Simosuchus

Last updated

Contents

Simosuchus
Temporal range: Late Cretaceous, 70–66  Ma
Ꞓ
O
S
D
C
P
T
J
K
Pg
N
↓
Simosuchus clarki, ROM.jpg
Reconstructed skeleton in Royal Ontario Museum
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Clade: Archosauria
Clade: Pseudosuchia
Clade: Crocodylomorpha
Clade: Crocodyliformes
Clade:† Notosuchia
Clade:† Ziphosuchia
Genus:† Simosuchus
Buckley et al., 2000
Species
  • †S. clarkiBuckley et al., 2000 (type)

Simosuchus is an extinct genus of notosuchian crocodyliforms 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 one isolated multicuspid tooth of this genus was discovered in Kallamedu Formation of India. [1]

The teeth of S. clarki were shaped like maple leaves, which coupled with its short and deep snout suggests it was not a carnivore like most other crocodylomorphs. In fact, these features have led many palaeontologists to consider it a herbivore.

History

Skull FMNH PR 2597 August 1, 2012 - Simosuchus clarki Skull on Display at the Royal Ontario Museum (FMNH PR 2597).jpg
Skull FMNH PR 2597

The first specimen of Simosuchus clarki, which served as the basis for its initial description in 2000, included a complete skull and lower jaw, the front of the postcranial skeleton, and parts of the posterior postcranial skeleton. Five more specimens were later described, representing the majority of the skeleton. Many isolated teeth have also been found in the Mahajanga Basin. Most remains of Simosuchus were found as part of the Mahajanga Basin Project, directed by the Université d'Antananarivo and Stony Brook University. Material was usually found in clays that were part of flow deposits in the Anembalemba Member of the Maevarano Formation. [2]

Description

Size comparison to a human Simosuchus scale.svg
Size comparison to a human

Simosuchus was small, about 0.75 metres (2.5 ft) long based on the skeletons of mature individuals. [3] [4] In contrast to most other crocodyliforms, which have long, low skulls, Simosuchus has a distinctively short snout. The snout resembles that of a pug, giving the genus its name, which means "pug-nosed crocodile" in Greek. [5] The shape of skulls differs considerably between specimens, with variation in ornamentation and bony projections. These differences may be indications of sexual dimorphism. The front (preorbita) portion of the skull is angled downwards. Simosuchus likely held its head so that the preorbital area was angled about 45° from horizontal. The teeth line the front of the jaws and are shaped like maple leaves. At the back of the skull, the occipital condyle (which articulates with the neck vertebrae) is downturned. 45 autapomorphies, or features unique to Simosuchus, can be found in the skull alone. [6]

Life restoration Simosuchus.jpg
Life restoration

In most respects, the postcranial skeleton of Simosuchus resembles that of other terrestrial crocodyliforms. There are several differences, however, that have been used to distinguish it from related forms. The scapula is broad and tripartite (three-pronged). On its surface, there is a laterally directed prominence. The deltopectoral crest, a crest on the upper end of the humerus, is small. The glenohumeral condyle of the humerus, which connects to the pectoral girdle in the shoulder joint, has a distinctive rounded ellipsoid shape. The limbs are robust. The radius and ulna of the forearm fit tightly together. The front feet are small with large claws, and the back feet are also reduced in size. There is a small crest along the anterior edge of the femur. On the pelvis, the anterior process of the ischium is spur-like. [7]

Most of the spinal column of Simosuchus is known. There are eight cervical vertebrae in the neck, at least fifteen dorsal vertebrae in the back, two sacral vertebrae at the hip, and no more than twenty caudal vertebrae in the tail. The number of vertebrae in the tail is less than that of most crocodyliforms, giving Simosuchus a very short tail. [3]

Like other crocodyliforms, Simosuchus was covered in bony plates called osteoderms. These form shields over the back, underside, and tail. Unusually among crocodyliforms, Simosuchus also has osteoderms covering much of the limbs. Osteoderms covering the back, tail, and limbs are light and porous, while the osteoderms covering the belly are plate-like and have an inner structure resembling spongy diploë. Simosuchus has a tetraserial paravertebral shield over its back, meaning that there are four rows of tightly locking paramedial osteoderms (osteoderms to either side of the midline of the back). To either side of the shield, there are four rows of accessory parasagittal osteoderms. These accessory osteoderms tightly interlock with one another. [8]

