Mourasuchus

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Mourasuchus
Temporal range: Mid-Late Miocene (Friasian-Huayquerian),
16–5.3  Ma [1]
Mourasuchus pattersoni - skull - Urumaco Formation - Venezuela.jpg
Skull of Mourasuchus pattersoni
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Clade: Archosauromorpha
Clade: Archosauriformes
Order: Crocodilia
Family: Alligatoridae
Subfamily: Caimaninae
Genus:† Mourasuchus
Price 1964
Species
  • †M. amazonensisPrice 1964
  • †M. atopusLangston 1966
  • †M. arendsiBocquentin-Villanueva 1984
  • †M. pattersoniCidade et al. 2017
Synonyms

Family-level:

  • Nettosuchidae Langston 1965

Genus-level:

  • CaraindasuchusGasparini 1985
  • NettosuchusLangston 1965

Mourasuchus is an extinct genus of giant, aberrant caiman from the Miocene of South America. Its skull has been described as duck-like, being broad, flat, and very elongate, superficially resembling Stomatosuchus from the Late Cretaceous. [2] [3]

Contents

History of discovery

Mourasuchus was first described by Price in 1964 based on a strange and nearly complete skull from the Solimões Formation of Amazonian Brazil, calling it Mourasuchus amazoniensis. [4] Unaware of Price's discovery, Langston described "Nettosuchus" atopus ("Absurd Duck Crocodile") only a year later based on fragmentary cranial, mandibular and postcranial remains from the middle Miocene La Venta Lagerstätte, a part of the Honda Group. Although he did recognize its similarities to caimans and alligators, Langston reasoned that its bizarre anatomy warranted its own monotypic family, naming it Nettosuchidae. [5] After being informed about the existence of Mourasuchus by Mr. W.D. Sills, Langston wrote a follow-up publication acknowledging his taxon to be a junior synonym of Mourasuchus. [6]

A third species was found in the Urumaco Formation of Venezuela in 1984 and named Mourasuchus arendsi by Bocquentin-Villanueva. Following this discovery, "Carandaisuchus" nativus was named in 1985 based on fossils of the Ituzaingó Formation of Argentina, as well as remains from Brazil and Bolivia. [7] However, by 1990 Caraindaisuchus had been lumped into Mourasuchus [8] and Scheyer & Delfino (2016) eventually concluded that M. nativus was merely a junior synonym of M. arendsi, [9] bringing the number of species from four to three. It wasn't long until the genus returned to four species however, with M. pattersoni (named after Anglo-American paleontologist Bryan Patterson) being described by Cidade et al. the following year. [10] Indeterminate Mourasuchus fossils were found in the Yecua Formation of Bolivia. [11]

Description

Occlusal pits in M. atopus, M. arendsi & M. pattersoni Mourasuchus pattersoni - external naris - Urumaco Formation - Venezuela.jpg
Occlusal pits in M. atopus, M. arendsi & M. pattersoni
South America laea relief location map.jpg
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Mourasuchus fossil localities
White pog.svg Ituzaingó Formation
Yellow ffff80 pog.svg Yecua Formation
Orange ff8040 pog.svg Urumaco Formation
Yellow ffff00 pog.svg Solimões Formation
Yellow pog.svg Honda Group
Gold pog.svg Pebas Formation

Mourasuchus is well known for its strange anatomy, its skull exceptionally dorsoventrally flattened, broad, and overall platyrostral in shape, differing greatly from any other caiman currently known, most closely resembling the enigmatic crocodyliform Stomatosuchus from the Late Cretaceous Bahariya Formation. The nares are elevated and so is the high postrostral cranium and short skull table. The orbital margins are likewise raised above the rostrum with thickened anteromedial margins formed by a knob-like swelling of the frontal and prefrontal bones. Overall, the orbits are smaller than the infratemporal fenestrae. The teeth were generally small and conical, numbering upwards of 40 teeth on each side of the upper and lower jaw and interlocking perfectly. The posterior teeth of the dentary and maxilla are laterally compressed. The premaxilla of Mourasuchus show noticeably large occlusal pits, especially noticeable in M. arendsi. However, such pits can be observed in a variety of extant and fossil taxa and are not considered to be diagnostic between species of Mourasuchus.

