Notosuchia is a suborder of primarily Gondwanan mesoeucrocodylian crocodylomorphs that lived during the Jurassic and Cretaceous. Some phylogenies recover Sebecosuchia as a clade within Notosuchia, others as a sister group (see below); if Sebecosuchia is included within Notosuchia its existence is pushed into the Middle Miocene, about 11 million years ago. 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.[2]

Notosuchia
Temporal range: Middle JurassicLate Cretaceous, 167–66 Ma[1] Middle Miocene (11 Ma) including Sebecosuchia
August 1, 2012 - Simosuchus on Display at the Royal Ontario Museum.jpg
Mounted skeleton of the notosuchian Simosuchus clarki in the Royal Ontario Museum
Scientific classification e
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Clade: Mesoeucrocodylia
Clade: Metasuchia
Clade: Ziphosuchia
Suborder: Notosuchia
Gasparini, 1971
Families and genera

DescriptionEdit

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.

ClassificationEdit

TaxonomyEdit

GeneraEdit

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

Six 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

 

Razanandrongobe

Middle Jurassic (Bathonian)

  Madagascar

Sakaraha Formation

The earliest known member of the group.[1]

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

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

PhylogenyEdit

Cladograms of Notosuchia
Ortega et al., 2000[3]
Notosuchia 

Notosuchus

Libycosuchus

Baurusuchus

Iberosuchus

Sebecus

Itaborai crocodile

Bretesuchus

Pol, 2003[4]
Notosuchia 

Uruguaysuchus

Simosuchus

Malawisuchus

Candidodon

Notosuchus

Comahuesuchus

Chimaerasuchus

Sphagesaurus

 Sebecosuchia 

Baurusuchus

Bretesuchus

Iberosuchus

Larsson and Sues, 2007[5]
Metasuchia 
 Notosuchia

Notosuchus

Malawisuchus

Araripesuchus

Baurusuchus

Neosuchia

 Sebecia 

Pabwehshi

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.[3] 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.[4] More recently, a phylogenetic analysis by Larsson and Sues (2007) resulted in the naming of a new clade, Sebecia, to include sebecids and peirosaurids.[5] 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 as of 2014, 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.[6]

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

This cladogram represents the results of the most comprehensive analysis of notosuchian relationships to date, performed in the description of Antaeusuchus taouzensis by Nicholl et al. 2021. It is largely based on the matrix from the above Pol et al. 2014 study, but also adding character scores from Leardi et al. 2015, Fiorelli et al. 2016, Leardi et al. 2018, and Martinez et al. 2018. The final matrix consisted of 121 taxa scored for 443 morphological traits.[7]

Notosuchia
Uruguaysuchidae

Araripesuchus tsangatsangana

Anatosuchus

Araripesuchus wegeneri

Araripesuchus buitreraensis

Araripesuchus gomesii

Araripesuchus patagonicus

Uruguaysuchus

Mahajangasuchidae

Kaprosuchus

Mahajangasuchus

Peirosauridae

Stolokrosuchus

Bayomesasuchus

Hamadasuchus

Antaeusuchus

Miadanasuchus

Barrosasuchus

Gasparinisuchus

Rukwasuchus

Uberabasuchus

Lomasuchus

Montealtosuchus

Candidodon

Ziphosuchia

Libycosuchus

Simosuchus

Neuquensuchus

Pakasuchus

Malawisuchus

Xenodontosuchia
Sphagesauria

Morrinhosuchus

Llanosuchus

Notosuchus

Labidiosuchus

Mariliasuchus

Sphagesauridae

Adamantinasuchus

Yacarerani

Caipirasuchus

Sphagesaurus

Caryonosuchus

Armadillosuchus

Chimaerasuchus

Comahuesuchus

Razanandrongobe

Sebecosuchia
Baurusuchidae

Cynodontosuchus

Pissarrachampsa

Campinasuchus

Stratiosuchus

Baurusuchus

Bergisuchus

Iberosuchus

Sebecidae

Ayllusuchus

Bretesuchus

Barinasuchus

Ogresuchus

Sebecus

ReferencesEdit

  1. ^ a b Dal Sasso C, Pasini G, Fleury G, Maganuco S. (2017) Razanandrongobe sakalavae, a gigantic mesoeucrocodylian from the Middle Jurassic of Madagascar, is the oldest known notosuchian. PeerJ 5:e3481 https://doi.org/10.7717/peerj.3481
  2. ^ Gasparini, Z. (1971). "Los Notosuchia del Cretácico de América del Sur como un nuevo Infraorden de los Mesosuchia (Crocodilia)". Ameghiniana. 8: 83–103.
  3. ^ a b 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.
  4. ^ a b 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. S2CID 86592576.
  5. ^ a b 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.
  6. ^ 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. Bibcode:2014PLoSO...993105P. doi:10.1371/journal.pone.0093105. PMC 3973723. PMID 24695105.
  7. ^ Nicholl CS, Hunt ES, Ouarhache D, Mannion PD (2021). "A second peirosaurid crocodyliform from the Mid-Cretaceous Kem Kem Group of Morocco and the diversity of Gondwanan notosuchians outside South America". Royal Society Open Science. 8 (10): Article ID 211254. doi:10.1098/rsos.211254.