Classification

Cladograms of Notosuchia
Turner and Calvo, 2005 [9]
Notosuchia  
Pol et al., 2012 [10]
Notosuchia

Simosuchus was first considered to be a basal member of the clade Notosuchia, and was often considered to be closely related to Uruguaysuchus from the Late Cretaceous of Uruguay and Malawisuchus from the Early Cretaceous of Malawi. Later phylogenetic studies have placed it closer to the genus Libycosuchus and in a more derived position than some other notosuchians such as Uruguaysuchus. [11] In its initial description by Buckley et al. (2000), Simosuchus was placed in the family Notosuchidae. Its sister taxon was Uruguaysuchus, and the two were allied with Malawisuchus . These taxa were placed in Notosuchidae along with Libycosuchus and Notosuchus . Most of the following phylogenetic analyses resulted in a similar placement of Simosuchus and other genera within Notosuchia. Turner and Calvo (2005) also found a clade including Simosuchus, Uruguaysuchus, and Malawisuchus in their study.

The phylogenetic analysis of Carvalho et al. (2004), based on different character values than previous studies, produced a very different relationship among Simosuchus and other notosuchians. Simosuchus, along with Uruguaysuchus and Comahuesuchus, were placed outside Notosuchia. Simosuchus was found to be the sister taxon of the Chinese genus Chimaerasuchus in the family Chimaerasuchidae. Like Simosuchus, Chimaerasuchus has a short snout and was probably herbivorous. Both genera were placed outside Notosuchia in the larger clade Gondwanasuchia. Uruguaysuchus, previously considered to be a basal notosuchian and a close relative of Simosuchus, was placed in its own family, Uruguaysuchidae, also outside Notosuchia. Malawisuchus was found to be a member of Peirosauroidea, specifically a member of the family Itasuchidae. [12]

The following cladogram simplified after a comprehensive analysis of notosuchians which focused on Simosuchus clarki presented by Alan H. Turner and Joseph J. W. Sertich in 2010. [11]

Notosuchia

*Note: Based on a specimen that was reassigned from Peirosaurus . [13]

Paleobiology

Reconstructed skull in the Field Museum of Natural History Simosuchus clarki skull.jpg
Reconstructed skull in the Field Museum of Natural History

Simosuchus was probably a herbivore, with its complex dentition resembling that of herbivorous iguanids. [14] Like other notosuchians, it was fully terrestrial, and the short tail would have had little use in swimming. [3]

The osteoderm shield was inflexible, restricting lateral movement in Simosuchus as a possible adaptation to an entirely terrestrial lifestyle. Robust legs are also consistent with terrestrial locomotion. The deltopectoral crest on the humerus and the anterior crest on the femur served as attachment points for strong limb muscles. The hindlimbs of Simosuchus were semierect, unlike the fully erect posture of most other notosuchians. [7]

Possible fossorial lifestyle

A fossorial, or burrowing, lifestyle for Simosuchus has been suggested in its initial description based on the robust limbs and short snout, which appears shovel-like, and the underslung lower jaw that would prevent friction when the animal opens its jaws during burrowing. There are also areas on the skull that may have attached to strong neck muscles that would have been well suited for burrowing. [5] [15]

Outdated restoration done before discovery of whole-body skeleton Simosuchus BW.jpg
Outdated restoration done before discovery of whole-body skeleton

