Chalawan Thailandicus
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''Chalawan'' (from th, ชาละวัน ) 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 ...
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 pholidosaurid
mesoeucrocodylia Mesoeucrocodylia is the clade that includes Eusuchia and crocodyliforms formerly placed in the paraphyletic group Mesosuchia. The group appeared during the Early Jurassic, and continues to the present day. Diagnosis It was long known that M ...
n known from the
Late Jurassic The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 163.5 ± 1.0 to 145.0 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987. In European lithostratigraphy, the name ...
or
Early Cretaceous The Early Cretaceous ( geochronological name) or the Lower Cretaceous (chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145  Ma to 100.5 Ma. Geology Pro ...
Phu Kradung Formation The Early Cretaceous Phu Kradung Formation is the lowest member of the Mesozoic Khorat Group which outcrops on the Khorat Plateau in Isan, Thailand. This geological formation consists of micaceous, brown to reddish-brown siltstone beds with minor ...
of
Nong Bua Lamphu Province Nong Bua Lamphu ( th, หนองบัวลำภู, , ) is one of Thailand's seventy-six provinces (''changwat'') lies in upper northeastern Thailand also called Isan. Neighbouring provinces are (from north clockwise) Udon Thani, Khon Kaen, ...
,
northeastern Thailand Northeast Thailand or Isan (Isan/ th, อีสาน, ; lo, ອີສານ; also written as Isaan, Isarn, Issarn, Issan, Esan, or Esarn; from Pali ''īsānna'' or Sanskrit ईशान्य ''īśānya'' "northeast") consists of 20 provin ...
. 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 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 teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
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, capital of Nong Bua Lamphu Province. It is on the central eastern border of the province, approximately 45 kilometers south-west of the city of Udon Thani and from there, accessed by route 210. The town l ...
, in the upper part of the
Phu Kradung Formation The Early Cretaceous Phu Kradung Formation is the lowest member of the Mesozoic Khorat Group which outcrops on the Khorat Plateau in Isan, Thailand. This geological formation consists of micaceous, brown to reddish-brown siltstone beds with minor ...
. This mandible was at first known from the posterior portion of the bone, described and named by
Eric Buffetaut The given name Eric, Erich, Erikk, Erik, Erick, or Eirik is derived from the Old Norse name ''Eiríkr'' (or ''Eríkr'' in Old East Norse due to monophthongization). The first element, ''ei-'' may be derived from the older Proto-Norse ''* ain ...
and Rucha Ingavat in
1980 Events January * January 4 – U.S. President Jimmy Carter proclaims a grain embargo against the USSR with the support of the European Commission. * January 6 – Global Positioning System time epoch begins at 00:00 UTC. * January 9 – ...
as a new species of the
goniopholidid Goniopholididae is an extinct family of moderate-sized semi-aquatic neosuchian crocodyliformes. Their bodyplan and morphology are convergent on living crocodilians. They lived across Laurasia (Asia, Europe and North America) between the Middle Ju ...
''
Sunosuchus ''Sunosuchus'' is an extinct genus of goniopholidid mesoeucrocodylian. Fossils are known from China, Kyrgyzstan, and Thailand and are Jurassic in age, although some may be Early Cretaceous. Four species are currently assigned to the genus: the ...
'', ''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 Pholidosauridae is an extinct 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 Tunisia), North Amer ...
. 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 ''Combinatio nova'', abbreviated ''comb. nov.'' (sometimes ''n. comb.''), is Latin for "new combination". It is used in taxonomic biology literature when a new name is introduced based on a pre-existing name. The term should not to be confused wi ...
'' ''Chalawan thailandicus''. The generic name is derived from the name of Chalawan, a giant in the Thai folktale of ''
Krai Thong ''Krai Thong'' or ''Kraithong'' ( th, ไกรทอง, ) is a Thai folktale, originating from Phichit Province. It tells the story of Chalawan, a crocodile lord who abducts a daughter of a wealthy Phichit man, and Kraithong, a merchant from No ...
'' 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 even more loosely to inclu ...
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. Another solid form of carbon known as graphite is the Chemical stability, chemically stable form of car ...
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 . Complete animal could reached over in body length, although other estimates suggest . The specimen shows a combination of goniopholid 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 Saskatche ...
'' and ''
Oceanosuchus ''Oceanosuchus'' is a genus of pholidosaurid mesoeucrocodylian, a type of marine crocodylomorph. It is known from a skull and partial skeleton found in early Cenomanian-age rocks from Normandy, France. The rostrum of the skull was relatively s ...
'' 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 (plural diastemata, from Greek διάστημα, 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 ...
. This along with the hook-like shape, whereas traditional goniopholids 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 dust" (from grc-gre, παλύνω, palynō, "strew, sprinkle" and '' -logy'') or of "particles that are strewn". A classic palynologist analyses particulate samples collected from the air, from water, or from deposit ...
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 is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...
PIN 4174‒1, a fragmented
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, the ...
, 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 vertebra. The condyles are oval or reniform (kidney-shaped) in shape, and their anteri ...
and nearly complete mandibles. It was collected from "Layer 2" of the
Tithonian In the geological timescale, the Tithonian is the latest age of the Late Jurassic Epoch and the uppermost stage of the Upper Jurassic Series. It spans the time between 152.1 ± 4 Ma and 145.0 ± 4 Ma (million years ago). It is preceded by the K ...
(Late Jurassic) Ulan Malgait beds, in the Shar Teeg locality, of the
Govi-Altai Province Govi-Altai ( mn, Говь-Алтай / , ) is an aimag (province) in western Mongolia. Transportation The Altai Airport (LTI/ZMAT) has one paved runway and is served by regular flights to Arvaikheer, Bayankhongor and Ulaanbaatar Ulaanbaatar ...
of Outer
Mongolia Mongolia; Mongolian script: , , ; lit. "Mongol Nation" or "State of Mongolia" () is a landlocked country in East Asia, bordered by Russia to the north and China to the south. It covers an area of , with a population of just 3.3 million, ...
, 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 have ...
of ''S. thailandicus'', and possibly even these of
Goniopholididae Goniopholididae is an extinct family of moderate-sized semi-aquatic neosuchian crocodyliformes. Their bodyplan and morphology are convergent on living crocodilians. They lived across Laurasia (Asia, Europe and North America) between the Middle Ju ...
, 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 t ...
of ''
Kansajsuchus ''Kansajsuchus'' is an extinct genus of paralligatorid mesoeucrocodylian. It is based on PIN 2399/301, a right premaxilla, one of the bones of the tip of the snout. This specimen was found in rocks of the lower Santonian-age Upper Cretaceo ...
'' from
Tajikistan Tajikistan (, ; tg, Тоҷикистон, Tojikiston; russian: Таджикистан, Tadzhikistan), officially the Republic of Tajikistan ( tg, Ҷумҳурии Тоҷикистон, Jumhurii Tojikiston), is a landlocked country in Centr ...
. 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 tr ...
. 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 Early Cretaceous crocodylomorphs of Asia Early Cretaceous reptiles of Asia Cretaceous Thailand Fossils of Thailand Fossil taxa described in 1980 Fossil taxa described in 2013 Taxa named by Éric Buffetaut Taxa named by Varavudh Suteethorn Prehistoric pseudosuchian genera