The mandible is U-shaped with a sharp transition between the first five teeth and the rest of the dentary teeth. The first to fifth tooth are located on the curved anterior portion of the mandible, while the bone is straight from the sixth onward. The mandibular symphysis is short and slender, only extending to the first posterior alveoli, meaning the animal most likely had a relatively weak bite. The osteoderms of Mourasuchus show conspicuous spines on their dorsal surface. [12]

The humerus of Mourasuchus is long and slender and interpreted to possibly indicate weakened forelimbs, potentially corresponding with a predominantly aquatic lifestyle. [13]

Size

Due to the fragmentary remains of Mourasuchus the body size is difficult to determine. Mourasuchus skulls range up to a meter (3.3 ft) in length, with the holotype skull of M. pattersoni being 108.1 centimetres (42.6 in) long while that of M. arendsi could reach a length of 108.5 centimetres (42.7 in) long. The largest skull belongs to M. amazonensis at 113.5 centimetres (44.7 in). In a 2020 study, Cidade et al. attempted to determine the body sizes of the four recognized Mourasuchus species based on the head: body ratio of modern genera like Caiman latirostris and Alligator mississippiensis . Their results recovered a mean body size of 6.3 m (21 ft) and 1.2 metric tons (1.3 short tons) for M. atopus, the smallest species, and upwards of 9.5 m (31 ft) and 4.4 metric tons (4.9 short tons) for M. amazonensis. [13] However, a later study contradicted these results. In 2022 Paiva and colleagues argued that the dorsal cranial length was a poor basis for size estimates compared to the width of the skull. Additionally, they found that prior studies commonly included juvenile animals in their data, muddying the results. This study calculated a length between 2.5–3.1 m (8 ft 2 in – 10 ft 2 in) for Mourasuchus atopus, 3.8–4.8 m (12–16 ft) for Mourasuchus arendsi, 4.3–5.5 m (14–18 ft) for Mourasuchus pattersoni and for Mourasuchus amazoniensis, the largest species, 4.7–5.98 m (15.4–19.6 ft). These results were achieved by specifically restricting the dataset to extant species of caimans, while calculations using measurements from all of Crocodilia generally rendered greater sizes but may be overestimates. Still, the authors note that the generally smaller stature of modern caimans may have influenced the results in a similar fashion. [14]

Phylogeny

Although Mourasuchus has been firmly placed within Caimaninae by authors, the exact relationship it has with other crocodilians of this family and even the relationship between the different species of the genus had long been unclear. Some studies have proposed that Mourasuchus was closely related to the Eocene Orthogenysuchus from North America, and more distantly to the giant caiman Purussaurus which it shared its habitat with. However, more recent papers indicate that ongoing preparation conducted on Orthogenysuchus significantly influences the scoring of this taxon's characters, leading to some authors removing the animal from analysis until further publications. The cladogram below shows the phylogenetic tree used by Bona et al. (2012). [15]

The following tree is based on the results recovered by Cidade et al. (2017), excluding the North American Orthogenysuchus and including the then newly named Mourasuchus pattersoni while also following the synonymy of M. nativus with M. arendsi. Like Bona et al. before them, the authors recover Mourasuchus and Purussaurus as closely related clades, this time as sister genera with Centenariosuchus just outside this shared clade. [10]

Paleobiology

Diet

Mourasuchus atopus size Mourasuchus atopus size.jpg
Mourasuchus atopus size

Much like with the unrelated but morphologically similar Stomatosuchidae, the feeding ecology of Mourasuchus is enigmatic and poorly understood, with a variety of hypotheses having been proposed to make sense of its specialized anatomy. With slender and short mandibular rami and perfectly occluding, slender and almost homodont teeth, Mourasuchus was not built to capture and hold prey like modern crocodilians. [8] [5] In his 1965 publication on the crocodilians of Cenozoic Columbia, Langston proposed three ideas on how Mourasuchus may have fed. He suggests that it may have waited with opened jaws for approaching prey, fed on aquatic and floating plant material or searched for prey by sifting through substrate at the bottom of bodies of water. [5] While his other hypothesis have gone without much attention, the later of the three has been discussed by later publications, referring to this ecology as either "filter feeding", "straining technique" or "gulp feeding". Cidade et al. follow Langston's proposed feeding hypothesis, speculating that Mourasuchus may have fed on small invertebrates, mollusks, crustaceans, and small fish, specializing in trying to ingest as much prey at once as possible. This they suggest explains the platyrostral-broad skull morphology, which differs from the usual crocodilian skull morphology that typically either favors longirostrine forms providing speediness or robust-snouted forms with increased bite force. [8] Langston also proposed a throat pouch based on comments Franz Nopsca made regarding Stomatosuchus, however there is no evidence for such a structure in Mourasuchus nor is its presence certain in stomatosuchids. As the prey Mourasuchus would have fed on in accordance with the straining technique hypothesis would likely be found either in free water or substrate, the animal would have also captured inedible material during feeding. For this a type of filtering mechanism or behavior would be beneficial, however nothing indicating as much has been found so far. Due to the absence of evidence for actual filtering, Cidade et al. instead favors the term "gulp feeding" for Mourasuchus hypothetical hunting behavior.