In 2010, Kley and colleagues have argued against this hypothesis on the basis of its morphology which doesn't show an adaptation as head-first burrowers, with its relatively large, wide head being disadvantageous and contrasting modern head-first burrowers. They also pointed out that the snout is not shovel-like, since it doesn't have a "dorsoventrally narrowed cutting edge" on it, but rather has a "relatively expansive anterior surface" which is almost completely flat. The lower jaw also wouldn't be "shielded" to a significant extent in response to the reaction force during burrowing; even the shielded portion, the premaxillary teeth which is crucial for feeding, would not have been able to endure the "extreme loading regimes associated with head-first burrowing." The upper jaw was also suggested to be not adapted for keeping its mouth shut when faced by reaction forces associated with burrowing. Its relatively large orbits were also inconsistent with a burrowing lifestyle, since large eyes have a risk of being damaged in head-first burrowing. [6]

In the same year, Sertich and Groenke noted that while the appendicular skeleton including the scapula and the forelimb doesn't show specific morphological adaptation for digging, it doesn't exclude the idea that Simosuchus was adept at burrowing, since many extant crocodylians that are able to burrow also don't show any of the specific morphological specializations. [7] Georgi and Krause, also in the same year, agreed that the burrowing hypothesis couldn't be confidently excluded, and claimed that the hypothesis needs more extensive comparative and functional analyses. They also suggested that its relatively short neck would have been a potential advantage for moving through a dense medium like a tunnel, and that it may have likely used its head on occasions to aid scratch-digging with its limbs. [3]

Paleobiogeography

It is unknown how Simosuchus arrived in Madagascar. A similar crocodyliform, Araripesuchus tsangatsangana , is also known from the Maevarano Formation, but its relation to Simosuchus is unclear. [16] It has been classified as both a notosuchian and a basal neosuchian in various phylogenetic analyses. [17] Nearly all notosuchians are known from Gondwana, the southern supercontinent that existed throughout much of the Mesozoic and encompassed South America, Africa, India, Australia, and Antarctica. Libycosuchus, regarded as one of the closest relatives of Simosuchus, lived in Egypt. Unresolved relationships among notosuchians along with an incomplete fossil record have made it difficult to determine the biogeographic origins of Simosuchus.

See also

Related Research Articles

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

Mahajangasuchus is an extinct genus of crocodyliform which had blunt, conical teeth. The type species, M. insignis, lived during the Late Cretaceous; its fossils have been found in the Maevarano Formation in northern Madagascar. It was a fairly large predator, measuring up to 4 metres (13 ft) long.

Trematochampsa is a dubious extinct genus of crocodyliform from the Late Cretaceous In Beceten Formation of Niger.

Elosuchus is an extinct genus of neosuchian crocodyliform that lived during the Middle Cretaceous of what is now Africa.

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.

<span class="mw-page-title-main">Notosuchia</span> Extinct suborder of reptiles

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 ; 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.

<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.

<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 crocodyliforms. 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>Armadillosuchus</i> Extinct genus of reptiles

Armadillosuchus is an extinct genus of sphagesaurid crocodylomorph. It was described in February 2009 from the late Campanian to early Maastrichtian Adamantina Formation of the Bauru Basin in Brazil, dating to approximately 70 Ma. Armadillosuchus was among the larger and more robust sphagesaurids, with a total length of approximately 2 metres (6.6 ft).

Stolokrosuchus is an extinct genus of crocodyliforms that lived during the Early Cretaceous. Its fossils, including a skull with a long thin snout and bony knobs on the prefrontal, have been found in Niger. Stolokrosuchus was described in 2000 by Hans Larsson and Boubacar Gado. The type species is S. lapparenti. They initially described it as related to Peirosauridae, if not a member of that family. One study has shown it to be related to Elosuchus. However, more recent works usually find Stolokrosuchus to be one of the basalmost neosuchian, only distantly related to the elosuchid or pholidosaurid, Elosuchus. It was a semiaquatic crocodylomorph.

<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.

<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.

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

Pakasuchus is a genus of notosuchian crocodyliform distinguished by its unusual mammal-like appearance, including mammal-like teeth that would have given the animal the ability to chew. It also had long, slender legs and a doglike nose. Fossils have been found in the Galula Formation of Rukwa Rift Basin of southwestern Tanzania, and were described in 2010 in the journal Nature. Pakasuchus is originally considered to lived approximately 105 million years ago, in the mid-Cretaceous, but later age of site is reconsidered to the late Cretaceous, Cenomanian to Campanian instead. The type species is P. kapilimai. Pakasuchus means "cat crocodile" in reference to its catlike skull.

