Sebecosuchia
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Sebecosuchia (meaning "
Sobek Sobek (), also known as Suchus (), was an ancient Egyptian deities, ancient Egyptian deity with a complex and elastic history and nature. He is associated with the Nile crocodile and is often represented as a crocodile-headed humanoid, if not a ...
crocodiles") is an extinct group of mesoeucrocodylian
crocodyliform Crocodyliformes is a clade of Crurotarsi, crurotarsan archosaurs, the group often traditionally referred to as "crocodilians". They are the first members of Crocodylomorpha to possess many of the features that define later relatives. They are the ...
s that includes the families Sebecidae and Baurusuchidae. The group was long thought to have first appeared in 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 ...
with the baurusuchids, but '' Razanandrongobe'' pushes the origin of Sebecosuchia to the Middle Jurassic. The last surviving members of the group, the sebecids, appear to have lasted until the
late Miocene The Late Miocene (also known as Upper Miocene) is a sub-epoch of the Miocene epoch (geology), Epoch made up of two faunal stage, stages. The Tortonian and Messinian stages comprise the Late Miocene sub-epoch, which lasted from 11.63 Ma (million ye ...
or
early Pliocene Early may refer to: Places in the United States * Early, Iowa, a city * Early, Texas, a city * Early Branch, a stream in Missouri * Early County, Georgia * Fort Early, Georgia, an early 19th century fort Music * Early B, stage name of Jamaican d ...
on the
Greater Antilles The Greater Antilles is a grouping of the larger islands in the Caribbean Sea, including Cuba, Hispaniola, Puerto Rico, and Jamaica, together with Navassa Island and the Cayman Islands. Seven island states share the region of the Greater Antille ...
. Fossils have been found primarily from
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 ...
but have also been found in
Europe Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...
,
North Africa North Africa (sometimes Northern Africa) is a region encompassing the northern portion of the African continent. There is no singularly accepted scope for the region. However, it is sometimes defined as stretching from the Atlantic shores of t ...
,
Madagascar Madagascar, officially the Republic of Madagascar, is an island country that includes the island of Madagascar and numerous smaller peripheral islands. Lying off the southeastern coast of Africa, it is the world's List of islands by area, f ...
, and the Indian subcontinent.