In a 2020 study Cidade et al. aimed to determine Mourasuchus ability to perform the death roll, a crucial behavior observed in modern crocodiles and alligators that is commonly observed during feeding, when crocodiles rapidly spin to rip pieces of flesh from their prey. The study calculated a very low "death roll" capability indicator (DRCI) similar to that seen in the Slender Snouted Crocodile, Indian Gharial and False Gharial, which suggests that Mourasuchus was very unlikely to be able to perform the death roll.

Paleoenvironment

Mourasuchus lived during an era of great crocodilian diversity in South America, being found in environments shared with genera inhabiting a diverse set of niches. In the Peruvian Pebas Formation M. atopus was found alongside the large bodied Purussaurus neivensis , a medium-sized species of Gryposuchus , G. pachakamue and three distinct taxa of durophagous caimans. The later species M. arendsi coexisted with an even greater host of contemporary crocodilians in the Urumaco Formation, sharing its habitat with the larger Purussaurus mirandai, several large-bodied gharials, multiple durophagous caimans and the true crocodile Charactosuchus . In some areas such as the Urumaco and Solimões Formation multiple species of Mourasuchus are known, with the former being home to both M. pattersoni and M. arendsi while the later has yielded M. arendsi and M. amazonensis. [8] The great diversity of crocodylomorphs in these Miocene-age (Tortonian stage, 8 million years ago) wetlands suggests that niche partitioning was efficient, which would have limited interspecific competition. [16]

Related Research Articles

<span class="mw-page-title-main">Alligatoridae</span> Family of crocodilians including alligators, caimans and kin

The family Alligatoridae of crocodylians includes alligators, caimans and their extinct relatives.

<i>Caiman</i> (genus) Genus of reptiles

Caiman is a genus of caimans within the alligatorid subfamily Caimaninae. They inhabit Central and South America. They are relatively small sized crocodilians, with all species reaching lengths of only a couple of meters and weighing 6 to 40 kg on average.

<i>Melanosuchus</i> Genus of caiman

Melanosuchus is a genus of caiman. The black caiman of South America is the sole extant (living) species, and is the largest living member of the subfamily Caimaninae, as well as the entire alligator family Alligatoridae.

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

Sarcosuchus is an extinct genus of crocodyliform and distant relative of living crocodilians that lived during the Early Cretaceous, from the late Hauterivian to the early Albian, 133 to 112 million years ago of what is now Africa and South America. The genus name comes from the Greek σάρξ (sarx) meaning flesh and σοῦχος (souchus) meaning crocodile. It was one of the largest pseudosuchians, with the largest specimen of S. imperator reaching approximately 9–9.5 metres (29.5–31.2 ft) long and weighing up to 3.45–4.3 metric tons. It is known from two species; S. imperator from the early Albian Elrhaz Formation of Niger, and S. hartti from the Late Hauterivian of northeastern Brazil. Other material is known from Morocco and Tunisia and possibly Libya and Mali.

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

Purussaurus is an extinct genus of giant caiman that lived in South America during the Miocene epoch, from the Friasian to the Huayquerian in the SALMA classification. It is known from skull material found in the Brazilian and Peruvian Amazon, Colombian Villavieja Formation, Panamanian Culebra Formation, Urumaco and Socorro Formations of northern Venezuela.

