Bergisuchus

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Bergisuchus
Temporal range: Middle Eocene
Early to Middle Lutetian
Bergisuchus dietrichbergi - mandibula fragment.jpg
Bergisuchus dietrichbergi holotype mandible, HLMD-Me 7003.
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:† Sebecosuchia
Clade:† Sebecia
Family:† Bergisuchidae
Rossmann, Rauhe & Ortega 2000
Genus:† Bergisuchus
Kuhn, 1968
Species:
†B. dietrichbergi
Binomial name
†Bergisuchus dietrichbergi
Kuhn, 1968

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.

As with other sebecosuchians, it is likely that Bergisuchus was a fast, terrestrial predator and not an aquatic ambush hunter like modern crocodylians. Its presence in Europe is also unusual, as later sebecosuchians were restricted entirely to South America, and so Bergisuchus indicates the group was once much more widespread in the early Cenozoic.

History of Discovery

Bergisuchus was first discovered by Dr. Dietrich Berg from the German Messel Pit in 1966, who originally classified it as an unnamed new species of sebecosuchian with close affinities to Sebecus , notable for being the first sebecosuchian remains to be recognised outside South America. [1] It was named and described two years later in 1968 by German palaeontologist Oskar Kuhn, who named the binomial Bergisuchus dietrichbergi in honour of Dr. Berg and combined it with the Greek suffix suchos for "crocodile". Bergisuchus is known from the holotype snout and lower jaw (HLMD-Me 7003) from the Messel Pit near Darmstadt first reported in 1966, dated to the Mammal Paleogene zone (MP) 11, and an additional incomplete pair of mandibles (GM XVIII-49) from the Geiseltal open-pit coal mine near Halle in the state of Saxony-Anhalt from the slightly younger MP 12. [2] The holotype is stored in the Hessisches Landesmuseum Darmstadt, while the paratype is housed in the Geiseltal Collection of the Center for Natural Science Collections at the Martin Luther University of Halle-Wittenberg, in Germany. [2] [3]

In 2015, osteoderms from karst deposits dated to the Middle Eocene in Lissieu, France, were tentatively suggested to belong to Bergisuchus sp., or alternatively to Iberosuchus sp. (another European sebecosuchian). This referral was based on the similarity between the osteoderms and those of Baurusuchus and other sebecosuchians, as the two genera are the only known European sebecosuchians. [4]

Description

Bergisuchus is only represented by the holotype snout and lower jaw (HLMD-Me 7003) and the referred pair of lower jaws (GM XVIII-49), so much of its anatomy is unknown. The snout is tall and laterally compressed, unlike the broad flat snouts of modern crocodylians, with tall nasals that form a raised, sharp midline along the length of the snout. Rossmann and colleagues reconstructed the incomplete premaxillae as tall and steep based on the dimensions of the maxilla, more similar in shape to those of Baurusuchus, Barinasuchus and Bretesuchus than to Sebecus. The premaxilla may also have sloped downwards, similar to that of Bretesuchus. [2] Overall, the snout is relatively short and deep compared to the long, low skull of Sebecus. [5] A deep notch is present between the maxilla and premaxilla to house the large lower canine tooth, along with a prominent bulge of bone above each. Based on the shape of the known snout, it's likely that Bergisuchus had separate nostril openings that faced laterally on the surface, as well as laterally facing eyes, unlike modern crocodylians. Also, it shares with Iberosuchus the unusual presence of a small antorbital fenestra, a feature that's invariably absent in both baurusuchids and sebecids. The surface of the maxilla is profusely sculpted with ridges and grooves, a feature that clearly distinguishes it from Iberosuchus. [2] [6]

