Notosuchia

Notosuchia
Temporal range: Early CretaceousMiddle Miocene, 110–11 Ma
Mounted skeleton of the notosuchian Simosuchus clarki in the Royal Ontario Museum
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Superorder: Crocodylomorpha
Clade: Metasuchia
Suborder: Notosuchia
Gasparini, 1971
Families

Notosuchia is a suborder of primarily Gondwanan mesoeucrocodylian crocodylomorphs that lived during the Cretaceous. Fossils have been found from South America, Africa, Asia, and Europe. Notosuchia was a clade of terrestrial crocodilians that evolved a range of feeding behaviours, including herbivory (Chimaerasuchus), omnivory (Simosuchus), and terrestrial hypercarnivory (Baurusuchus). It included many members with highly derived traits unusual for crocodylomorphs, including mammal-like teeth, flexible bands of shield-like body armor similar to those of armadillos (Armadillosuchus), and possibly fleshy cheeks and pig-like snouts (Notosuchus). The suborder was first named in 1971 by Zulma Gasparini and has since undergone many phylogenetic revisions.[1]

Description

Notosuchians were generally small, with slender bodies and erect limbs. The most distinctive characteristics are usually seen in the skull. Notosuchian skulls are generally short and deep. While most are relatively narrow, some are very broad. Simosuchus has a broadened skull and jaw that resembles a pug, while Anatosuchus has a broad, flat snout like that of a duck.

The teeth vary greatly between different genera. Many have heterodont dentitions that vary in shape across the jaw. Often, there are large canine-like teeth protruding from the front of the mouth and broader molar-like teeth in the back. Some genera, such as Yacarerani and Pakasuchus, have extremely mammal-like teeth. Their molars are complex and multicuspid, and are able to occlude or fit with one another. Some forms such as Malawisuchus had jaw joints that enabled them to move the jaw back and forth in a shearing motion rather than just up and down.

A derived group of notosuchians, the baurusuchids, differ considerably from other forms. They are very large in comparison to other notosuchians and are exclusively carnivorous. Baurusuchids have deep skulls and prominent canine-like teeth.

Classification

Taxonomy

Genera

The evolutionary interrelationships of Notosuchia are in flux, but the following genera are generally considered notosuchians:

Genus Age Location Unit Notes Images

Adamantinasuchus

Turonian - Santonian

 Brazil

Adamantina Formation

A carnivore with a very short, high skull and large eye sockets

Anatosuchus

Aptian - Albian

 Niger

Tegama Group

A small notosuchian under 1 metre (3.3 ft) long with a duck-like snout

Araripesuchus

Albian - Maastrichtian

 Madagascar
 Niger
 Brazil
 Argentina

Maevarano Formation
Echkar Formation
Santana Formation
Candeleros Formation

Five species are known, the most of any notosuchian

Armadillosuchus

Turonian - Santonian

 Brazil

Adamantina Formation

A sphagesaurid with armadillo-like armor shields.

Baurusuchus

Turonian

 Brazil

Adamantina Formation

A large hypercarnivore 3.5 to 4 metres (11 to 13 ft) in length

Caipirasuchus

Turonian – Santonian

 Brazil

Adamantina Formation

Campinasuchus

Turonian – Santonian

 Brazil

Adamantina Formation

Candidodon

Albian

 Brazil

Itapecuru Formation

Chimaerasuchus

Aptian - Albian

 China

Wulong Formation

The first notosuchian found with heterodont teeth, thought to be a herbivore

Comahuesuchus

Santonian

 Argentina

Bajo de la Carpa Formation

Cynodontosuchus

Coniacian – Santonian

 Argentina

Bajo de la Carpa Formation
Pichi Picun Leufu Formation

Libycosuchus

Cenomanian

 Egypt
 Nigeria

Bahariya formation
Malawisuchus Early Cretaceous

 Malawi

A possible burrower that could move its jaw back and forth while eating

Mariliasuchus

Campanian - Maastrichtian

 Brazil

Adamantina Formation

Morrinhosuchus

Turonian - Santonian

 Brazil

Adamantina Formation

Notosuchus

Coniacian - Santonian

 Argentina

Bajo de la Carpa Formation

A notosuchian that may have had a pig-like snout

Pakasuchus

Albian

 Tanzania

A notosuchian with very complex, mammal-like heterodont teeth.

Pissarrachampsa

Campanian – Maastrichtian

 Brazil

Vale do Rio do Peixe Formation

Simosuchus

Maastrichtian

 Madagascar

A broad-snouted omnivore with clove-shaped teeth

Sphagesaurus

Late Cretaceous

 Brazil

Adamantina Formation

An omnivorous notosuchian

Stratiotosuchus

Turonian – Santonian

 Brazil

Adamantina Formation

Uruguaysuchus

Santonian - Campanian

 Uruguay

Wargosuchus

Santonian

 Argentina

Bajo de la Carpa Formation

Yacarerani

Turonian - Santonian

 Bolivia

A notosuchian with rabbit-like incisors found in association with a probable nest

