Khunnuchelys Temporal range: Turonian-Campanian, 91–75 Ma PreꞒ Ꞓ O S D C P T J K Pg N

Holotype skull of Khunnuchelys lophorhothon
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Reptilia
Order:	Testudines
Suborder:	Cryptodira
Superfamily:	Trionychia
Family:	Trionychidae
Subfamily:	Trionychinae
Genus:	†Khunnuchelys Brinkman et al., 1993
Type species
†Khunnuchelys erinhotensis Brinkman et al., 1993
Species
†K. erinhotensis Brinkman et al., 1993 †K. kizylkumensis Brinkman et al., 1993 †K. lophorhothon Danilov et al., 2013
Synonyms
Synonyms of K. kizulkumensis^[1] Trionyx sp. Nessov, 1986 cf. Eurycephalochelys sp. Chkhikvadze and Shuvalov, 1988 cf. Axestemys riabinini Kordikova, 1994a Axestemys (Axestemys) sp. Kordikova, 1994b Synonyms of K. lophorhothon^[2] Trionychidae indet. Kordikova, 1994 cf. Lophorhothon sp. Nessov, 1995 Khunnuchelys sp. Nessov, 1997 Khunnuchelys sp. 1 Danilov and Vitek, 2012 Khunnuchelys sp. 2 Danilov and Vitek, 2012

Khunnuchelys was a genus of trionychine turtle from the Late Cretaceous of Asia. Three species are known, K. erinhotensis, the type species, K. kizylkumensis, and K. lophorhothon. K. erinhotensis is known from the Iren Dabasu Formation in China from the late Turonian until the middle Campanian. K. kizylkumensis is known from the late Turonian Bissekty Formation of Uzbekistan. The third species, described in 2013 by Danilov et al., is known from the early to middle Campanian aged Bostobe Formation of Kazakhstan.

History of naming

K. lophorothon is known from two skulls, and has a complex history of classifications. Originally, it was assigned to the dinosaur genus cf. Lophorhothon by Lev Alexandrovitch Nessov in 1995 as an unidentified species. Later, it was correctly transferred to the genus Khunnuchelys by Danilov and Vitek (2012), but was not described in any detail. The two skulls were named Khunnuchelys sp. 1 and Khunnuchelys sp. 2. by them. The finder of the second skull was Mikhail E. Voskoboynikov, who allowed it to be studied by Lev I. Khosatzky.^[2]

K. lophorothon emends the original classification of the species, as the species name if derived from the name of Lophorhothon, the dinosaur it was originally classified under.^[2]

Description

Skull

Khunnuchelys is an extinct turtle is which many skulls are known from. In fact, is the only skull-based trionychid from the Cretaceous.^[2] The side of the maxillary makes up a beak-like structure, an unusual feature of trionychids.^[1] One skull, ZIN PH 1/146, is relatively small, and is only partly preserved. The maxilla and prefrontals of the specimen are only partly known, although the jugals, basioccipitals, supraoccipital crest, and left quadrate are close to complete. The preserved skull length is 9 cm (3.5 in), and the total approximate skull is 17 cm (6.7 in) long. The specimen is unique among Khunnuchelys, as the ventral edge is generally well-preserved. The ventral edge shows that the skull of Khunnuchelys is arched from the front to the back.^[2]

Distinguishing characteristics

A set of features unique among trionychids were identified by Brinkman et al. in 1993 during the description of Khunnuchelys erinhotensis and K. kisylkumensis. They are listed below:^[2]

orbits facing strongly forwards;
the suborbital region of the maxilla is deep;
the palate is vaulted;
the triturating surfaces of the maxilla and dentary are beaklike;
the triturating surfaces of the maxillae meet at the midline, forming a mid-ventral ridge;
the internal nares are located far posteriorly and are roofed by the palatines and maxillae;
the anterolateral corner of the palatine and posterolateral corner of the maxilla are greatly enlarged to form about half of the triturating surface;
the maxilla and jugal are swollen to form a broadly convex surface in dorsal view;
the postorbital contacts the maxilla, excluding the jugal from the margin of the orbit;
the roof of the skull is constricted to expose the palatine in dorsal view;
the descending process of the parietal does not participate in the processus trochlearis oticum; a posteriorly facing occipital surface is bordered dorsally by a rounded ridge formed by the opisthotic and supraoccipital;
a concave surface is present on the occiput above the fenestra postotica and lateral to the foramen magnum;
the occipital surface of the exoccipital faces posteriorly;
and nervi hypoglossi exiting via a single foramen in the exoccipital.