The lower jaws are mostly only known from incomplete dentaries (as well as part of the splenial), and their surfaces are as similarly strongly sculpted as the upper jaws. The teeth are relatively narrow and serrated (ziphodont), similar to those of predatory theropod dinosaurs and unlike the conical teeth of modern crocodylians. Few teeth themselves are preserved, but they include a very large and prominent serrated 'pseudocanine' that fits into the notch of the upper jaw. This canine is approximately 2 cm tall, and sits on a prominently raised portion of the jaw bone, which arches down in front and behind it and so the rest of the jaw is quite shallow. [6] The rest of the teeth are mostly missing, but the size and shape of the remaining alveoli indicate the size and position of the other teeth in the jaws. The other teeth are around 3 times smaller in diameter than the canines, a unique characteristic of this genus, and are closely packed behind the canine in the lower jaw. Three small teeth are present in front of the canine, the first of which is larger and procumbent, facing up and forwards at the front of the jaw. The maxillary teeth are similarly small, the largest being the third tooth, and the rest of which getting progressively smaller behind it. Premaxillary teeth are unknown, but it possibly had another pair of larger 'pseudocanines' in the upper jaw. It is estimated that there were only 13 teeth in each lower jaw, and at least 13—14 in the upper jaw, a relatively small number compared to other sebecosuchians like Iberosuchus. The teeth are not as strongly compressed as other ziphodont crocodilians, and so may have been stronger and more resistant to stress. [2]

Rossmann and colleagues estimated the total body length of Bergisuchus to be around 1.5 metres (4.9 ft) based on other short-snouted, partially terrestrial crocodylians including Allognathosuchus and the extant dwarf caimans Paleosuchus . Despite its small size, they believed both specimens to represent mature adults, based on both the degree of fusion between the sutures in the skull and the extent of the dermal sculpting on the surface of the bones. As a sebecosuchian, Bergisuchus likely had long limbs that were positioned directly under its body and moved with a parasagittal gait, unlike the sprawling limbs of modern crocodylians. It is unknown if it had a reduced covering of osteoderms like some other sebecosuchians, but no osteoderms were found associated with either individual (although isolated osteoderms have been tentatively referred to this genus). [2] [4]

Classification

Bergisuchus was identified as a sebecosuchian by Dr. Berg when it was first discovered, closely allied to the South American genus Sebecus as "aff. Sebecus? n. sp.". [1] It was later variously assigned to the family Baurusuchidae by Steel in 1973, [7] [8] and also in the now defunct family Trematochampsidae by Buffetaut in 1988. [9] At the time it was unclear what diagnostic traits where unique to each group and which were shared between them, and so the fragmentary Bergisuchus could not be confidently identified and was sometimes placed in incertae sedis for this reason. [10] In 2000, the sebecosuchian affinities of Bergisuchus, particularly to Sebecus, were established by Rossmann and colleagues in a thorough re-examination of the material, and it was assigned to its own monotypic family, the Bergisuchidae, as it was considered too distinct from Sebecus to belong to the same family. [2]

The position of Bergisuchus as a close relative of Sebecus has since been supported by a number of phylogenetic analyses of Mesozoic and Cenozoic mesoeucrocodylians, although the interrelationships of Bergisuchus, Sebecidae and other clades are not settled. The cladograms below are simplified from two recent phylogenetic analyses of mesoeucrocodylians, that of Pol et al. (2014) and of Piacentini Pinheiro et al. (2018), displaying the alternative arrangements of Sebecidae in either Sebecosuchia and Sebecia. [Note 1] The former analysis found Bergisuchus and Iberosuchus as sebecosuchians closely related to but excluded from Sebecidae, while the latter analysis recovers Bergisuchus well within Sebecidae, as per the definitions used in the study:

Palaeobiology

Palaeoecology

Skeleton of the contemporary terrestrial crocodilian Boverisuchus. Boverisuchus magnifrons.jpg
Skeleton of the contemporary terrestrial crocodilian Boverisuchus .

The Messel Pit is famous for its well-preserved fossils, which include semi-aquatic crocodylians such as Asiatosuchus and Diplocynodon . Unlike these crocodylians, Bergisuchus was a small terrestrial hypercarnivore. Rossmann and colleagues speculated extensively about the palaeoecology of Bergisuchus. They suggested that Bergisuchus did not inhabit the fluvial and lacustrine habitats around the Messel and Geiseltal sites. They interpreted the rare, fragmented material as evidence for the bones being transported into these settings from elsewhere, compared to the more abundant and more complete crocodylians from these sites that were well preserved by the local conditions. They further speculated that Bergisuchus may have inhabited drier upland regions from these lakes and rivers, potentially coexisting alongside predatory creodont mammals in the Geiseltal area and avoiding competition with the similarly terrestrial but unrelated planocraniid crocodilians like Boverisuchus . [2]