History and phylogeny

Sebecosuchia was first constructed in 1937 by
George Gaylord Simpson George Gaylord Simpson (June 16, 1902 – October 6, 1984) was an American paleontologist. Simpson was perhaps the most influential paleontologist of the twentieth century, and a major participant in the modern synthesis, contributing '' Tempo ...
. In 1946 the concept was again used by American paleontologist Edwin Colbert to include '' Sebecus'' and Baurusuchidae. ''Sebecus'', which had been known from
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 ...
since 1937, was an unusual crocodyliform with a deep snout and teeth that were ziphodont, or serrated and laterally compressed. The family Baurusuchidae was named the year before and included the newly described ''Baurusuchus'', which was also a South American deep-snouted form. The clade Sebecosuchia was given a phylogenetic definition in the ''
PhyloCode The ''International Code of Phylogenetic Nomenclature'', known as the ''PhyloCode'' for short, is a formal set of rules governing phylogenetic nomenclature. Its current version is specifically designed to regulate the naming of clades, leaving the ...
'' by Juan Leardi and colleagues in 2024 as "the least inclusive clade containing '' Sebecus icaeorhinus'' and '' Baurusuchus pachecoi'' provided that it doesn’t include '' Araripesuchus gomesii'', '' Montealtosuchus arrudacamposi'', or ''
Crocodylus niloticus The Nile crocodile (''Crocodylus niloticus'') is a large crocodilian native to freshwater habitats in Africa, where it is present in 26 countries. It is widely distributed in sub-Saharan Africa, occurring mostly in the eastern, southern, and ce ...
'' (the Nile crocodile)". This definition ensures that Sebecosuchia self destructs if peirosaurids, uruguaysuchids, or modern crocodylians fall within the last common ancestor and all descendants of baurusuchids and sebecids. More recently, other crocodyliforms have been assigned to Sebecosuchia that cannot be placed into either family. These include the genera '' Eremosuchus'', named in 1989, and '' Pehuenchesuchus'', named in 2005. They are usually considered to be more basal sebecosuchians than the sebecids and baurusuchids. 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 ...
showing the possible phylogenetic position of Sebecosuchia modified from Turner and Calvo (2005). In this cladogram, Sebecidae is a
paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
assemblage of basal sebecosuchians while Baurusuchidae is monophyletic and includes the more derived sebecosuchians. In a phylogenetic study of crocodyliforms, Benton and Clark (1988) split up Sebecosuchia, finding baurusuchids to be basal
notosuchia Notosuchia is a clade of primarily Gondwanan mesoeucrocodylian Crocodylomorpha, crocodylomorphs that lived during the Jurassic and Cretaceous. Some phylogeny, phylogenies recover Sebecosuchia as a clade within Notosuchia, others as a sister group ...
ns while sebecids were basal
neosuchia Neosuchia is a clade within Mesoeucrocodylia that includes all modern extant crocodilians and their closest fossil relatives. It is defined as the most inclusive clade containing all Crocodylomorpha, crocodylomorphs more closely related to ''Croc ...
ns. Since that time, most studies have supported a monophyletic Sebecosuchia. In 2007, however, a phylogenetic study placed baurusuchids as basal metasuchians and sebecids as close relatives to a family of notosuchians called the Peirosauridae. Together, sebecids and peirosaurids made the new clade Sebecia. Below is a cladogram from that study, Larsson and Sues (2007): Two years later, Sereno and Larsson (2009) came to the same conclusion, except they placed baurusuchids as advanced notosuchians. More recently however, Turner and Sertich (2010) found support for Sebecosuchia in their analysis of notosuchian relationships. In their study, Sebecosuchia was a derived clade within Notosuchia. Iori and Carvalho (2011) came to similar conclusions, grouping ''Baurusuchus'' alongside Sebecidae. Below is the cladogram from Turner and Sertich (2010): Diego Pol and Jaime E. Powell (2011) came to the same conclusion, however their analysis couldn't find a
monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...
Baurusuchidae within Sebecosuchia. The following cladogram simplified after their analysis, with focus on Sebecosuchia.


Paleobiology

All sebecosuchians were carnivorous and terrestrial. The nares open at the very tip of the snout, suggesting that it lived on land rather than in water (in aquatic crocodyliforms, the nares usually open dorsally on top of the snout). The snout itself is laterally compressed, a feature shared with other terrestrial reptiles such as
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 ...
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 ...
s. The eye sockets are opened laterally rather than dorsally as in aquatic crocodyliforms. Moreover, there is a prominent
fourth trochanter The fourth trochanter is a shared characteristic common to archosaurs. It is a protrusion on the posterior-medial side of the middle of the femur shaft that serves as a muscle attachment, mainly for the '' musculus caudofemoralis longus'', the m ...
on the
femur The femur (; : femurs or femora ), or thigh bone is the only long bone, bone in the thigh — the region of the lower limb between the hip and the knee. In many quadrupeds, four-legged animals the femur is the upper bone of the hindleg. The Femo ...
for the attachment of muscles that would have aided in upright walking. Although they are now widely accepted to be terrestrial, sebecosuchians were once thought to be semiaquatic, spending part of their time in water. The laterally compressed snout of sebecosuchians may have enabled them to withstand high forces during biting. The teeth are also laterally compressed, pointed, and serrated. Their shape would have allowed them to easily penetrate and slice flesh. The
pterygoid bone The pterygoid is a paired bone forming part of the palate of many vertebrates, behind the palatine bone In anatomy, the palatine bones (; derived from the Latin ''palatum'') are two irregular bones of the facial skeleton in many animal specie ...
in the skull is strongly bent, allowing for a larger jaw adductor muscle to quickly close the jaws and give sebecosuchians a powerful bite.


References

{{Taxonbar, from=Q7442744 Notosuchia Terrestrial crocodylomorphs Late Cretaceous crocodylomorphs Paleogene crocodylomorphs Neogene crocodylomorphs Late Cretaceous first appearances Miocene extinctions