Piscogavialis

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Piscogavialis
Temporal range: Late Miocene,
11.6–5.3  Ma [1]
Piscogavialis 1.jpg
Skull
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Order: Crocodilia
Family: Gavialidae
Subfamily: Gryposuchinae
Genus: Piscogavialis
Kraus 1998
Type species
Piscogavialis jugaliperforatus

Piscogavialis is an extinct monospecific genus of gryposuchine gavialid crocodylian. The only species yet known is P. jugaliperforatus. Fossils of Piscogavialis have been found from the Mio-Pliocene Pisco Formation of the Sacaco Basin in southern Peru in 1998, [2] where it coexisted with the much smaller gavialid Sacacosuchus . [3]

Contents

Piscogavialis is known only from a single specimen, but it represents some of the best preserved gavialid material known from South America. The skull is preserved in three dimensions and is nearly complete. A mandible and some postcranial material have also been found in association with the skull. Several important features of the occipital region of the skull support a referral to the family Gavialidae, which also includes the extant gharial and false gharial.

Paleobiology

The strata from which remains of Piscogavialis have been found suggest that it lived in a coastal environment. [4] [5] Another extinct marine gavialid, Sacacosuchus , was described in 2022 and discovered in the same formation. Sacacosuchus was smaller, estimated to be up to 4.32 m (14.2 ft) long, whereas Piscogavialis was nearly double the size. [3]

During the Miocene, what is now the Pisco Basin in Peru was largely covered by a shallow marine transgression that connected to the open ocean. The environment was marked by shallow waters, protected bays with rocky shores and small islands. The south-east Pacific coast was inhabited by at least two species of crocodilians, Piscogavialis and Sacacosuchus, throughout most of this time period. The remains of both species were found alongside other animals typical for the marine environments of this region including various cetaceans, seals, seabirds, marine sloths and sharks. As the Peruvian sediments yielded bones of both adults and juvenile specimens, it is assumed that Sacacosuchus spent its entire life in saltwater. Although Piscogavialis was notably larger, it was also likely more specialized, feeding primarily on fast fish. Sacacosuchus, meanwhile, in spite of being only half the length of its contemporary relative, was adapted to a more generalist diet. [3]

During the Miocene, environmental factors worked together to highly favor marine gavialoids, in particular warm temperatures and abundant shallow water coastal environments suited to their lifestyle. Subsequently, their diversity was high during most of this time period with their range extending as far south as Chile. However, despite these initially favorable conditions, global temperatures continuously dropped throughout the late Neogene. Initially, both Piscogavialis and Sacacosuchus weren't heavily affected, with the water temperatures of their habitat continuing to support these ectotherms until at least the end of the Miocene period. Eventually, however, the extinction of Peru's marine gavialids was brought on by the disruption of these coastal ecosystems, caused by dropping sea levels and the uplifting of the Andes Mountains, coupled with the more drastically dropping global temperatures of the Pliocene. [3]

Phylogeny

A phylogenetic analysis conducted in a 2007 study found Gryposuchinae to include the genera Aktiogavialis , Gryposuchus , Ikanogavialis , Piscogavialis, and Siquisiquesuchus . Below is a cladogram from the 2007 analysis showing the phylogenetic relationships of gryposuchines among gavialoids: [5]

Gavialoidea  

Eothoracosaurus

Thoracosaurus

Eosuchus

Argochampsa

Gavialidae  

Eogavialis

Gavialis

Gryposuchinae  

Ikanogavialis

Piscogavialis

Siquisiquesuchus

Gryposuchus

Aktiogavialis

Alternatively, a 2018 tip dating study by Lee & Yates simultaneously using morphological, molecular (DNA sequencing), and stratigraphic (fossil age) data indicated that the members of Gryposuchinae may in fact be paraphyletic and rather an evolutionary grade towards Gavialis and the gharial, as shown in the cladogram below: [6]

Gavialidae

Gavialis gangeticus Gharial

Gavialis bengawanicus

Gavialis browni

Gryposuchus colombianus

Ikanogavialis

Gryposuchus pachakamue

Piscogavialis

Harpacochampsa

Toyotamaphimeia

Penghusuchus

Gavialosuchus

Tomistoma lusitanicum

Tomistoma schlegelii False gharial

Gryposuchinae

The below cladogram is from the 2022 Salas-Gismondi et al. study describing the newly discovered co-existing Sacacosuchus , and shows Piscogavialis as a more derived member of Gavialidae: [3]

