Susisuchus

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Susisuchus
Temporal range: Early Cretaceous
Susisuchus specimen.PNG
S. anatoceps specimen (A) and cross sectioned samples (B-C)
Scientific classification Red Pencil Icon.png
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Clade: Neosuchia
Family: Susisuchidae
Salisbury et al., 2003
Genus: Susisuchus
Salisbury et al., 2003
Species
  • S. anatocepsSalisbury et al., 2003 (type)
  • S. jaguaribensisFortier and Schultz, 2009

Susisuchus is an extinct genus of neosuchian mesoeucrocodylian crocodyliform from the Early Cretaceous of Brazil. Fossils have been found from the Nova Olinda Member of the Aptian-age Crato Formation in the Araripe and Lima Campos Basins of northeastern Brazil. Named in 2003, Susisuchus is the sole member of the family Susisuchidae, and is closely related to the clade Eusuchia, which includes living crocodilians. The type species is S. anatoceps, known from a single partial articulated skeleton that preserves some soft tissue. A second species, S. jaguaribensis, was named in 2009 from fragmentary remains.

Contents

Discovery

Size of S. anatoceps and other Brazilian Cretaceous crocodylomorphs Size of Brazilian fossil crocodylomorphs.PNG
Size of S. anatoceps and other Brazilian Cretaceous crocodylomorphs

The genus Susisuchus was first erected with the description of the type species S. anatoceps in 2003. S. anatoceps was found in the Aptian-age Crato Formation in the Araripe Basin, and was the first crocodyliform to be described from the formation. The holotype skeleton is the most complete known specimen of Susisuchus, including most of the bones except for those of the hindlimbs, and even traces of soft tissue. The specimen is thought to have been a desiccated carcass before it was buried and later fossilized. A second species, S. jaguaribensis, was named in 2009 from the Lima Campos Basin about 115 kilometres (71 mi) from where the skeleton of S. anatoceps was uncovered. [1]

A new crocodyliform specimen from the Crato Formation was described in 2009. It can be distinguished from the two other crocodyliforms known from the formation, Araripesuchus and the trematochampsid Caririsuchus , on the basis of characteristics of the fibula and proportions of the femur and tibia. Since the limb elements differ from all other known crocodyliforms from the Araripe Basin, the specimen was tentatively classified as cf. Susisuchus sp. [2]

Description

The holotype specimen of Susisuchus anatoceps includes a skull and lower jaw, forelimbs, parts of the axial skeleton, and some osteoderms. There are traces of soft tissue around both forelimbs and the digits of the right hand. S. jaguaribensis is known from far less material, but the fragmentary remains are enough to diagnose the species. Both species have a contact between the squamosal and parietal bones that is situated on the posterior border of the supratemporal fenestra, a hole in the skull table. [1]

Osteoderms

Segmentation of dorsal osteoderms over time in advanced neosuchians (Susisuchus second from left). Neosuchian osteoderms.svg
Segmentation of dorsal osteoderms over time in advanced neosuchians (Susisuchus second from left).

Susisuchus has a dorsal shield of osteoderms that is similar to those of the more derived clade Eusuchia, which includes living crocodilians. Susisuchus was one of the first mesoeucrocodylians to have a tetraserial paravertebral shield, which is formed from the dorsal osteoderms to make the back more rigid. In a tetraserial paravertebral shield, there are four separate rows of paravertebral osteoderms (osteoderms near the midline of the back). Earlier mesoeucrocodylians such as Bernissartia have biserial paravertebral shields, in which there are only two rows of paravertebral osteoderms. These two rows of large osteoderms became segmented into four smaller rows in Susisuchus. [3] Susisuchus also has two rows of accessory osteoderms to either side of the paravertebral osteoderms which are not part of the shield. [4]

The segmentation of the paravertebral shield allowed for a greater degree of lateral flexibility in the trunk of Susisuchus. This flexibility would have enabled lateral undulation while swimming, resulting in greater efficiency in locomotion. Because the width of the paravertebral shield was not compromised by this segmentation, the back would remain rigid enough to overcome the forces encountered during high-walking, or walking semi-erect. However, the ability to high-walk would have restricted the size of Susisuchus. If Susisuchus grew any larger than 50 kilograms (110 lb), the forces encountered during high-walking would have been too great for the paravertebral shield to counteract. [3]

Classification

Possible resting postures of S. anatoceps at various depths Resting postures of Susisuchus.PNG
Possible resting postures of S. anatoceps at various depths

Susisuchus is closely related to Pachycheilosuchus from Glen Rose, Texas, United States, and Isisfordia from Winton, Queensland, Australia. All three genera are highly derived Early Cretaceous mesoeucrocodylians. [4] In the initial description of Susisuchidae, Isisfordia was considered a possible member of the family, although the genus had not yet been named. However, with its naming in 2006, Isisfordia was placed within Eusuchia as the most basal member of the clade, while Susisuchus was placed outside Eusuchia as its sister taxon. Below is a cladogram from Salisbury et al. (2006) showing the relationships of Susisuchus and Isisfordia within Mesoeucrocodylia: [3]

Mesoeucrocodylia' 

Theriosuchus spp.

