''Ikanogavialis'' is an extinct

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 ...

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 ...

crocodilia 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 ...

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 ...

s have been found in the

Urumaco Formation 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. Location Th ...

Urumaco Urumaco is a town in Falcón State in Venezuela. It is of interest to paleontologists due to its rich fossil history. The arid climate of the region means that the fossils are not hidden by vegetation. The fossils were first made known to scienc ...

Venezuela Venezuela (; ), officially the Bolivarian Republic of Venezuela ( es, link=no, República Bolivariana de Venezuela), is a country on the northern coast of South America, consisting of a continental landmass and many islands and islets in th ...

and the

Solimões Formation The Pebas Formation is a lithostratigraphic unit of Miocene age, found in western Amazonia. The formation extends over , including parts of Brazil, Peru, Ecuador and Colombia.Wesselingh et al., 2006 It is interpreted as representing the deposits ...

Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...

. The strata from which remains are found are late

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 ...

in age, rather than

Pliocene The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58Holocene The Holocene ( ) is the current geological epoch. It began approximately 11,650 cal years Before Present (), after the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene togethe ...

on Muyua or

Woodlark Island Woodlark Island, known to its inhabitants simply as Woodlark or Muyua, is the main island of the Woodlark Islands archipelago, located in Milne Bay Province and the Solomon Sea, Papua New Guinea. Although no formal census has been conducted sinc ...

Papua New Guinea Papua New Guinea (abbreviated PNG; , ; tpi, Papua Niugini; ho, Papua Niu Gini), officially the Independent State of Papua New Guinea ( tpi, Independen Stet bilong Papua Niugini; ho, Independen Stet bilong Papua Niu Gini), is a country i ...

Description

''Ikanogavialis'' had a dorsoventrally deep snout and a distinctive notch between the

dentary 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

maxilla The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The t ...

ry alveoli. The external nares projected anterodorsally from the rostrum. This can be seen as a

plesiomorphic In phylogenetics, a plesiomorphy ("near form") and symplesiomorphy are synonyms for an ancestral character shared by all members of a clade, which does not distinguish the clade from other clades. Plesiomorphy, symplesiomorphy, apomorphy, and ...

characteristic in crocodilians, but given that the earliest gavialoids possessed dorsally projecting external nares, this feature can be seen as having been a reversal from the gavialoid

apomorphy In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to have ...

back to the crocodilian plesiomorphy rather than having been directly obtained from an early crocodilian ancestor.

Species

The

type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...

of ''Ikanogavialis'' is ''I. gameroi''. It was named in 1970 from material found from the

Urumaco formation 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. Location Th ...

, South America. Additionally, a

Pleistocene The Pleistocene ( , often referred to as the ''Ice age'') is the geological Epoch (geology), epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fina ...

gavialoid found on

was initially named ''Gavialis papuensis'' in 1905, but has since been proposed to be a member of ''Ikanogavialis'', although the poor quality of the preserved material makes it difficult to determine.

Paleobiology

''Ikanogavialis'' may have lived in a coastal paleoenvironment along with other gavialids such as ''

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 ...

''. The strata of the Urumaco formation were deposited in both marine and fluvial settings, although it is unclear to which portion both genera belong. Other gavialoids such as ''

Siquisiquesuchus ''Siquisiquesuchus'' (meaning " Siquisique crocodile" after the town in Lara, Venezuela, near where the first described specimens were found) is an extinct genus of gavialid crocodilian. It is known from cranial remains and a few postcranial bon ...

'' and ''

Piscogavialis ''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 Bas ...

'' are known to have lived in coastal environments, and it is likely that extant freshwater

gavialoids 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 ...

such as ''

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 ...

'' may have originated from these coastal forms. ''Ikanogavialis'' also existed with many other crocodilians in Venezuela during the late Miocene including the giant

caiman A caiman (also cayman as a variant spelling) is an alligatorid belonging to the subfamily Caimaninae, one of two primary lineages within the Alligatoridae family, the other being alligators. Caimans inhabit Mexico, Central and South America fro ...

Purussaurus ''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 ...

'' and an extinct species of '' Melanosuchus''. "''Gavialis''" / ''Ikanogavialis papuensis'' was similarly fully marine, having been found in association with sea turtles and

sirenia The Sirenia (), commonly referred to as sea-cows or sirenians, are an order of fully aquatic, herbivorous mammals that inhabit swamps, rivers, estuaries, marine wetlands, and coastal marine waters. The Sirenia currently comprise two distinct f ...

ns. It represents the youngest fully marine crocodilian to date.

Phylogeny

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 ''Aktiogavialis'' is an extinct genus of crocodylian that lived from the Oligocene until the Miocene in what is now the Caribbean. Two species have been described: ''Aktiogavialis puertoricensis'' from the Middle Oligocene of Puerto Rico and ''Ak ...

'', ''

'', ''Ikanogavialis'', ''

'', and ''

''. 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

: 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

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 ''

'' 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:

Notes and references

External links

''Ikanogavialis''
in the

Paleobiology Database The Paleobiology Database is an online resource for information on the distribution and classification of fossil animals, plants, and microorganisms. History The Paleobiology Database (PBDB) originated in the NCEAS-funded Phanerozoic Marine Pale ...

{{Taxonbar, from=Q5995365 Gavialidae Prehistoric pseudosuchian genera Neogene crocodylomorphs Miocene reptiles of South America Montehermosan Huayquerian Chasicoan Mayoan Neogene Brazil Fossils of Brazil Neogene Venezuela Fossils of Venezuela Prehistoric marine reptiles Holocene extinctions Fossil taxa described in 1970 Pleistocene crocodylomorphs