Notosuchia

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Notosuchia
Temporal range: Middle JurassicMiddle Miocene, 167–11  Ma [1]
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 OOjs UI icon edit-ltr.svg
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Clade: Pseudosuchia
Clade: Crocodylomorpha
Clade: Mesoeucrocodylia
Clade: Metasuchia
Clade: Ziphosuchia
Suborder: Notosuchia
Gasparini, 1971
Families and genera

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]

Contents

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.

Recent research found Araripesuchus wegeneri , Armadillosuchus arrudai , Baurusuchus , Iberosuchus macrodon , and Stratiotosuchus maxhechti were ectothermic organisms [3]

Classification

Taxonomy

Genera

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

GenusAgeLocationUnitNotesImages
Adamantinasuchus TuronianSantonian Flag of Brazil.svg  Brazil Adamantina Formation A carnivore with a very short, high skull and large eye sockets Adamantinasuchus BW.jpg
Anatosuchus AptianAlbian Flag of Niger.svg  Niger Tegama Group A small notosuchian under 1 metre (3.3 ft) long with a duck-like snout Anatosuchus.jpg
Araripesuchus AlbianMaastrichtian Six species are known, the most of any notosuchian Araripesuchus wegeneri.jpg
Armadillosuchus TuronianSantonian Flag of Brazil.svg  Brazil Adamantina Formation A sphagesaurid with armadillo-like armor shields. Armadillosuchus.jpg
Baurusuchus Turonian Flag of Brazil.svg  Brazil Adamantina Formation A large hypercarnivore 3.5 to 4 metres (11 to 13 ft) in length Baurusuchus BW.jpg
Caipirasuchus TuronianSantonian Flag of Brazil.svg  Brazil Adamantina Formation
Campinasuchus TuronianSantonian Flag of Brazil.svg  Brazil Adamantina Formation
Candidodon Albian Flag of Brazil.svg  Brazil Itapecuru Formation
Chimaerasuchus AptianAlbian Flag of the People's Republic of China.svg  China Wulong Formation The first notosuchian found with heterodont teeth, thought to be a herbivore
Comahuesuchus Santonian Flag of Argentina.svg  Argentina Bajo de la Carpa Formation Comahuesuchus BW.jpg

Cynodontosuchus

ConiacianSantonian Flag of Argentina.svg  Argentina
Libycosuchus Cenomanian
Bahariya Formation
Malawisuchus Early Cretaceous Flag of Malawi.svg  Malawi A possible burrower that could move its jaw back and forth while eating
Mariliasuchus CampanianMaastrichtian Flag of Brazil.svg  Brazil Adamantina Formation Mariliasuchus BW.jpg
Morrinhosuchus TuronianSantonian Flag of Brazil.svg  Brazil Adamantina Formation Morrinhosuchus.jpg
Notosuchus ConiacianSantonian Flag of Argentina.svg  Argentina Bajo de la Carpa Formation A notosuchian that may have had a pig-like snout Notosuchus BW.jpg
Pakasuchus Albian Flag of Tanzania.svg  Tanzania A notosuchian with very complex, mammal-like heterodont teeth. Pakasuchus.jpg
Pissarrachampsa CampanianMaastrichtian Flag of Brazil.svg  Brazil Vale do Rio do Peixe Formation Pissarrachampsa sera.png
Razanandrongobe Middle Jurassic (Bathonian)Flag of Madagascar.svg  Madagascar Sakaraha Formation The earliest known member of the group. [1]
Simosuchus Maastrichtian Flag of Madagascar.svg  Madagascar A broad-snouted omnivore with clove-shaped teeth Simosuchus clarki skull.jpg
Sphagesaurus Late Cretaceous Flag of Brazil.svg  Brazil Adamantina Formation An omnivorous notosuchian
Stratiotosuchus TuronianSantonian Flag of Brazil.svg  Brazil Adamantina Formation Stratiotosuchus maxhechti.jpg
Uruguaysuchus SantonianCampanian Flag of Uruguay.svg  Uruguay Uruguaysuchus BW.jpg
Wargosuchus Santonian Flag of Argentina.svg  Argentina Bajo de la Carpa Formation
Yacarerani Turonian-Santonian Bandera de Bolivia (Estado).svg  Bolivia Cajones Formation A notosuchian with rabbit-like incisors found in association with a probable nest Yacarerani.jpg