The characteristics would not be useful if Khunnuchelys was not a trionychid, but Brinkman et al. found that a classification as a trionychid was likely because of the features listed below:^[2]

it is similar to Conchochelys in the presence of a deeply excavated palate, extensively developed secondary palate with broad contact of the maxillae at the midline, grooves extending backwards from the internal choanal openings, and short basioccipital region
and it is similar to Dogania in that the prefrontal makes a small contribution to the orbit margin between the frontal and maxilla.

K. lophorothon is distinguished from the other species by the features listed below, found by Danilov et al. in its description:^[2]

in that the palatines not only form the lateral walls of the grooves that open posteriorly from the apertura narium interna, but also form part of the floor of these grooves, leaving the grooves less ventrally exposed.

Both species can be differentiated from K. erinhotensis:^[2]

in that the upper margin of the external nares is located dorsal to the ventral orbit margins, and in that the foramen jugularis posterius is confluent with the fenestra postotica.

Classification

Khunnuchelys is a turtle that is grouped within Trionychinae, a subfamily of Trionychidae. The species "Trionyx" kansaiensis is known from shell fragments, that are from the same region of Kazakhstan as K. lophorothon, and might be from the later, in which case, "T." kansaiensis might be synonymous with K. lophorothon. Brinkman et al. found that Khunnuchelys was closely related to Conchochelys, but even though it is likely, Danilov et al. did not find any features to prove their close relationship.^[2]

References

^ ^a ^b Vitek, N.A.; Danilov, I.G. (2013). "Soft-shelled turtles (Trionychidae) from the Bissekty Formation (Upper Cretaceous: Turonian) of Uzbekistan: Skull-based taxa and probable skull-shell associations" (PDF). Cretaceous Research. 43 (2013): 48–58. doi:10.1016/j.cretres.2013.02.009. hdl:2152/41078. S2CID 128940175. Archived from the original (PDF) on 2021-08-31. Retrieved 2014-02-24.
^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j Danilov, I.G.; Vitek, N.S.; Averianov, A.O.; Glinskiy, V.N. (2013). "A new softshelled trionychid turtle of the genus Khunnuchelys from the Upper Cretaceous Bostobe Formation of Kazakhstan" (PDF). Acta Palaeontologica Polonica. doi:10.4202/app.2013.0045. Archived from the original (PDF) on 2014-02-28.

Trionychidae family

Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Subclass: Anapsida
Order: Testudines
Superfamily: Trionychia

Genera

Species of the Trionychidae family

Amyda

Asiatic softshell turtle
†Amyda gregaria
Amyda ornata

Apalone

†Axestemys

†Axestemys byssina
†Axestemys cerevisia
†Axestemys montinsana
†Axestemys quinni
†Axestemys splendida
†Axestemys vittata

Chitra

Cyclanorbis

Cycloderma

Dogania

Malayan softshell turtle

†Gilmoremys

†Gilmoremys gettyspherensis
†Gilmoremys lancensis

†Hutchemys

†Hutchemys arctochelys
†Hutchemys rememdium
†Hutchemys tetanetron
†Hutchemys sterea
†Hutchemys acupictus

†Khunnuchelys

†Khunnuchelys erinhotensis
†Khunnuchelys kizylkumensis
†Khunnuchelys lophorhothon

Lissemys

Nilssonia

†Palaeoamyda

†Palaeoamyda messeliana

Palea

Wattle-necked softshell turtle

Pelochelys

Pelodiscus

Rafetus

Trionyx

Phylogenetic arrangement of turtles based on turtles of the world 2017 update: Annotated checklist and atlas of taxonomy, synonymy, distribution, and conservation status. †=extinct.