<span class="mw-page-title-main">Orinoco crocodile</span> Species of reptile

The Orinoco crocodile is a critically endangered crocodile. Its population is very small, and they can only be found in the Orinoco river basin in Venezuela and Colombia. Extensively hunted for their skins in the 19th and 20th centuries, it is one of the most endangered species of crocodiles. It is a very large species of crocodilian; males have been reported up to 6.8 m in the past, weighing over 900 kg (2,000 lb), but such sizes do not exist today, 5.2 m being a more widely accepted maximum size. A large male today may attain 4.2 m in length and can weigh up to 450 kg (1,000 lb), while females are substantially smaller with the largest likely to weigh around 225 kg (496 lb). Sexual dimorphism is not as profound as in other crocodilian species. The coloration is light even in adults.

<span class="mw-page-title-main">Cuvier's dwarf caiman</span> Species of reptile

Cuvier's dwarf caiman is a small crocodilian in the alligator family from northern and central South America. It is found in Bolivia, Brazil, Colombia, Ecuador, French Guiana, Guyana, Paraguay, Peru, Suriname, Trinidad and Venezuela. It lives in riverine forests, flooded forests near lakes, and near fast-flowing rivers and streams. It can traverse dry land to reach temporary pools and tolerates colder water than other species of caimans. Other common names for this species include the musky caiman, the dwarf caiman, Cuvier's caiman, and the smooth-fronted caiman. It is sometimes kept in captivity as a pet and may be referred to as the wedge-head caiman by the pet trade community.

<span class="mw-page-title-main">Caiman</span> Subfamily of reptiles

A caiman is an alligatorid belonging to the subfamily Caimaninae, one of two primary lineages within the Alligatoridae family, the other being alligators. Caimans inhabit Mexico and Central and South America from marshes and swamps to mangrove rivers and lakes. They have scaly skin and live a fairly nocturnal existence. They are relatively small-sized crocodilians with an average maximum weight of 6 to 40 kg depending on species, with the exception of the black caiman, which can grow more than 4 m (13 ft) in length and weigh in excess of 1,000 kg. The black caiman is the largest caiman species in the world and is found in the slow-moving rivers and lakes that surround the Amazon basin. The smallest species is the Cuvier's dwarf caiman, which grows to 1.2 to 1.5 m long. There are six different species of caiman found throughout the watery jungle habitats of Central and Southern America. The average length for most of the other caiman species is about 2 to 2.5 m long.

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

Necrosuchus is an extinct genus of caiman from modern day Argentina that lived during the Paleocene epoch. It inhabited the fluvio-lacustrine environment of the Patagonian Salamanca Formation.

<i>Langstonia</i> Extinct species of reptile

Langstonia is an extinct genus of notosuchian crocodylomorph of the family Sebecidae. It lived in the middle Miocene, in the "Monkey Beds" of the Colombian Villavieja Formation. Langstonia was named in 2007 by Alfredo Paolillo and Omar Linares for fossils originally described by Langston in 1965 as Sebecus huilensis. Thus, the type species is L. huilensis.

<i>Gryposuchus</i> Extinct genus of gavialoid crocodilian

Gryposuchus is an extinct genus of gavialid crocodilian. Fossils have been found from Argentina, Colombia, Venezuela, Brazil and the Peruvian Amazon. The genus existed during the Miocene epoch. One recently described species, G. croizati, grew to an estimated length of 10 metres (33 ft). Gryposuchus is the type genus of the subfamily Gryposuchinae, although a 2018 study indicates that Gryposuchinae and Gryposuchus might be paraphyletic and rather an evolutionary grade towards the gharial.

Ikanogavialis is an extinct genus of gavialid crocodilian. Fossils have been found in the Urumaco Formation in Urumaco, Venezuela and the Solimões Formation of Brazil. The strata from which remains are found are late Miocene in age, rather than Pliocene as was once thought. A possible member of this genus survived into the Late Holocene on Muyua or Woodlark Island in Papua New Guinea.

Orthogenysuchus is an extinct genus of caimanine alligatorids. Fossils have been found from the Wasatch Beds of the Willwood Formation of Wyoming, deposited during the early Eocene. The type species is O. olseni. The holotype, known as AMNH 5178, is the only known specimen belonging to the genus and consists of a skull lacking the lower jaws. The braincase is filled in by the matrix and most of the suture lines between bones are indiscernible, making comparisons with other eusuchian material difficult.