The reduced ziphodont dentition of Bergisuchus with large teeth concentrated at the front of the jaws is somewhat similar to that of baurusuchids such as Stratiotosuchus , which have reduced their maxillary tooth count down to only 5 teeth. This has been suggested to be a specialisation for hunting large-sized prey items by imparting more powerful bites at the very front of the jaws. This is unlike the condition of most other sebecids, which have more generalised dentitions with similarly sized teeth throughout the jaw, which may have preferred smaller to mid-sized prey. [13] [14] The deeper snout and less compressed, stronger teeth of Bergisuchus also suggest that it was capable of withstanding greater forces relatively than planocraniids, and is inferred to be capable of catching and dismembering prey by thrashing its head in any direction, unlike planocraniids. [2]

The unusual differentiated heterodont teeth were interpreted as evidence for a relatively complex method for processing food, and they suggested that Bergisuchus used the large canines at the front of the jaws as fangs and the smaller rear teeth to "chew" and process food before swallowing. They also suggested that the huge, exaggerated canines could possibly have been used as sexual signals or as weapons in intraspecific competition, rather than predation. [2]

Palaeobiogeography

The presence of Bergisuchus in Europe has been regarded as evidence for a connection between the faunas from South America and Europe in the early Eocene, as other Cenozoic sebecosuchians are mostly known from South America. This link has been supported by the presence of other South American lineages contemporaneous with Bergisuchus in the Messel pit, including purported phorusrhacid birds and herpetotheriid marsupials. [15] Eocene European ecosystems containing sebecosuchians like Bergisuchus have been compared to those of later South America, with predator guilds composed of mammals, phorusrhacids and sebecosuchians, supporting this affinity. [4] Furthermore, Bergisuchus and Iberosuchus do not appear to be closely related to the Cretaceous sebecosuchian Doratodon , which was found across Europe during the Late Cretaceous. This appears to indicate that Bergisuchus and other European sebecosuchians did not descend from Cretaceous European sebecosuchians like Doratodon, but was instead part of a separate invasion of sebecosuchians into Europe during the Palaeogene from South America. [16] This is further supported by the presence of the Algerian Eocene sebecosuchian Eremosuchus in Africa, which has been suggested to be evidence that sebecosuchians reached Europe from South America via Africa, as opposed to through North America where no sebecosuchian remains have been found. [17] [18]

Notes

  1. ↑ Sebecosuchia is cladistically defined as the node containing Sebecidae and Baurusuchidae, assuming a close relationship between the two. Sebecia is a similar concept but for the node containing Sebecidae and Peirosauridae. The validity of each clade depends on the position of Sebecidae within Mesoeucrocodylia, often involving its relationship to Notosuchia.

Related Research Articles

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

Doratodon is an extinct genus of Late Cretaceous crocodylomorph that may have been a member of the Sebecosuchia. Doratodon was a relatively small animal with ziphodont teeth, meaning the teeth had flattened sides and serrated edges. Two species of Doratodon are known to science: D. carcharidens from Austria and Hungary, the type species; and D. ibericus from Spain. Teeth similar to those of Doratodon are also known from Italy and Romania, though they cannot be confidently assigned to this genus.

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

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.

Atacisaurus is an extinct dubious genus of gavialoid crocodylian. Fossils have been found in the Grès de Carcassonne Member of the Sables du Castrais Formation in Laure-Minervois, France that date back to the Middle Eocene.

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

Boverisuchus is an extinct genus of planocraniid crocodyliforms known from the middle Eocene of Germany and western North America. It was a relatively small crocodyliform with an estimated total length of approximately 2.2–3.6 metres (7.2–11.8 ft).

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

Iberosuchus is a genus of extinct sebecosuchian mesoeucrocodylian found in Western Europe from the Eocene. Remains from Portugal was described in 1975 by Antunes as a sebecosuchian crocodilian. This genus has one species: I. macrodon. Iberosuchus was a carnivore, unlike the crocodilians today, they are not aquatic and are instead terrestrial.

<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">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>Barinasuchus</i> Extinct genus of reptiles

Barinasuchus is an extinct genus of sebecid mesoeucrocodylian. Its fossils have been found in middle Eocene-age rocks of the Divisadero Largo Formation of Argentina, middle Miocene-age rocks of the Ipururo Formation of Peru, and middle Miocene-age rocks of the Parángula Formation of Venezuela.