Goniopholis spp.

Bernissartia fagesii

Susisuchus anatoceps

Eusuchia  

Isisfordia duncani

Hylaeochampsa vectiana

Allodaposuchus precedens

Crocodylia

Susisuchus has amphicoelus thoracic, lumbar and caudal (tail) vertebrae, meaning that the ends of the centra of these vertebrae are concave. The presence of this feature in a crocodyliform with a tetraserial paravertebral shield is unusual, and has been used to erect the family Susisuchidae, of which Susisuchus is the only member. [4] Susisuchus can be considered a transitional form because it has a tetraserial paravertebral shield characteristic of eusuchians, but retains more primitive amphicoelus vertebrae characteristic of more basal crocodylomorphs.

With the description of S. jaguaribensis as a second species of Susisuchus, the genus has been phylogenetically redefined as a node-based taxon including the last common ancestor of Susisuchus anatoceps and Susisuchus jaguaribensis and all of its descendants. [1]

Related Research Articles

Mesoeucrocodylia Clade of reptiles

Mesoeucrocodylia is the clade that includes Eusuchia and crocodyliforms formerly placed in the paraphyletic group Mesosuchia. The group appeared during the Early Jurassic, and continues to the present day.

<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.

Chimaerasuchidae 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>Mariliasuchus</i> Extinct genus of reptiles

Mariliasuchus is an extinct genus of Late Cretaceous notosuchian mesoeucrocodylian found near Marilia, Brazil. The first bone remains were found and collected in 1995 by Brazilian paleontologist William Nava, in red rocks from the fossiliferous Adamantina Formation. Four years later, it was described as Mariliasuchus amarali, by Brazilian paleontologists Ismar de Souza Carvalho and Reinaldo J. Bertini.

Hyposaurus is a genus of extinct marine dyrosaurid crocodyliform. Fossils have been found in Paleocene aged rocks of the Iullemmeden Basin in West Africa, Campanian–Maastrichtian Shendi Formation of Sudan and Maastrichtian through Danian strata in New Jersey, Alabama and South Carolina. Isolated teeth comparable to Hyposaurus have also been found in Thanetian strata of Virginia. It was related to Dyrosaurus. The priority of the species H. rogersii has been debated, however there is no sound basis for the recognition of more than one species from North America. The other North American species are therefore considered nomina vanum.

Pachycheilosuchus is an extinct genus of mesoeucrocodylian from the Early Cretaceous of Texas, United States. Previously known, in part, as the "Glen Rose form", this crocodylomorph is notable for its procoelous vertebrae, otherwise found only in derived eusuchian crocodilians, a thick margin on the maxillae, and a shield of armor on the neck formed by the fusion of six individual scutes.

Calsoyasuchus is a genus of goniopholidid mesoeucrocodylian that lived in the Early Jurassic. Its fossilized remains were found in the Sinemurian-Pliensbachian-age Kayenta Formation on Navajo Nation land in Coconino County, Arizona, United States. Formally described as C. valliceps, it is known from a single incomplete skull which is unusually derived for such an early crocodile relative. This genus was described in 2002 by Ronald Tykoski and colleagues; the species name means "valley head" and refers to a deep groove along the midline of the nasal bones and frontal bones.

Rugosuchus is an extinct genus of neosuchian crocodyliform from the late Early Cretaceous of China. It is known from most of a skull, a partial postcranial skeleton, and a second partial skeleton including part of the hips. It was described by Xiao-Chun Wu and colleagues in 2001, with R. nonganensis as the type species. At the time of its description, it was the most complete crocodyliform from northeastern China, and only the second known.

Eusuchia Clade of reptiles

The Eusuchia are a clade of crocodylomorphs that first appeared in the Early Cretaceous with Hylaeochampsa. Along with Dyrosauridae and Sebecosuchia, they were the only crocodyliformes who survived the K-T extinction. Since the other two clades died out 35 and 11 million years ago respectively, all living crocodilian species are eusuchians, as are many extinct forms.

<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 other five related species: A. wegeneri and A. rattoides, discovered in Niger, A. tsangatsangana, discovered in Madagascar, A. gomesii, discovered in Brazil and another species discovered in Argentina, A. patagonicus.

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.

Eutretauranosuchus is an extinct genus of goniopholidid crocodyliform. E. delfsi is the only known species within the genus.