Phylogeny

Cladograms of Notosuchia
Ortega et al., 2000 [4]
Notosuchia 

Notosuchus

Libycosuchus

Baurusuchus

Iberosuchus

Sebecus

Itaborai crocodile

Bretesuchus

Pol, 2003 [5]
Notosuchia 

Uruguaysuchus

Simosuchus

Malawisuchus

Candidodon

Notosuchus

Comahuesuchus

Chimaerasuchus

Sphagesaurus

 Sebecosuchia 

Baurusuchus

Bretesuchus

Iberosuchus

Larsson and Sues, 2007 [6]
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. [4] 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. [5] More recently, a phylogenetic analysis by Larsson and Sues (2007) resulted in the naming of a new clade, Sebecia, to include sebecids and peirosaurids. [6] 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. [7]

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. [8]

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

Related Research Articles

<i>Simosuchus</i> Extinct genus of reptiles

Simosuchus is an extinct genus of notosuchian crocodylomorphs from the Late Cretaceous of Madagascar. It is named for its unusually short skull. Fully grown individuals were about 0.75 metres (2.5 ft) in length. The type species is Simosuchus clarki, found from the Maevarano Formation in Mahajanga Province, although some fossils have been found in India.

<span class="mw-page-title-main">Chimaerasuchidae</span> Extinct family of reptiles

Chimaerasuchidae is a family of mesoeucrocodylians. It was erected as a clade in 2004 by Carvalho et al and included Chimaerasuchus from the Early Cretaceous of China and possibly also Simosuchus from the Late Cretaceous of Madagascar. The validity of the clade has been questioned in later studies that found the two genera to be more distantly related.

<i>Razanandrongobe</i> Genus of fossil reptiles related to crocodilians

Razanandrongobe is a genus of carnivorous ziphosuchian crocodyliform from the Middle Jurassic of Madagascar. It contains the type and only species Razanandrongobe sakalavae, named in 2004 by Simone Maganuco and colleagues based on isolated bones found in 2003. The remains, which included a fragment of maxilla and teeth, originated from the Bathonian-aged Sakaraha Formation of Mahajanga, Madagascar. While they clearly belonged to a member of the Archosauria, Maganuco and colleagues refrained from assigning the genus to a specific group because the fragmentary remains resembled lineages among both the theropod dinosaurs and crocodylomorphs.

<span class="mw-page-title-main">Peirosauridae</span> Extinct family of reptiles

Peirosauridae is a Gondwanan family of mesoeucrocodylians that lived during the Cretaceous period. It was a clade of terrestrial crocodyliforms that evolved a rather dog-like skull, and were terrestrial carnivores. It was phylogenetically defined in 2004 as the most recent common ancestor of Peirosaurus and Lomasuchinae and all of its descendants. Lomasuchinae is a subfamily of peirosaurids that includes the genus Lomasuchus.

<span class="mw-page-title-main">Baurusuchidae</span> Extinct family of reptiles

Baurusuchidae is a Gondwanan family of mesoeucrocodylians that lived during the Late Cretaceous. It is a group of terrestrial hypercarnivorous crocodilians from South America and possibly Pakistan. Baurusuchidae has been, in accordance with the PhyloCode, officially defined as the least inclusive clade containing Cynodontosuchus rothi, Pissarrachampsa sera, and Baurusuchus pachecoi. Baurusuchids have been placed in the suborder Baurusuchia, and two subfamilies have been proposed: Baurusuchinae and Pissarrachampsinae.

<i>Araripesuchus</i> Extinct genus of reptiles

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 notosuchians, characterized by their varied teeth types and distinct skull elements. This genus consists of six species: A. buitreraensis, discovered in Argentina, A. wegeneri, discovered in Cameroon and Niger, A. rattoides, discovered in Niger, A. tsangatsangana, discovered in Madagascar, A. gomesii, discovered in Brazil and another species discovered in Argentina, A. patagonicus. It has been argued that the phylogenetic position of this genus is uncertain, and that taxonomic revision is required.

