Chalawan (genus)
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''Chalawan'' (from ) is an
extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
pholidosaurid Pholidosauridae is an extinct Family (biology), family of aquatic neosuchian mesoeucrocodylian crocodylomorphs. Fossils have been found in Europe (Denmark, England, France, Germany, Spain and Sweden), Africa (Algeria, Niger, Mali, Morocco and Tu ...
mesoeucrocodylian known from the
Late Jurassic The Late Jurassic is the third Epoch (geology), epoch of the Jurassic Period, and it spans the geologic time scale, geologic time from 161.5 ± 1.0 to 143.1 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic stratum, strata.Owen ...
or
Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name) is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 143.1 ...
Phu Kradung Formation of Nong Bua Lamphu Province,
northeastern Thailand Northeast Thailand or Isan (Isan/, ; ; also written as Isaan, Isarn, Issarn, Issan, Esan, or Esarn; from Pāli ''isāna'' or Sanskrit ईशान्य ''īśānya'' "northeast") consists of 20 provinces in northeastern Thailand. Isan is T ...
. It contains a single species, ''Chalawan thailandicus'', with ''Chalawan shartegensis'' as a possible second species.


Discovery and naming

The first fossil of ''Chalawan'' was a nearly complete
lower jaw In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone i ...
collected in the early 1980s from a road-cut near the town of
Nong Bua Lamphu Nong Bua Lam Phu () is a town in Thailand Thailand, officially the Kingdom of Thailand and historically known as Siam (the official name until 1939), is a country in Southeast Asia on the Mainland Southeast Asia, Indochinese Peninsula. Wi ...
, in the upper part of the Phu Kradung Formation. This mandible was at first known from the posterior portion of the bone, described and named by Eric Buffetaut and Rucha Ingavat in
1980 Events January * January 4 – U.S. President Jimmy Carter proclaims a United States grain embargo against the Soviet Union, grain embargo against the USSR with the support of the European Commission. * January 6 – Global Positioning Sys ...
as a new species of the goniopholidid '' Sunosuchus'', ''Sunosuchus thailandicus''. Shortly afterwards more material of the same specimen was found and described. In the early 2000s another locality of the Phu Kradung Formation was discovered near the village of Kham Phok, Mukdahan Province, yielding amongst other remains the anterior tip of a crocodylomorph mandible which was assigned to ''Sunosuchus thailandicus'' and described the pholidosaurid affinity of the specimen. This interpretation remained tentative until the discovery of previously unnoticed skull material associated with the Kham Phok mandible (specimen PRC102-143) which led to the reassignment of ''Chalawan'' to the Pholidosauridae. Based on the new cranial material, Jeremy E. Martin, Komsorn Lauprasert, Eric Buffetaut, Romain Liard and Varavudh Suteethorn erected the new genus ''Chalawan'' and the ''
combinatio nova In biological taxonomy, a combinatio nova (abbreviated comb. nov. or n. comb.) refers to the formal renaming of an organism's scientific name when it is transferred to a different genus, reclassified within a different species, or its taxonomic ...
'' ''Chalawan thailandicus''. The generic name is derived from the name of Chalawan, a giant in the Thai folktale of '' Krai Thong'' that could take the form of a
crocodile Crocodiles (family (biology), family Crocodylidae) or true crocodiles are large, semiaquatic reptiles that live throughout the tropics in Africa, Asia, the Americas and Australia. The term "crocodile" is sometimes used more loosely to include ...
with
diamond Diamond is a Allotropes of carbon, solid form of the element carbon with its atoms arranged in a crystal structure called diamond cubic. Diamond is tasteless, odourless, strong, brittle solid, colourless in pure form, a poor conductor of e ...
teeth.