Crocodyloidea

Gavialoidea

Kentisuchus spenceri

Dollosuchoides densmorei

Maroccosuchus zennaroi

Maomingosuchus petrolica

Gavialidae

Tomistoma schlegelii, false gharial

Tomistoma lusitanicum

Gavialosuchus eggenburgensis

Sacacosuchus cordovai

Thecachampsa antiqua

Thecachampsa carolinensis

Thecachampsa americana

Tomistoma dowsoni

Tomistoma coppensi

Tomistoma cairense

Paratomistoma courti

MUSM 1513

Argochampsa krebsi

Aktiogavialis caribesi

Aktiogavialis puertoricensis

Eogavialis

Gavialis bengawanicus

Gavialis gangeticus, gharial

Siwalik Gavialis

Siquisiquesuchus venezuelensis

Piscogavialis jugaliperforatus

Ikanogavialis gameroi

Dadagavialis gunai

Gryposuchus pachakamue

Gryposuchus neogaeus

Gryposuchus croizati

Gryposuchus colombianus

Related Research Articles

<span class="mw-page-title-main">Gavialinae</span> Subfamily of gharial crocodylians

Gavialinae is a subfamily of large semiaquatic crocodilian reptiles, resembling crocodiles, but with much thinner snouts. Gavialinae is one of the two major subfamilies within the family Gavialidae - the other being the subfamily Tomistominae, which contains the false gharial and extinct relatives.

<span class="mw-page-title-main">Gavialidae</span> Family of gharial crocodylians

Gavialidae is a family of large semiaquatic crocodilians with elongated, narrow snouts. Gavialidae consists of two living species, the gharial and the false gharial, both occurring in Asia. Many extinct members are known from a broader range, including the recently extinct Hanyusuchus. Gavialids are generally regarded as lacking the jaw strength to capture the large mammalian prey favoured by crocodiles and alligators of similar size so their thin snout is best used to catch fish, however the false gharial has been found to have a generalist diet with mature adults preying upon larger vertebrates, such as ungulates.

<i>Tomistoma</i> Genus of crocodilians

Tomistoma is a genus of gavialid crocodilians. They are noted for their long narrow snouts used to catch fish, similar to the gharial. Tomistoma contains one extant (living) member, the false gharial, as well as potentially several extinct species: T. cairense, T. lusitanicumT. coppensi, and T. dowsoni. However, these species may need to be reclassified to different genera as studies have shown them to be paraphyletic, for example: previously assigned species T. taiwanicus from Taiwan, is reclassified to the genus Toyotamaphimeia, and T. dowsoni should be excluded from Tomistoma based on phylogenetic analysis.

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

Gavialosuchus is an extinct genus of gavialoid crocodylian from the early Miocene of Europe. Currently only one species is recognized, as a few other species of Gavialosuchus have since been reclassified to other genera.

Argochampsa is an extinct genus of eusuchian crocodylomorph, usually regarded as a gavialoid crocodilian, related to modern gharials. It lived in the Paleocene of Morocco. Described by Hua and Jouve in 2004, the type species is A. krebsi, with the species named for Bernard Krebs. Argochampsa had a long narrow snout, and appears to have been marine in habits.

Dollosuchoides, colloquially known as the Crocodile of Maransart, is an extinct monospecific genus of gavialoid crocodilian, traditionally regarded as a member of the subfamily Tomistominae. Fossils have been found in the Brussel Formation of Maransart, Belgium and date back to the middle Eocene.

Eogavialis is an extinct genus of eusuchian crocodylomorph, usually regarded as a gavialoid crocodylian. It superficially resembles Tomistoma schlegelii, the extant false gharial, and consequently material from the genus was originally referred to Tomistoma. Indeed, it was not until 1982 that the name Eogavialis was constructed after it was realised that the specimens were from a more basal form.

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

Eothoracosaurus is an extinct monospecific genus of eusuchian crocodylomorphs found in Eastern United States which existed during the Late Cretaceous period. Eothoracosaurus is considered to belong to an informally named clade called the "thoracosaurs", named after the closely related Thoracosaurus. Thoracosaurs in general were traditionally thought to be related to the modern false gharial, largely because the nasal bones contact the premaxillae, but phylogenetic work starting in the 1990s instead supported affinities within gavialoid exclusive of such forms. Even more recent phylogenetic studies suggest that thoracosaurs might instead be non-crocodilian eusuchians.

<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.

Maroccosuchus zennaroi is an extinct gavialoid crocodylian from the Early Eocene of Morocco, traditionally regarded as a member of the subfamily Tomistominae.

Paratomistoma is an extinct monospecific genus of gavialoid crocodylian. It is based on the holotype specimen CGM 42188, a partial posterior skull and lower jaw discovered at Wadi Hitan, Egypt, in Middle Eocene-age rocks of the Gehannam Formation. The skull is unfused but considered morphologically mature. Paratomistoma was named in 2000 by Christopher Brochu and Philip Gingerich; the type species is P. courti in honor of Nicholas Court, who found CGM 42188. They performed a phylogenetic analysis and found Paratomistoma to be a derived member of Tomistominae, related to the false gharial. It may have been a marine or coastal crocodilian.