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>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.

Gasparinisuchus is an extinct genus of peirosaurid notosuchian known from the Late Cretaceous of Neuquén and Mendoza Provinces, western central Argentina. It contains a single species, Gasparinisuchus peirosauroides.

Rukwasuchus is an extinct genus of peirosaurid crocodyliforms known from the middle Cretaceous Galula Formation of southwestern Tanzania. It contains a single species, Rukwasuchus yajabalijekundu.

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

Anthracosuchus is an extinct genus of dyrosaurid crocodyliform from the Paleocene of Colombia. Remains of Anthracosuchus balrogus, the only known species, come from the Cerrejón Formation in the Cerrejón mine, and include four fossil specimens with partial skulls. Anthracosuchus differs from other dyrosaurids in having an extremely short (brevirostrine) snout, widely spaced eye sockets with bony protuberances around them, and osteoderms that are smooth and thick. It is one of the most basal dyrosaurids along with Chenanisuchus and Cerrejonisuchus.

Barrosasuchus is a genus of peirosaurid notosuchian from the Santonian of Argentina and part of the extensive peirosaurid record of Late Cretaceous Patagonia. It contains one species, Barrosasuchus neuquenianus. B. neuquenianus is known from an almost complete skull and the majority of the articulated postcranial skeleton, making it the best preserved Patagonian peirosaurid.

