Piscogavialis
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''Piscogavialis'' is an
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
monospecific In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unispec ...
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of gryposuchine
gavialid Gavialidae is a family of large semiaquatic crocodilians with elongated, narrow snouts. Gavialidae consists of two living species, the gharial (''Gavialis gangeticus'') and the false gharial (''Tomistoma schlegelii''), both occurring in Asia. Man ...
crocodylia Crocodilia (or Crocodylia, both ) is an order of mostly large, predatory, semiaquatic reptiles, known as crocodilians. They first appeared 95 million years ago in the Late Cretaceous period (Cenomanian stage) and are the closest living ...
n. The only species yet known is ''P. jugaliperforatus''. Fossils of ''Piscogavialis'' have been found from the
Mio Mio or MIO may refer to: shortened form of Mioritic Shepherd dog or Mioritic sheepdog; a Romanian mountain dog. Places * Mio, Michigan, a town in the US Music * ''Mío'', 2011 album by David Bustamante * "Mío", 1992 song by Paulina Rubio Bra ...
-
Pliocene The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58Pisco Formation The Pisco Formation is a geologic formation located in Peru, on the southern coastal desert of Ica, Peru, Ica and Arequipa. The approximately thick formation was deposited in the Pisco Basin, spanning an age from the Miocene, Middle Miocene up to ...
of the Sacaco Basin in southern
Peru , image_flag = Flag of Peru.svg , image_coat = Escudo nacional del Perú.svg , other_symbol = Great Seal of the State , other_symbol_type = Seal (emblem), National seal , national_motto = "Fi ...
in
1998 1998 was designated as the ''International Year of the Ocean''. Events January * January 6 – The '' Lunar Prospector'' spacecraft is launched into orbit around the Moon, and later finds evidence for frozen water, in soil in permanently ...
,''Piscogavialis''
at
Fossilworks Fossilworks is a portal which provides query, download, and analysis tools to facilitate access to the Paleobiology Database The Paleobiology Database is an online resource for information on the distribution and classification of fossil animals ...
.org
where it coexisted with the much smaller gavialid '' Sacacosuchus''. ''Piscogavialis'' is known only from a single specimen, but it represents some of the best preserved gavialid material known from
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...
. The skull is preserved in three dimensions and is nearly complete. A
mandible In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
and some
postcranial Postcrania (postcranium, adjective: postcranial) in zoology and vertebrate paleontology is all or part of the skeleton apart from the skull. Frequently, fossil remains, e.g. of dinosaurs or other extinct tetrapods, consist of partial or isolated sk ...
material have also been found in association with the skull. Several important features of the
occipital The occipital bone () is a cranial dermal bone and the main bone of the occiput (back and lower part of the skull). It is trapezoidal in shape and curved on itself like a shallow dish. The occipital bone overlies the occipital lobes of the cereb ...
region of the skull support a referral to the
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
Gavialidae Gavialidae is a family (biology), family of large semiaquatic crocodilians with elongated, narrow snouts. Gavialidae consists of two extant taxon, living species, the gharial (''Gavialis gangeticus'') and the false gharial (''Tomistoma schlegelii ...
, which also includes the
extant Extant is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Extant taxon, a taxon which is not extinct, ...
gharial The gharial (''Gavialis gangeticus''), also known as gavial or fish-eating crocodile, is a crocodilian in the family Gavialidae and among the longest of all living crocodilians. Mature females are long, and males . Adult males have a distinct b ...
and
false gharial The false gharial (''Tomistoma schlegelii''), also known by the names Malayan gharial, Sunda gharial and tomistoma is a freshwater crocodilian of the family Gavialidae native to Peninsular Malaysia, Borneo, Sumatra and Java. It is listed as Vu ...
.