Phylogeny

Cladograms of Notosuchia
Ortega et al., 2000[2]
Notosuchia 

Notosuchus




Libycosuchus




Baurusuchus




Iberosuchus




Sebecus




Itaborai crocodile



Bretesuchus








Pol, 2003[3]
Notosuchia 

Uruguaysuchus




Simosuchus





Malawisuchus



Candidodon





Notosuchus




Comahuesuchus





Chimaerasuchus



Sphagesaurus



 Sebecosuchia 

Baurusuchus




Bretesuchus



Iberosuchus










Larsson and Sues, 2007[4]
Metasuchia 
 Notosuchia

Notosuchus



Malawisuchus





Araripesuchus





Baurusuchus




Neosuchia


 Sebecia 

Pabwehshi



 Sebecidae 

Sebecus



Bretesuchus




Peirosauridae








       Taxa previously assigned to Notosuchia


The clade Notosuchia has undergone many recent phylogenetic revisions. In 2000, Notosuchia was proposed to be one of two groups within the clade Ziphosuchia, the other being Sebecosuchia, which included deep snouted forms such as baurusuchids and sebecids.[2] The definition of Notosuchia by Sereno et al. (2001) is similar to that of Ziphosuchia as it includes within it Sebecosuchia. Pol (2003) also includes Sebecosuchia within Notosuchia.[3] More recently, a phylogenetic analysis by Larsson and Sues (2007) resulted in the naming of a new clade, Sebecia, to include sebecids and peirosaurids.[4] Baurusuchidae was considered to be polyphyletic in this study, with Pabwehshi being a basal member of Sebecia and Baurusuchus being the sister taxon to the clade containing Neosuchia and Sebecia. Thus, Sebecosuchia was no longer within Notosuchia and not considered to be a true clade, while Notosuchia was found to be a basal clade of Metasuchia.

The following cladogram simplified after the most comprehensive analysis of notosuchians to date, presented by Pol et al. in 2014. It is based mainly on the data matrix published by Pol et al. (2012) which is itself a modified version of previous analyses. Thirty-one additional characters were added from other comprehensive analyses of notosuchians, e.g. Turner and Sertich (2010), Andrade et al. (2011), Montefeltro et al. (2011), Larsson and Sues (2007), and Novas et al. (2009), and 34 characters were noval, resulting in a matrix that includes 109 crocodyliforms and outgroup taxa which are scored based on 412 morphological traits.[5]

Notosuchia


Uruguaysuchidae

Araripesuchus tsangatsangana




Anatosuchus



Araripesuchus wegeneri





Araripesuchus buitreraensis




Araripesuchus gomesii




Araripesuchus patagonicus



Uruguaysuchus








Stolokrosuchus



Mahajangasuchidae

Kaprosuchus



Mahajangasuchus



Peirosauridae

Hamadasuchus




Gasparinisuchus



Lomasuchus



Montealtosuchus



Uberabasuchus









Candidodon


Ziphosuchia

Libycosuchus




Simosuchus





Malawisuchus



Pakasuchus






Morrinhosuchus




Notosuchus




Coringasuchus



Labidiosuchus



Mariliasuchus


Sphagesauridae


Adamantinasuchus



Yacarerani






Caipirasuchus stenognathus




Caipirasuchus paulistanus



Caipirasuchus montealtensis






Sphagesaurus




Armadillosuchus



Caryonosuchus











Chimaerasuchus




Comahuesuchus


Sebecosuchia

Pabwehshi


Baurusuchidae

Cynodontosuchus




Pissarrachampsa




Stratiotosuchus




Campinasuchus




Baurusuchus albertoi




Baurusuchus pachecoi



Baurusuchus salgadoensis











Bergisuchus



Iberosuchus



Sebecidae


Lorosuchus



Pehuenchesuchus





Barinasuchus





Ayllusuchus



Bretesuchus





Lumbrera form



Langstonia



Sebecus



Zulmasuchus

















References

  1. Gasparini, Z. (1971). "Los Notosuchia del Cretácico de América del Sur como un nuevo Infraorden de los Mesosuchia (Crocodilia)". Ameghiniana 8: 83–103.
  2. 1 2 Ortega, F. Z.; Buscalioni, A. D.; Calvo, J. O. (2000). "A new species of Araripesuchus (Crocodylomorpha, Mesoeucrocodylia) from the Lower Cretaceous of Patagonia (Argentina)". Journal of Vertebrate Paleontology 20 (1): 57–76. doi:10.1671/0272-4634(2000)020[0057:ANSOAC]2.0.CO;2.
  3. 1 2 Pol, D. (2003). "New Remains of Sphagesaurus huenei (Crocodylomorpha: Mesoeucrocodylia) from the Late Cretaceous of Brazil". Journal of Vertebrate Paleontology 23 (4): 817–831. doi:10.1671/A1015-7.
  4. 1 2 Larsson, H. C. E.; Sues, H.-D. (2007). "Cranial osteology and phylogenetic relationships of Hamadasuchus rebouli (Crocodyliformes: Mesoeucrocodylia) from the Cretaceous of Morocco". Zoological Journal of the Linnean Society 149 (4): 533–567. doi:10.1111/j.1096-3642.2007.00271.x.
  5. Pol, D.; Nascimento, P. M.; Carvalho, A. B.; Riccomini, C.; Pires-Domingues, R. A.; Zaher, H. (2014). "A New Notosuchian from the Late Cretaceous of Brazil and the Phylogeny of Advanced Notosuchians". PLoS ONE 9 (4): e93105. doi:10.1371/journal.pone.0093105. PMC 3973723. PMID 24695105.
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