References

  1. ↑ Prasad, Guntupalli V. R.; Verma, Omkar; Flynn, John J.; Goswami, Anjali (2013). "A new Late Cretaceous vertebrate fauna from the Cauvery Basin, South India: implications for Gondwanan paleobiogeography". Journal of Vertebrate Paleontology. 33 (6): 1260–1268. doi:10.1080/02724634.2013.777348. S2CID   131192480.
  2. ↑ Krause, D.W.; Sertich, J.J.W.; Rogers, R.R.; Kast, S.C.; Rasoamiaramanana, A.H.; Buckley, G.A. (2010). "Overview of the discovery, distribution, and geological context of Simosuchus clarki (Crocodyliformes: Notosuchia) from the Late Cretaceous of Madagascar". Journal of Vertebrate Paleontology. 30 (6, Supplement): 4–12. doi:10.1080/02724634.2010.516784. S2CID   129124296.
  3. 1 2 3 4 Georgi, J.A.; Krause, D.W. (2010). "Postcranial axial skeleton of Simosuchus clarki (Crocodyliformes: Notosuchia) from the Late Cretaceous of Madagascar". Journal of Vertebrate Paleontology. 30 (6, Supplement): 99–121. doi:10.1080/02724634.2010.519172. S2CID   84710186.
  4. ↑ Michael J. Benton (2015). Vertebrate Palaeontology. John Wiley & Sons, Ltd. p. 249. ISBN   9781118407554 . Retrieved 25 September 2022.
  5. 1 2 Buckley, G.A.; Brochu, C.A.; Krause, D.W.; Pol, D. (2000). "A pug-nosed crocodyliform from the Late Cretaceous of Madagascar". Nature. 405 (6789): 941–944. Bibcode:2000Natur.405..941B. doi:10.1038/35016061. PMID   10879533. S2CID   4426288.
  6. 1 2 Kley, N.J.; Sertich, J.J.W.; Turner, A.H.; Krause, D.W.; O'Connor, P.M.; Georgi, J.A. (2010). "Craniofacial morphology of Simosuchus clarki (Crocodyliformes: Notosuchia) from the Late Cretaceous of Madagascar". Journal of Vertebrate Paleontology. 30 (6, Supplement): 13–98. doi:10.1080/02724634.2010.532674. S2CID   84437972. CC-BY icon.svg Text was copied from this source, which is available under a Creative Commons Attribution 4.0 International License.
  7. 1 2 3 Sertich, J.J.W.; Groenke, J.R. (2010). "Appendicular skeleton of Simosuchus clarki (Crocodyliformes: Notosuchia) from the Late Cretaceous of Madagascar". Journal of Vertebrate Paleontology. 30 (6, Supplement): 122–153. doi:10.1080/02724634.2010.516902. S2CID   86113454.
  8. ↑ Hill, R.V. (2010). "Osteoderms of Simosuchus clarki (Crocodyliformes: Notosuchia) from the Late Cretaceous of Madagascar". Journal of Vertebrate Paleontology. 30 (6, Supplement): 154–176. doi:10.1080/02724634.2010.518110. S2CID   85720441.
  9. ↑ Turner, A.H.; Calvo, J.O. (2005). "A new sebecosuchian crocodyliform from the Late Cretaceous of Patagonia". Journal of Vertebrate Paleontology. 25 (1): 87–98. doi:10.1671/0272-4634(2005)025[0087:ANSCFT]2.0.CO;2. S2CID   86257810.
  10. ↑ Diego Pol; Juan M. Leardi; Agustina Lecuona & Marcelo Krause (2012). "Postcranial anatomy of Sebecus icaeorhinus (Crocodyliformes, Sebecidae) from the Eocene of Patagonia". Journal of Vertebrate Paleontology. 32 (2): 328–354. doi:10.1080/02724634.2012.646833. S2CID   86565764.
  11. 1 2 Turner, A.H.; Sertich, J.W. (2010). "Phylogenetic history of Simosuchus clarki (Crocodyliformes: Notosuchia) from the Late Cretaceous of Madagascar". Journal of Vertebrate Paleontology. 30 (6, Supplement): 177–236. doi:10.1080/02724634.2010.532348. S2CID   86737170.
  12. ↑ Carvalho, I.S.; Ribeiro, L.C.B.; Avilla, L.S. (2004). "Uberabasuchus terrificus sp. nov., a new Crocodylomorpha from the Bauru Basin (Upper Cretaceous), Brazil" (PDF). Gondwana Research. 7 (4): 975–1002. Bibcode:2004GondR...7..975C. doi:10.1016/S1342-937X(05)71079-0. ISSN   1342-937X. Archived from the original (PDF) on 2011-07-06.
  13. ↑ Agustín G. Martinelli; Joseph J.W. Sertich; Alberto C. Garrido & Ángel M. Praderio (2012). "A new peirosaurid from the Upper Cretaceous of Argentina: Implications for specimens referred to Peirosaurus torminni Price (Crocodyliformes: Peirosauridae)". Cretaceous Research. 37: 191–200. doi:10.1016/j.cretres.2012.03.017.
  14. ↑ Melstrom, Keegan; Irmis, Randall (2019-06-01). "Repeated Evolution of Herbivorous Crocodyliforms during the Age of Dinosaurs". Current Biology. 29 (14): 2389–2395.e3. doi: 10.1016/j.cub.2019.05.076 . PMID   31257139.
  15. ↑ Krause, D.W.; O'Connor, P.M.; Rogers, K.C.; Sampson, S.D.; Buckley, G.A.; Rogers, R.R. (2006). "Late Cretaceous terrestrial vertebrates from Madagascar: Implications for Latin American biogeography". Annals of the Missouri Botanical Garden. 93 (2): 178–20. doi:10.3417/0026-6493(2006)93[178:LCTVFM]2.0.CO;2. S2CID   9166607.
  16. ↑ Turner, A.H. (2006). "Osteology and phylogeny of a new species of Araripesuchus (Crocodyliformes: Mesoeucrocodylia) from the Late Cretaceous of Madagascar". Historical Biology. 18 (3): 255–369. doi:10.1080/08912960500516112. S2CID   83717721.
  17. ↑ Sereno, P. C.; Larsson, H. C. E. (2009). "Cretaceous crocodyliforms from the Sahara". ZooKeys (28): 1–143. doi: 10.3897/zookeys.28.325 .
This page is based on this Wikipedia article
Text is available under the CC BY-SA 4.0 license; additional terms may apply.
Images, videos and audio are available under their respective licenses.