<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.

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>Bretesuchus</i> Extinct genus of reptiles

Bretesuchus is an extinct genus of sebecosuchian mesoeucrocodylian within the family Sebecidae known from northwestern Argentina. It was a large apex predator.

Eremosuchus is an extinct genus of sebecosuchian mesoeucrocodylian. Fossils have been found from El Kohol, Algeria of Eocene age. It had serrated, ziphodont teeth.

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>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>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.

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

Planocraniidae is an extinct family of eusuchian crocodyliforms known from the Paleogene of Asia, Europe and North America. The family was coined by Li in 1976, and contains three genera, Boverisuchus, Duerosuchus and Planocrania. Planocraniids were highly specialized crocodyliforms that were adapted to living on land. They had extensive body armor, long legs, and blunt claws resembling hooves, and are sometimes informally called "hoofed crocodiles".

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

Sahitisuchus is an extinct genus of sebecid mesoeucrocodylian known from Rio de Janeiro State of southeastern Brazil. It contains a single species, Sahitisuchus fluminensis. It is a terrestrial sebecid, however also adopted to a semi-aquatic lifestyle to some degree, most probably coexisting with the semi-aquatic alligatorid Eocaiman itaboraiensis.

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

Dentaneosuchus is a genus of large bodied sebecid crocodylomorph from the Middle Eocene of Issel and Réalmont (France). Originally described as Atacisaurus crassiproratus, the discovery of additional remains led to it being placed in a separate genus in 2023. It was tentatively recovered as the basalmost member of the family Sebecidae. Because of this Dentaneosuchus could play an important part in deciphering the origins and dispersal of European sebecids, as their presence on the continent, far away from their primary range in South America, is still not entirely resolved. It reached a similar size to the enormous Barinasuchus, making it not only one of the biggest sebecids but also the biggest terrestrial carnivore of Cenozoic Europe. Dentaneosuchus would have been an apex predator of its environment, capable of taking large prey such as Lophiodon. However, for as of yet unknown reasons crocodylomorphs would lose their spot as top predator in this part of the world by the end of the Eocene, with Dentaneosuchus representing one of the last members of its group in Europe.