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 crocodylomorphs more closely related to Crocodylus niloticus than to Notosuchus terrestris. Neosuchia is very diverse and may be polyphyletic, as the clade has undergone many revisions since it was first named in 1988. Members of Neosuchia generally share a crocodilian-like bodyform adapted to freshwater aquatic life, as opposed to the terrestrial habits of more basal crocodylomorph groups. The earliest neosuchian is suggested to be the Early Jurassic Calsoyasuchus, which lived during the Sinemurian and Pliensbachian stages, often identified as a member of Goniopholididae, though this is disputed, and the taxon may lie outside Neosuchia, which places the earliest records of the group in the Middle Jurassic.

Gilchristosuchus is an extinct genus of neosuchian crocodyliform. Its fossils have been found in the upper Milk River Formation of Alberta, Canada, in rocks of either latest Santonian or earliest Campanian age. Gilchristosuchus was described in 1993 by Wu and Brinkman. The type species is G. palatinus, in reference to its distinctive palatine bones.

Brillanceausuchus is an extinct genus of atoposaurid crocodylomorph. Fossils have been found in Early Cretaceous–age rocks of Cameroon. The genus is notable for the position of the secondary choana within its palate. Parts of the pterygoid bones make up the rostral margin of the choana and thus separate it from the palatines, a feature also seen in the more advanced neosuchian suborder Eusuchia. This characteristic was once thought to be characteristic of Eusuchia, but its presence in Brillanceausuchus suggests that the trait is homoplasic, thus making the evolution of the position of the choana within crocodilians more complex than previously thought.

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

Isisfordia is an extinct genus of crocodyliform closely related to crocodilians that lived in Australia during the Middle Cretaceous (Albian–Cenomanian).

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

Pholidosaurus is an extinct genus of neosuchian crocodylomorph. It is the type genus of the family Pholidosauridae. Fossils have been found in northwestern Germany. The genus is known to have existed during the Berriasian-Albian stages of the Early Cretaceous. Fossil material found from the Annero and Jydegård Formations in Skåne, Sweden and on the island of Bornholm, Denmark, have been referred to as a mesoeucrocodylian, and possibly represent the genus Pholidosaurus.

Shantungosuchus is an extinct genus of Early Cretaceous crocodyliform found in China. It includes three species: Shantungosuchus chuhsienensis and S. brachycephalus, which were both described by Yang Zhongjian – usually referred to as "Young" – in 1961 and 1982, and S. hangjinensis, which was described by Xiao-Chun Wu et al in 1994. S. chuhsienensis is the type for this genus.

Paluxysuchus is an extinct genus of neosuchian crocodyliform known from the Early Cretaceous Twin Mountains Formation of north-central Texas. It contains a single species, Paluxysuchus newmani. Paluxysuchus is one of three crocodyliforms known from the Early Cretaceous of Texas, the others being Pachycheilosuchus and an unnamed species referred to as the "Glen Rose Form". Paluxysuchus has a long, flat skull that is probably transitional between the long and narrow skulls of many early neosuchians and the short and flat skulls of later neosuchians.

Hulkepholis is an extinct genus of goniopholidid mesoeucrocodylian from the Early Cretaceous of southern England and eastern Spain. It contains two species, the type species, Hulkepholis willetti, and also H. plotos. Hulkepholis is most closely related to both species of Anteophthalmosuchus.

References

  1. 1 2 3 Fortier, D.C.; Schultz, C.L. (2009). "A new neosuchian crocodylomorph (Crocodyliformes, Mesoeucrocodylia) from the Early Cretaceous of north-east Brazil" (PDF). Palaeontology. 52 (5): 991–1007. doi: 10.1111/j.1475-4983.2009.00894.x .
  2. Figueiredo, R.G.; Kellner, A.W.A. (2009). "A new crocodylomorph specimen from the Araripe Basin (Crato Member, Santana Formation), northeastern Brazil". Paläontologische Zeitschrift. 83 (2): 323–331. doi:10.1007/s12542-009-0016-6. S2CID   129166567.
  3. 1 2 3 Salisbury, S.W.; Molnar, R.E.; Frey, E.; Willis, P.M.A. (2006). "The origin of modern crocodyliforms: new evidence from the Cretaceous of Australia". Proceedings of the Royal Society B. 273 (1600): 2439–2448. doi:10.1098/rspb.2006.3613. PMC   1634899 . PMID   16959633.
  4. 1 2 3 Salisbury, S.W.; Frey, E.; Martill, D.M.; Buchy, M.C. (2003). "A new crocodilian from the Lower Cretaceous Crato Formation of north-eastern Brazil". Palaeontographica Abteilung A. 270 (1–3): 3–47. ISSN   0375-0442.