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

Stomatosuchidae is an extinct family of neosuchian crocodylomorphs. It is defined as the most inclusive clade containing Stomatosuchus inermis but not Notosuchus terrestris, Simosuchus clarki, Araripesuchus gomesii, Baurusuchus pachecoi, Peirosaurus torminni, or Crocodylus niloticus. Two genera are known to belong to Stomatosuchidae: Stomatosuchus, the type genus, and Laganosuchus. Fossils have been found from Egypt, Morocco, and Niger. Both lived during the Cenomanian stage of the Late Cretaceous. The skulls of stomatosuchids are said to be platyrostral because they have unusually flattened, elongate, duck-shaped craniums with U-shaped jaws. This platyrostral condition is similar to what is seen in the "nettosuchid" Mourasuchus, which is not closely related to stomatosuchids as it is a more derived alligatoroid that existed during the Miocene.

Globidentosuchus is an extinct genus of basal caimanine crocodylian known from the late Middle to Late Miocene of the Middle and the Upper Members of the Urumaco Formation at Urumaco, Venezuela. Its skull was very short and robust, with large units of spherical teeth used to break the shells of molluscs as part of its durophagus diet. It is thought to be one of the most basal Caimanines, even sharing some traits with alligatorids.

<i>Caiman wannlangstoni</i> Extinct species of reptile

Caiman wannlangstoni is an extinct species of caiman that lived in what is now the Amazon Basin and surrounding areas during the Middle and Late Miocene. Fossils of C. wannlangstoni have been found in the Pebas Formation near Iquitos in Peru and include partial skulls and isolated skull bones. Other fossils were uncovered from the Urumaco Formation in Venezuela and the Laventan Honda Group of Colombia. The species was first described in 2015. Features that in combination distinguish C. wannlangstoni from other caimans include a deep snout, a wavy upper jaw margin, a large and upward-directed narial opening, and blunt teeth at the back of the jaws. Based on the sizes of the skulls, its estimated body length is about 211 to 227 centimetres.

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

Gnatusuchus is an extinct genus of caiman represented by the type species Gnatusuchus pebasensis from the Middle Miocene Pebas Formation of Peru. Gnatusuchus lived about 13 million years ago (Ma) in a large wetland system called the Pebas mega-wetlands that covered over one million square kilometers of what is now the Amazon Basin.

The Urumaco Formation is a formation in Venezuela that includes deposits from the Late Miocene. It is the site of several "giant forms": the turtles, crocodiles, sloths and rodents of Urumaco are among the largest of their groups.

Caiman brevirostris is an extinct species of caiman that lived during the Late Miocene, around 11.6 million years ago, to the end of the Miocene 5.3 million years ago in Acre and Amazonas, Brazil as well as Urumaco, Venezuela. Several specimens have been referred to the species, but only 3 of them are confidently placed in the species. C. brevirostris was originally named in 1987 on the basis of a single, incomplete rostrum with an associated mandibular ramus that had been found in Acre, Brazil. C. brevirostris is very distinct among Caiman species and caimaninae overall in that it preserves a characteristically short and robust skull that bears blunt posterior teeth that were built to break down harder foods. This was an adaption for durophagy, likely to crush shells of mollusks and clams which were common in the wetlands that C. brevirostris resided in.

Acresuchus is an extinct monospecific genus of medium-sized caiman from the Late Miocene of western Brazil and Venezuela. The genus contains a single species, Acresuchus pachytemporalis. Acresuchus is a close relative of the giant caiman Purussaurus.