References

  1. 1 2 Martin, Jeremy (2016). "New material of the ziphodont mesoeucrocodylian Iberosuchus from the Eocene of Languedoc, southern France" (PDF). Annales de Paléontologie. 102 (2): 135–144. Bibcode:2016AnPal.102..135M. doi:10.1016/j.annpal.2016.05.002.
  2. 1 2 3 4 5 6 7 8 9 10 Rossmann, T.; Rauhe, M.; Ortega, F. (2000). "Studies on Cenozoic crocodiles: 8. Bergisuchus dietrichbergi Kuhn (Sebecosuchia: Bergisuchidae n. fam.) from the Middle Eocene of Germany, some new systematic and biological conclusions". Paläontologische Zeitschrift. 74 (3): 379–392. Bibcode:2000PalZ...74..379R. doi:10.1007/BF02988108. S2CID   129507618.
  3. ↑ Steinheimer, Frank D.; Hastings, Alexander K. (2018). Beck L.; Joger U. (eds.). HALLE: The Geiseltal Collection of Martin Luther University, Halle-Wittenberg. Paleontological Collections of Germany, Austria and Switzerland. Natural History Collections. Springer, Cham. pp. 271–280. doi:10.1007/978-3-319-77401-5_25. ISBN   9783319774008.
  4. 1 2 3 Martin, Jeremy (2015). "A sebecosuchian in a middle Eocene karst with comments on the dorsal shield in Crocodylomorpha". Acta Palaeontologica Polonica. 3 (60): 673–680.
  5. ↑ Molnar, R. (2010). "A new reconstruction of the skull of Sebecus icaeorhinus (Crocodyliformes: Sebecosuchia) from the Eocene of Argentina". Brazilian Geographical Journal: Geosciences and Humanities Research Medium. 1 (2). Uberlândia: 314–330. CiteSeerX   10.1.1.300.8530 .
  6. 1 2 Ortega, F.; Buscaloni, A.D; Gasaparini, Z. (1996). "Reinterpretation and new denomination of Atacisaurus crassiproratus (Middle Eocene; Issel, France) as cf. Iberosuchus (Crocodylomorpha, Metasuchia)". Geobios. 29 (3): 353–364. Bibcode:1996Geobi..29..353O. doi:10.1016/S0016-6995(96)80037-4.
  7. ↑ Steel, R. (1973). Kuhn, O.; Wellnhofer, P. (eds.). Crocodylia. Handbuch der Paläoherpetologie (in German). Vol. 16. Fischer. ISBN   978-3437301414.
  8. ↑ Carroll, Robert L. (1988). Vertebrate Paleontology and Evolution. W. H. Freeman and Company. p.  620. ISBN   978-0716718222. OCLC   922750908.
  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. ↑ Gasparini, Z. (1984). "New Tertiary Sebecosuchia (Crocodylia: Mesosuchia) from Argentina". Journal of Vertebrate Paleontology . 4 (1): 85–95. Bibcode:1984JVPal...4...85G. doi:10.1080/02724634.1984.10011988.
  11. ↑ Pol, Diego; Nascimento, Paulo M.; Carvalho, Alberto B.; Riccomini, Claudio; Pires-Domingues, Ricardo A.; Zaher, Hussam (2014). "A New Notosuchian from the Late Cretaceous of Brazil and the Phylogeny of Advanced Notosuchians". PLoS ONE . 9 (4): e93105. Bibcode:2014PLoSO...993105P. doi: 10.1371/journal.pone.0093105 . PMC   3973723 . PMID   24695105.
  12. ↑ Piacentini Pinheiro, A.E.; Pereira, P.V.L.G.dC.; de Souza, R.G.; Brum, A.S.; Lopes, R.T.; Machado, A.S.; Bergqvist, L.P.; Simbras, F.M. (2018). "Reassessment of the enigmatic crocodyliform "Goniopholis" paulistanus Roxo, 1936: Historical approach, systematic, and description by new materials". PLoS ONE . 13 (8): e0199984. Bibcode:2018PLoSO..1399984P. doi: 10.1371/journal.pone.0199984 . PMC   6070184 . PMID   30067779.
  13. ↑ Leardi, J. M.; Pol, D.; Gasparini, Z. (2018). "New Patagonian baurusuchids (Crocodylomorpha; Notosuchia) from the Bajo de la Carpa Formation (Upper Cretaceous; Neuquén, Argentina): New evidences of the early sebecosuchian diversification in Gondwana". Comptes Rendus Palevol. 7 (8): 504–521. Bibcode:2018CRPal..17..504L. doi: 10.1016/j.crpv.2018.02.002 .
  14. ↑ Pinheiro, A. E. P.; Bertini, R. J.; Andrade, M. D.; Neto, R. M. (2008). "A new specimen of Stratiotosuchus maxhechti (Baurusuchidae, Crocodyliformes) from the Adamantina Formation (Upper Cretaceous), Southeastern Brazil". Revista Brasileira de Paleontologia. 11 (1): 37–50. CiteSeerX   10.1.1.734.6541 . doi:10.4072/rbp.2008.1.04.
  15. ↑ Agusti, Jordi; Antón, Mauricio (2002). "The Eocene: Reaching the Climax". Mammoths, Sabertooths, and Hominids: 65 Million Years of Mammalian Evolution in Europe. Columbia University Press. pp. 48–50. ISBN   978-0-231-11640-4. OCLC   61129178.
  16. ↑ Rabi, M.; Sebők, N. (2015). "A revised Eurogondwana model: Late Cretaceous notosuchian crocodyliforms and other vertebrate taxa suggest the retention of episodic faunal links between Europe and Gondwana during most of the Cretaceous". Gondwana Research . 28 (3): 1197–1211. Bibcode:2015GondR..28.1197R. doi:10.1016/j.gr.2014.09.015.
  17. ↑ Buffetaut, E. (1982). "A ziphodont mesosuchian crocodile from the Eocene of Algeria and its implications for vertebrate dispersal". Nature . 300 (5888): 176–178. Bibcode:1982Natur.300..176B. doi:10.1038/300176a0. S2CID   4323999.
  18. ↑ Buffetaut, E. (1989). "A new ziphodont mesosuchian crocodile from the Eocene of Algeria". Palaeontographica. 208: 1–10.
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