus'' were redescribed by
Philip J. Currie Philip John Currie (born March 13, 1949) is a Canadian palaeontologist and museum curator who helped found the Royal Tyrrell Museum of Palaeontology in Drumheller, Alberta and is now a professor at the University of Alberta in Edmonton. In the ...
(1982). He diagnosed the genus on the basis of two
autapomorphies In phylogenetics, an autapomorphy is a distinctive feature, known as a derived trait, that is unique to a given taxon. That is, it is found only in one taxon, but not found in any others or outgroup taxa, not even those most closely related to t ...
: It possesses high and rectangular neural spines of the
dorsal vertebrae In vertebrates, thoracic vertebrae compose the middle segment of the vertebral column, between the cervical vertebrae and the lumbar vertebrae. In humans, there are twelve thoracic vertebrae and they are intermediate in size between the cervical ...
and the height of
neural spine The spinal column, a defining synapomorphy shared by nearly all vertebrates,Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic i ...
of mid-caudal vertebra about 35% greater than length of associated centrum, and about 75% length of associated haemal arch and spine. The largest "tangasaurid" (sensu Currie, 1982) is ''Hovasaurus'' with an estimated maximum snout-vent length of about . The largest specimen of ''Tangasaurus'' is 20% smaller than that, but its incomplete ossification suggests that larger specimens probably existed. Currie (1982) united to subfamilies within the Tangasauridae: Kenyasaurinae (that he named to include ''
Kenyasaurus ''Kenyasaurus'' is an extinct genus of basal tangasaurid known from the Early Triassic period of Coast Province, southeastern Kenya. It contains a single species, ''Kenyasaurus mariakaniensis''. Discovery ''Kenyasaurus'' is known only from the ...
'' and ''
Thadeosaurus ''Thadeosaurus'' is an extinct genus of diapsid reptile belonging to the family Younginidae. Fossils have been found in the Lower Sakamena Formation ( Sakamena Group) of the Morondava Basin, Madagascar in 1981, and date to the late Permian to ...
'') and Tangasaurinae (to include Tangasauridae sensu Haughton, 1930). He allied Tangasauridae and ''Youngina'' together within superfamily Younginoidea which he named. Currie (1980) named ''
Acerosodontosaurus ''Acerosodontosaurus'' is an extinct genus of neodiapsid reptiles that lived during the Upper Permian of Madagascar. The only species of ''Acerosodontosaurus'', ''A. piveteaui'', is known from a natural mold of a single partial skeleton including ...
'', and allied it with Younginoidea in the clade Younginiformes. More recent works that use
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
analyses usually suggest that neither Younginoidea nor Younginiformes are
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
. Constanze Bickelmann, Johannes Müller and Robert R. Reisz (2009) redescribed ''Acerosodontosaurus'' and suggested an aquatic lifestyle for it. Their analysis is figured below, and it found support for two distinct families within "Younginiformes": the aquatic Tangasauridae, and the terrestrial Younginidae (in partial
polytomy An internal node of a phylogenetic tree is described as a polytomy or multifurcation if (i) it is in a rooted tree and is linked to three or more child subtrees or (ii) it is in an unrooted tree and is attached to four or more branches. A tr ...
with Tangasauridae). More resolved results were obtained by Robert R. Reisz, Sean P. Modesto and Diane M. Scott (2011) in their description of ''Orovenator''. However, those results required the exclusion of the fragmentary taxa ''Galesphyrus'', ''Kenyasaurus'', ''Palaeagama'' and ''Saurosternon'' from their analysis. The
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 ...
below shows ''Tangasaurus'' phylogenetic position among other neodiapsids following Reisz ''et al.'', 2011.


Etymology

''Tangasaurus'' was first described and named by
Sidney H. Haughton Sidney Henry Haughton FRS (7 May 1888 – 24 May 1982) was an English-born South African paleontologist and geologist best known for his description of the sauropodomorph dinosaur '' Melanorosaurus'' in 1924, and his work on the geology of ...
in
1924 Events January * January 12 – Gopinath Saha shoots Ernest Day, whom he has mistaken for Sir Charles Tegart, the police commissioner of Calcutta, and is arrested soon after. * January 20– 30 – Kuomintang in China hol ...
and the
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...
is ''Tangasaurus mennelli''. The generic name is derived from the name of the
Tanga Region Tanga Region (''Mkoa wa Tanga'' in Swahili) is one of Tanzania's 31 administrative regions. The region covers an area of . The region is comparable in size to the combined land area of the nation state of Burundi. The regional capital is the mun ...
in which the only known specimens were found, and
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
''sauros'', meaning "lizard". The specific name honors F. P. Mennell, the geologist who reported and collected the specimens of ''Tangasaurus''.


References

{{Taxonbar, from=Q2908535 Prehistoric neodiapsids Prehistoric reptile genera Fossil taxa described in 1924 Permian reptiles of Africa Taxa named by Sidney H. Haughton