Pabwehshi is an extinct genus of mesoeucrocodylian. It is based on GSP-UM 2000, a partial snout and corresponding lower jaw elements, with another snout assigned to it. These specimens were found in Maastrichtian-age Upper Cretaceous rocks of the Vitakri and Pab Formations in Balochistan, Pakistan, and represent the first diagnostic crocodyliform fossils from Cretaceous rocks of South Asia. Pabwehshi had serrated interlocking teeth in its snout that formed a "zig-zag" cutting edge. Pabwehshi was named in 2001 by Jeffrey A. Wilson and colleagues. The type species is P. pakistanensis, in reference to the nation where it was found. It was traditionally classified as a baurusuchid closely related to Cynodontosuchus and Baurusuchus. Larsson and Sues (2007) found close affinity between Pabwehshi and the Peirosauridae within Sebecia. Montefeltro et al.Pabwehshi has a sagittal torus on its maxillary palatal shelves – a character that is absent in baurusuchids – but they did not include Pabwehshi in their phylogenetic analysis.

Trematochampsidae is an extinct family of mesoeucrocodylian crocodylomorphs. Fossils are present from Madagascar, Morocco, Niger, Argentina, and Brazil. Possible trematochampsids have been found from Spain and France, but classification past the family level is indeterminant. The trematochampsids first appeared during the Barremian stage of the Early Cretaceous and became extinct during the late Maastrichtian stage of the Late Cretaceous.

<i>Bergisuchus</i> Extinct genus of reptiles

Bergisuchus is an extinct genus of small sebecosuchian mesoeucrocodylian known primarily from the Eocene Messel Pit in Germany. Few fossils of Bergisuchus have been discovered, only a single incomplete snout, a few partial lower jaws and some teeth. Despite being fragmentary, the jaw bones are enough to indicate that Bergisuchus had a short, deep, narrow snout and serrated teeth, quite unlike the broad flat snouts of modern crocodylians.

Ilchunaia is an extinct genus of sebecid mesoeucrocodylian. Fossils have been found from the Divisadero Largo Formation of Argentina dating back to the Late Eocene, and a locality in Mendoza, Argentina dating back to the Oligocene. Little material is known from the genus, with only the anterior portion of the skull being present to study.

<i>Kaprosuchus</i> Genus of crocodyliform from the Late Cretaceous period

Kaprosuchus is an extinct genus of mahajangasuchid crocodyliform. It is known from a single nearly complete skull collected from the Upper Cretaceous Echkar Formation of Niger. The name means "boar crocodile" from the Greek κάπρος, kapros ("boar") and σοῦχος, soukhos ("crocodile") in reference to its unusually large caniniform teeth which resemble those of a boar. It has been nicknamed "BoarCroc" by Paul Sereno and Hans Larsson, who first described the genus in a monograph published in ZooKeys in 2009 along with other Saharan crocodyliformes such as Anatosuchus and Laganosuchus. The type species is K. saharicus.

Mahajangasuchidae is an extinct family of notosuchian crocodyliforms. It currently contains two genera, Mahajangasuchus and Kaprosuchus, both of which lived during the Late Cretaceous in Gondwana. It is defined as the most inclusive clade containing Mahajangasuchus insignis but not Notosuchus terrestris, Simosuchus clarki, Araripesuchus gomesii, Baurusuchus pachecoi, Peirosaurus torminni, Goniopholis crassidens, Pholidosaurus schaumbergensis, or Crocodylus niloticus. Phylogenetically, Mahajangasuchidae is placed just outside pholidosaurids and more derived neosuchians.

<span class="mw-page-title-main">Metasuchia</span> Clade of reptiles

Metasuchia is a major clade within the superorder Crocodylomorpha. It is split into two main groups, Ziphosuchia and Neosuchia. Notosuchia is an extinct group that contains primarily small-bodied Cretaceous taxa with heterodont dentition. Neosuchia includes the extant crocodylians and basal taxa, such as peirosaurids and pholidosaurids. It is phylogenetically defined by Sereno et al. (2001) as a clade containing Notosuchus terrestris, Crocodylus niloticus, and all descendants of their common ancestor.