Testudines

Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Clade: Diapsida
Order: Testudines

Genus

Chelonioidea
(Sea turtles)

Cheloniidae	†Allopleuron Caretta †Carolinochelys Chelonia †Eochelone Eretmochelys †Gigantatypus †Glarichelys †Itilochelys Lepidochelys †Mexichelys †Miocaretta Natator †Pacifichelys †Syllomus †Tasbacka
Dermochelyidae	†Arabemys †Corsochelys †Cosmochelys Dermochelys †Eosphargis †Mesodermochelys †Psephophorus
	†Euclastes †Peritresius †Procolpochelys †Protosphargis †Puppigerus

Kinosternoidea

Dermatemydidae	Dermatemys
Kinosternidae	Claudius †Hoplochelys Kinosternon Staurotypus Sternotherus

Testudinoidea

Emydidae	†Acherontemys Chrysemys Clemmys Deirochelys Emys Actinemys Emydoidea Glyptemys Graptemys Malaclemys Pseudemys Terrapene Trachemys †Wilburemys
Geoemydidae	Batagur †Banhxeochelys Cuora Cyclemys Geoclemys Geoemyda Hardella Heosemys Leucocephalon Malayemys Mauremys Melanochelys Morenia Notochelys Orlitia Pangshura Rhinoclemmys Sacalia Siebenrockiella Vijayachelys
Platysternidae	Platysternon
Testudinidae	Aldabrachelys Astrochelys Centrochelys Chelonoidis Chersina Cylindraspis †Cymatholcus †Floridemys Geochelone Gopherus †Hadrianus †Hesperotestudo Homopus Indotestudo Kinixys Malacochersus Manouria †Megalochelys †Oligopherus Psammobates Pyxis †Solitudo Stigmochelys †Stylemys Testudo

Trionychia

Carettochelyidae	†Allaeochelys †Anosteira Carettochelys
Trionychidae	Amyda Apalone †Axestemys Chitra Cyclanorbis Cycloderma †Drazinderetes Dogania †Gilmoremys †Hutchemys †Khunnuchelys Lissemys Nilssonia Palea †Palaeoamyda Pelochelys Pelodiscus Rafetus Trionyx
	†Basilochelys †Sinaspideretes

Chelydridae	Chelydra †Chelydrops †Chelydropsis †Emarginachelys †Macrocephalochelys Macrochelys †Planiplastron †Protochelydra
†Nanhsiungchelyidae	†Anomalochelys †Basilemys †Jiangxichelys
†Protostegidae	†Alienochelys †Archelon †Atlantochelys †Bouliachelys †Calcarichelys †Cratochelone †Desmatochelys †Iserosaurus †Notochelone †Ocepechelon †Pneumatoarthrus †Protostega †Rhinochelys †Santanachelys †Terlinguachelys
	†Adocus †Argillochelys †Bashuchelys †Ctenochelys †Prionochelys †Toxochelys

Pleurodira

†Araripemydidae	†Araripemys
†Bothremydidae	†Araiochelys †Arenila †Azabbaremys †Bothremys †Cearachelys †Chedighaii †Chupacabrachelys †Eotaphrosphys †Foxemys †Galianemys †Ilatardia †Inaechelys †Itapecuruemys †Jainemys †Kinkonychelys †Kurmademys †Labrostochelys †Nigeremys †Phosphatochelys †Polysternon †Puentemys †Rosasia †Rhothonemys †Sankuchemys †Taphrosphys †Ummulisani †Zolhafah
Chelidae	Acanthochelys Chelodina Chelus Elseya Elusor Emydura Hydromedusa †Lomalatachelys Mesoclemmys Myuchelys Phrynops Platemys †Prochelidella Pseudemydura Rheodytes Rhinemys †Yaminuechelys
Pelomedusidae	Pelomedusa Pelusios
Podocnemididae	†Albertwoodemys †Bauruemys †Brontochelys †Caninemys †Carbonemys †Cerrejonemys †Cordichelys Erymnochelys †Lapparentemys †Latentemys Peltocephalus Podocnemis †Stupendemys
†Sahonachelyidae	†Sahonachelys †Sokatra

	†Caribemys †Caririemys †Tacuarembemys

Phylogenetic arrangement of turtles based on Turtles of the World 2017 Update: Annotated Checklist and Atlas of Taxonomy, Synonymy, Distribution, and Conservation Status. † = extinct.

See also List of Testudines families