Siquisiquesuchus is an extinct genus of gavialid crocodilian. It is known from cranial remains and a few postcranial bones found in Miocene-age rocks of the Castillo Formation in northwestern Venezuela.

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

Penghusuchus is an extinct genus of gavialid crocodylian. It is known from a skeleton found in Middle to Upper Miocene rocks of Penghu Island, off Taiwan. The taxon was described in 2009 by Shan and colleagues; the type species is P. pani. It may be related to two other fossil Asian gavialids: Toyotamaphimeia machikanensis of Japan and Hanyusuchus sinensis of South China. It was a medium-sized gavialid with an estimated total length of 4.5 metres (15 ft).

Gryposuchinae is an extinct subfamily of gavialid crocodylians. Gryposuchines lived mainly in the Miocene of South America. However, "Ikanogavialis" papuensis may have survived more recently, into the Late Pleistocene/Holocene. Most were long-snouted coastal forms. The group was named in 2007 and includes genera such as Gryposuchus and Aktiogavialis, although a 2018 study indicates that the group might be paraphyletic and rather an evolutionary grade towards the gharial.

<span class="mw-page-title-main">Gavialoidea</span> Superfamily of large reptiles

Gavialoidea is one of three superfamilies of crocodylians, the other two being Alligatoroidea and Crocodyloidea. Although many extinct species are known, only the gharial Gavialis gangeticus and the false gharial Tomistoma schlegelii are alive today, with Hanyusuchus having become extinct in the last few centuries.

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

Tomistominae is a subfamily of crocodylians that includes one living species, the false gharial. Many more extinct species are known, extending the range of the subfamily back to the Eocene epoch. In contrast to the false gharial, which is a freshwater species that lives only in southeast Asia, extinct tomistomines had a global distribution and lived in estuaries and along coastlines.

Tomistoma cairense is an extinct species of gavialoid crocodilian from the Lutetian stage of the Eocene era. It lived in North East Africa, especially Egypt. Remains of T. cairense have been found in the Mokattam Formation, in Mokattam, Egypt. Tomistoma cairense did not have a Maxilla process within their lacrimal gland, whereas all extant (living) crocodilians do.

Dadagavialis is an extinct monospecific genus of gavialid crocodylian that lived during the Early Miocene in what is now Panama. It was described in 2018, and was proposed to be a member of Gryposuchinae. However, other studies have shown Gryposuchinae to be paraphyletic and rather an evolutionary grade towards the living gharial, and thus Dadagavialis might just be classified as a member of Gavialidae.


Sacacosuchus is an extinct monospecific genus of marine gavialid that lived along the coast of the south-east Pacific from approximately 19 to 6.3 million years ago. Its fossils have been found in the Chilcatay and Pisco Formations of Peru, where it coexisted with the much larger Piscogavialis. Based on its skull, Sacacosuchus was most likely a generalist feeder with an estimated total body length of 4.32 m (14.2 ft). Its extinction is thought to have been caused by a combination of factors including falling sea levels and global cooling.

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. Piscogavialis at Fossilworks.org
  3. 1 2 3 4 5 Salas-Gismondi, R.; Ochoa, D.; Jouve, S.; Romero, P.E.; Cardich, J.; Perez, A.; DeVries, T.; Baby, P.; Urbina, M.; Carré, M. (2022-05-11). "Miocene fossils from the southeastern Pacific shed light on the last radiation of marine crocodylians". Proceedings of the Royal Society B. 289. doi:10.1098/rspb.2022.0380. PMC   9091840 .
  4. Kraus, R. M. (1998). "The cranium of Piscogavialis jugaliperforatus n.gen., n.sp. (Gavialidae, Crocodylia) from the Miocene of Peru". Paläontologische Zeitschrift. 72 (3): 389–406. doi:10.1007/bf02988368. S2CID   84214781.
  5. 1 2 Vélez-Juarbe, J.; Brochu, C. A.; Santos, H. (2007). "A gharial from the Oligocene of Puerto Rico: transoceanic dispersal in the history of a non-marine reptile". Proceedings of the Royal Society B. 274 (1615): 1245–1254. doi:10.1098/rspb.2006.0455. PMC   2176176 . PMID   17341454.
  6. Michael S. Y. Lee; Adam M. Yates (27 June 2018). "Tip-dating and homoplasy: reconciling the shallow molecular divergences of modern gharials with their long fossil". Proceedings of the Royal Society B . 285 (1881). doi: 10.1098/rspb.2018.1071 . PMC   6030529 . PMID   30051855.