References

  1. ↑ Rio, Jonathan P.; Mannion, Philip D. (6 September 2021). "Phylogenetic analysis of a new morphological dataset elucidates the evolutionary history of Crocodylia and resolves the long-standing gharial problem". PeerJ . 9: e12094. doi: 10.7717/peerj.12094 . PMC   8428266 . PMID   34567843.
  2. ↑ Brochu, C. A. (1999). "Phylogenetics, Taxonomy, and Historical Biogeography of Alligatoroidea". Society of Vertebrate Paleontology Memoir. 6: 9–100. doi:10.2307/3889340. JSTOR   3889340.
  3. ↑ Villanueva, J. B.; Souza Filho, J. P. (1990). "O crocodiliano sul-americano Carandaisuchus como sinonímia de Mourasuchus (Nettosuchidae)". Revista Brasileira de Geociências . 20: 230–233. doi: 10.25249/0375-7536.1990230233 .
  4. ↑ Price, L. I. (1964). "Sôbre o crânio de um grande crocodilídeo extinto do alto Rio Juruá, Estado do Acre". An. Acad. Bras. Ciênc. 36 (1): 59–66.
  5. 1 2 3 Langston, W. (1965). Fossil crocodilians from Colombia and the Cenozoic history of the Crocodilia in South America. University of California Publications in Geological Sciences 52. University of California Press. pp. 1–152.
  6. ↑ Langston, W. (1966). "Mourasuchus Price, Nettosuchus Langston, and the family Nettosuchidae (Reptilia: Crocodilia)". Copeia. 1966 (4): 882–885. doi:10.2307/1441424. JSTOR   1441424.
  7. ↑ Gasparini, Z. B. (1985). "Un nuevo cocodrilo (Eusuchia) Cenozoico de América del Sur". Coletânea de Trabalhos Paleontológicos do IIX Congresso Brasileiro de Paleontologia, MME-DNPM. 27: 51–53.
  8. 1 2 3 4 Bocquentin, Jean-Claude; De Souza Filho, Jonas Pereira (1990). "O Crogodiliano Sul-Americano Carandaisuchus como sinonímia de Mourasuchus (Nettosuchidae)". Revista Brasileira de Geociências. 20: 230–233. doi: 10.25249/0375-7536.1990230233 .
  9. ↑ Scheyer, T.M., and M. Delfino. 2016. The late Miocene caimanine fauna (Crocodylia: Alligatoroidea) of the Urumaco Formation, Venezuela. Palaeontologia Electronica 19. 1–57. Accessed 9 October 2018.
  10. 1 2 Cidade, G.M.; A. Solórzano; A.D. Rincón; D. Riff, and A.S. Hsiou. 2017. A new Mourasuchus (Alligatoroidea, Caimaninae) from the late Miocene of Venezuela, the phylogeny of Caimaninae and considerations on the feeding habits of Mourasuchus. PeerJ 5. 1–37. Accessed 9 October 2018. Archived 9 October 2018 at the Wayback Machine
  11. ↑ Tineo, D.E.; P. Bona; L.M. Pérez; G.D. Vergani; G. González; D.G. Poiré; Z. Gasparini, and P. Legarreta. 2015. Palaeoenvironmental implications of the giant crocodylian Mourasuchus (Alligatoridae, Caimaninae) in the Yecua Formation (late Miocene) of Bolivia. Alcheringa 39. 1–12. Accessed 9 October 2018.
  12. ↑ Langston (2008). "Notes on a partial skeleton of Mourasuchus (Crocodylia, Nettosuchidae) from the Upper Miocene of Venezuela". Arquivos do Museu Nacional, Rio de Janeiro. 66 (1): 125–143.
  13. 1 2 Cidade, Giovanne M.; Rincón, Ascanio D.; Solórzano, Andrés (2020). "New cranial and postcranial elements of Mourasuchus (Alligatoroidea: Caimaninae) from the late Miocene of Venezuela and their palaeobiological implications". Historical Biology . 33 (10): 2387–2399. doi:10.1080/08912963.2020.1795844. S2CID   225395230.
  14. ↑ Paiva, Ana Laura S.; Godoy, Pedro L.; Souza, Ray B. B.; Klein, Wilfried; Hsiou, Annie S. (13 August 2022). "Body size estimation of Caimaninae specimens from the miocene of South America". Journal of South American Earth Sciences. 118: 103970. Bibcode:2022JSAES.11803970P. doi: 10.1016/j.jsames.2022.103970 . ISSN   0895-9811. S2CID   251560425.
  15. ↑ Bona, Paula; Degrange, Frederico J.; Fernández, Marta S. (2012). "Skull Anatomy of the Bizarre Crocodylian Mourasuchus nativus (Alligatoridae, Caimaninae)". Anatomical Record. 296 (2): 227–239. doi:10.1002/ar.22625. hdl: 11336/24984 . PMID   23193096. S2CID   32793915.
  16. ↑ Salas Gismondi, R.; P.O. Antoine; P. Baby; S. Brusset; M. Benammi; N. Espurt; D. de Franceschi; F. Pujos, and J. Tejada and M. Urbina. 2007. Middle Miocene crocodiles from the Fitzcarrald Arch, Amazonian Peru In: Díaz-Martínez, E. and Rábano, I. (eds.), 4th European Meeting on the Palaeontology and Stratigraphy of Latin America. Cuadernos del Museo Geominero, Instituto Geológico y Minero de España 8. . Accessed 9 October 2018.
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