Description

The holotype mandible is very robust, and its tip is spoon shaped and wider than the portion of the jaw immediately behind it. The mandible of PRC102-143 is similar in size, though less complete, and preserves the same spatulate appearance. The 3rd and 4th dentary alveoli are raised and contiguous and hosted fang like teeth, after which the dentary constricts, much like seen in ''Sarcosuchus''. However unlike ''Sarcosuchus'', Chalawan does not possess a diastema between the enlarged 4th and small 5th dentary alveoli. The referred cranial material suggests a possible skull length of . The complete animal could reach over in body length, although other estimates suggest . The specimen shows a combination of goniopholidid and pholidosaur features and has a robust tubular and slightly flattened rostrum widening abruptly around the jugals. The nasals are separated from the nares by the premaxilla, which tapers at its posterior end and extends far between the maxilla, and the supratemporal fenestrae are moderately sized, making up 25% of the skull table, differing from ''
Terminonaris ''Terminonaris'' is a genus of extinct pholidosaurid crocodyliforms that lived during the Late Cretaceous (Cenomanian and TuronianWu X-C, Russell AP, & Cumbaa SL. 2001. ''Terminonaris'' (Archosauria: Crocodyliformes): new material from Saskatchew ...
'' and '' Oceanosuchus'' whose fenestrae take up more of the skull table's surface. The premaxillary teeth are located on the anterior half of the premaxilla and when seen in lateral view the anterior margin extends further down than the rest of the bone, creating a characteristic "beak" shape also seen in ''Sarcosuchus'' and ''Terminonaris robusta''. The transverse area of the premaxilla is separated from the rest of the bone by a
diastema A diastema (: diastemata, from Greek , 'space') is a space or gap between two teeth. Many species of mammals have diastemata as a normal feature, most commonly between the incisors and molars. More colloquially, the condition may be referred to ...
. This along with the hook-like shape, whereas traditional goniopholidids possess a notch in this area. Both the presence of a diastema and the hook-like anterior margin of the premaxilla are considered to be pholidosaurid characteristics. The exact age of ''Chalawan'' is uncertain, as the Phu Kradung Formation may either date to the Late Jurassic or Early Cretaceous. While vertebrate fossil discoveries point at a Late Jurassic origin,
palynology Palynology is the study of microorganisms and microscopic fragments of mega-organisms that are composed of acid-resistant organic material and occur in sediments, sedimentary rocks, and even some metasedimentary rocks. Palynomorphs are the mic ...
data instead suggests that the Formation represents Early Cretaceous sediments.Racey and Goodall (2009). "Late Palaeozoic and Mesozoic Ecosystems in SE Asia". Geological Society. London. Special Publication 315 Pp 69-84.


Phylogeny

The phylogenetic tree below depicts Pholidosauridae as recovered by Jouve & Jalil (2020) obtained from a matrix including 224 characters. In this tree ''Chalawan'' was recovered as a sister taxon to the genus ''Sarcosuchus''.


''"Sunosuchus" shartegensis''

Halliday ''et al.'' (2013) tentatively synonymized ''"Sunosuchus" shartegensis'' (Efimov, 1988) as ''"Sunosuchus"'' cf. ''thailandicus''. ''"S." shartegensis'' is known solely from the
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
PIN 4174‒1, a fragmented
skull The skull, or cranium, is typically a bony enclosure around the brain of a vertebrate. In some fish, and amphibians, the skull is of cartilage. The skull is at the head end of the vertebrate. In the human, the skull comprises two prominent ...
, comprising the rostrum, the preorbital region of the skull roof, the quadrates and parts of the quadratojugal, the
occipital condyle The occipital condyles are undersurface protuberances of the occipital bone in vertebrates, which function in articulation with the superior facets of the Atlas (anatomy), atlas vertebra. The condyles are oval or reniform (kidney-shaped) in shape ...
and nearly complete mandibles. It was collected from "Layer 2" of the
Tithonian In the geological timescale, the Tithonian is the latest age (geology), age of the Late Jurassic Epoch and the uppermost stage (stratigraphy), stage of the Upper Jurassic Series. It spans the time between 149.2 ±0.7 annum, Ma and 143.1 ±0.6 (mi ...
(Late Jurassic) Ulan Malgait beds, in the Shar Teeg locality, of the
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of Outer
Mongolia Mongolia is a landlocked country in East Asia, bordered by Russia to the north and China to the south and southeast. It covers an area of , with a population of 3.5 million, making it the world's List of countries and dependencies by po ...
, embedded in grey clay. Halliday ''et al.'' (2013) stated that ''"S." shartegensis'' shares some features with other species of ''Sunosuchus'', and can not be differentiated from the holotype of ''S. thailandicus''. Nevertheless, it lacks definitive
synapomorphies 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 ...
of ''S. thailandicus'', and possibly even these of Goniopholididae, suggesting that it might belong to a different species. Using an updated version of Andrade ''et al.'' (2011) phylogenetic analysis, Halliday ''et al.'' (2013) found ''"S." shartegensis'' to be the
sister taxon In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
of '' Kansajsuchus'' from
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. The addition of ''S. thailandicus'' (based on the holotype) to the analysis did not confirm the referral of ''"S." shartegensis'' to ''"S."'' cf. ''thailandicus'', as it resulted in a large
polytomy An internal node of a phylogenetic tree is described as a polytomy or multifurcation if (i) it is in a rooted tree and is linked to three or more child subtrees or (ii) it is in an unrooted tree and is attached to four or more branches. A tree ...
. As the paper synonymizing the two crocodylomorphs appeared shortly before Martin ''et al.s description of the cranial material, the taxonomic status of ''"S." shartegensis'' remains unknown.


References

{{Taxonbar, from=Q15221406 Tethysuchia Early Cretaceous crocodylomorphs of Asia Early Cretaceous reptiles of Asia Cretaceous Thailand Fossils of Thailand Fossil taxa described in 2013 Taxa named by Éric Buffetaut Taxa named by Varavudh Suteethorn Prehistoric pseudosuchian genera