<i>Sebecus</i> Extinct genus of reptiles

Sebecus is an extinct genus of sebecid crocodylomorph from Eocene of South America. Like other sebecosuchians, it was entirely terrestrial and carnivorous. The genus is currently represented by two species, the type S. icaeorhinus and S. ayrampu. Several other species have been referred to Sebecus, but were later reclassified as their own genera.

<span class="mw-page-title-main">Sebecia</span> Extinct clade of reptiles

Sebecia is an extinct clade of mesoeucrocodylian crocodyliforms that includes peirosaurids and sebecids. It was first constructed in 2007 to include Hamadasuchus, Peirosauridae, and Sebecus. It was initially considered to be the sister taxon of the clade Neosuchia, which includes living crocodilians, although some later studies have placed it within Neosuchia as a basal clade. Sebecians were terrestrial crocodyliforms characterized by their deep snouts and ziphodont dentition. They first appeared in the Late Cretaceous, survived the Cretaceous–Paleogene extinction event, and became extinct in the Miocene epoch.

Sebecosuchia is an extinct group of mesoeucrocodylian crocodyliforms that includes the families Sebecidae and Baurusuchidae. The group was long thought to have first appeared in the Late Cretaceous with the baurusuchids and become extinct in the Miocene with the last sebecids, but Razanandrongobe pushes the origin of Sebecosuchia to the Middle Jurassic. Fossils have been found primarily from South America but have also been found in Europe, North Africa, Madagascar, and the Indian subcontinent.

<i>Stratiotosuchus</i> Prehistoric genus of reptiles

Stratiotosuchus is an extinct genus of baurusuchid mesoeucrocodylian from the Adamantina Formation in Brazil. It lived during the Late Cretaceous. The first fossils were found in the 1980s, and the type species Stratiotosuchus maxhechti was named in 2001. A hyperpredator, it and other baurusuchids may have filled niches occupied elsewhere by theropod dinosaurs.

<i>Pissarrachampsa</i> Extinct genus of reptiles

Pissarrachampsa is an extinct genus of baurusuchid mesoeucrocodylian from the Late Cretaceous of Brazil. It is based on a nearly complete skull and a referred partial skull and lower jaw from the ?Campanian - ?Maastrichtian-age Vale do Rio do Peixe Formation of the Bauru Group, found in the vicinity of Gurinhatã, Brazil.

<i>Antaeusuchus</i> Genus of extinct animals

Antaeusuchus taouzensis is a species of peirosaurid notosuchian from the Late Cretaceous Kem Kem Group of Morocco. It was described in 2021, and it is the only species in the genus Antaeusuchus. It is the fourth notosuchian described from the region and the second Kem Kem peirosaurid after Hamadasuchus. A 2023 study proposes that Antaeusuchus may not be distinct enough to warrant its own genus and that it instead represents another species of Hamadasuchus.

Titanochampsa is a genus of large mesoeucrocodylian from the Maastrichtian Marilia Formation of Brazil. Although only known from a single skull roof, the material shows that Titanochampsa was not a member of Notosuchia, which were previously believed to have been the only crocodyliforms present in the strata of the Bauru Group. Body size estimates vary greatly and range between 2.98–5.88 m due to the incomplete nature of the holotype fossil. The overall anatomy of the skull roof, alongside its size and possible affinities with Neosuchians, may suggest that it was a semi-aquatic ambush hunter similar to modern crocodilians.

References

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  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. Cubo, J.; Aubier, P.; Faure-Brac, M. G.; Martet, G.; Pellarin, R.; Pelletan, I.; Sena, M. V. A. (2022). "Paleohistological inferences of thermometabolic regimes inNotosuchia (Pseudosuchia: Crocodylomorpha) revisited". Paleobiology: 1–11. doi:10.1017/pab.2022.28.
  4. 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.
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  6. 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 .
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  8. 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 . PMC   8511751 . PMID   34659786.