Paleobiology

The strata from which remains of ''Piscogavialis'' have been found suggest that it lived in a coastal environment. Another extinct marine gavialid, '' Sacacosuchus'', was described in 2022 and discovered in the same formation. '' Sacacosuchus'' was smaller, estimated to be up to long, whereas ''Piscogavialis'' was nearly double the size. During the
Miocene The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recen ...
, what is now the
Pisco Basin Pisco Basin ( es, Cuenca de Pisco) is a sedimentary basin extending over in southwestern Peru.Solís Mundaca, 2018, p.1 The basin has a thick sedimentary fill, which is about half the thickness of more northern foreland basins in Peru. The old ...
in
Peru , image_flag = Flag of Peru.svg , image_coat = Escudo nacional del Perú.svg , other_symbol = Great Seal of the State , other_symbol_type = Seal (emblem), National seal , national_motto = "Fi ...
was largely covered by a shallow
marine transgression A marine transgression is a geologic event during which sea level rises relative to the land and the shoreline moves toward higher ground, which results in flooding. Transgressions can be caused by the land sinking or by the ocean basins filling ...
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
cetacea Cetacea (; , ) is an infraorder of aquatic mammals that includes whales, dolphins, and porpoises. Key characteristics are their fully aquatic lifestyle, streamlined body shape, often large size and exclusively carnivorous diet. They propel them ...
ns, 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. 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 Chile, officially the Republic of Chile, is a country in the western part of South America. It is the southernmost country in the world, and the closest to Antarctica, occupying a long and narrow strip of land between the Andes to the east a ...
. However, despite these initially favorable conditions, global temperatures continuously dropped throughout the late
Neogene The Neogene ( ), informally Upper Tertiary or Late Tertiary, is a geologic period and system that spans 20.45 million years from the end of the Paleogene Period million years ago ( Mya) to the beginning of the present Quaternary Period Mya. ...
. Initially, both ''Piscogavialis'' and ''Sacacosuchus'' weren't heavily affected, with the water temperatures of their habitat continuing to support these
ectotherm An ectotherm (from the Greek () "outside" and () "heat") is an organism in which internal physiological sources of heat are of relatively small or of quite negligible importance in controlling body temperature.Davenport, John. Animal Life a ...
s 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 The Andes, Andes Mountains or Andean Mountains (; ) are the List of mountain ranges#Mountain ranges by length, longest continental mountain range in the world, forming a continuous highland along the western edge of South America. The range i ...
, coupled with the more drastically dropping global temperatures of the
Pliocene The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58


Phylogeny

A
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
analysis conducted in a 2007 study found
Gryposuchinae Gryposuchinae is an extinct subfamily (biology), subfamily of Gavialidae, gavialid crocodylians. Gryposuchines lived mainly in the Miocene of South America. However, ''Ikanogavialis, "Ikanogavialis" papuensis'' may have survived more recently, in ...
to include the genera '' Aktiogavialis'', ''
Gryposuchus ''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 (Colhuehuapian to Huayquerian). One recently d ...
'', ''
Ikanogavialis ''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 t ...
'', ''Piscogavialis'', and '' Siquisiquesuchus''. Below is a
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...
from the 2007 analysis showing the phylogenetic relationships of gryposuchines among gavialoids: Alternatively, a 2018
tip dating Tip dating is a technique used in molecular dating that allows the inference of time-calibrated phylogenetic trees. Its defining feature is that it uses the ages of the samples to provide time information for the analysis, in contrast with traditio ...
study by Lee & Yates simultaneously using morphological, molecular (
DNA sequencing DNA sequencing is the process of determining the nucleic acid sequence – the order of nucleotides in DNA. It includes any method or technology that is used to determine the order of the four bases: adenine, guanine, cytosine, and thymine. Th ...
), and
stratigraphic Stratigraphy is a branch of geology concerned with the study of rock layers (strata) and layering (stratification). It is primarily used in the study of sedimentary and layered volcanic rocks. Stratigraphy has three related subfields: lithostrati ...
(
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
age) data indicated that the members of
Gryposuchinae Gryposuchinae is an extinct subfamily (biology), subfamily of Gavialidae, gavialid crocodylians. Gryposuchines lived mainly in the Miocene of South America. However, ''Ikanogavialis, "Ikanogavialis" papuensis'' may have survived more recently, in ...
may in fact be
paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...
and rather an
evolutionary grade A grade is a taxon united by a level of morphological or physiological complexity. The term was coined by British biologist Julian Huxley, to contrast with clade, a strictly phylogenetic unit. Definition An evolutionary grade is a group of ...
towards ''
Gavialis ''Gavialis'' is a genus of crocodylians that includes the living gharial ''Gavialis gangeticus'' and one known extinct species, '' Gavialis bengawanicus.'' ''G. gangeticus'' comes from the Indian Subcontinent, while ''G. bengawanicus'' is known ...
'' and the
gharial The gharial (''Gavialis gangeticus''), also known as gavial or fish-eating crocodile, is a crocodilian in the family Gavialidae and among the longest of all living crocodilians. Mature females are long, and males . Adult males have a distinct b ...
, as shown in the cladogram below: 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 Gavialidae is a family (biology), family of large semiaquatic crocodilians with elongated, narrow snouts. Gavialidae consists of two extant taxon, living species, the gharial (''Gavialis gangeticus'') and the false gharial (''Tomistoma schlegelii ...
:


References

{{Taxonbar, from=Q7198259, from2=Q112845633 Gavialidae Prehistoric pseudosuchian genera Miocene crocodylomorphs Miocene reptiles of South America Montehermosan Huayquerian Neogene Peru Fossils of Peru Fossil taxa described in 1998