Largocephalosaurus

''Largocephalosaurus'' is an extinct genus of basal saurosphargid, a marine reptile known from the Middle Triassic (Anisian age) Guanling Formation of Yunnan and Guizhou Provinces, southwestern China. It contains a type species, ''Largocephalosaurus polycarpon'', and a second species ''L. qianensis''.

''L. polycarpon'' was initially interpreted as an eosauropterygian sauropterygian closely related to European pachypleurosaurs and nothosaurids. However following additional preparation of the postcranial skeleton and the discovery of a second better known species ''L. qianensis'', Li ''et al.'' (2014) reinterpreted ''Largocephalosaurus'' as a basal member of the family Saurosphargidae and a close relative of ''Saurosphargis'' and '' Sinosaurosphargis''. According to Li ''et al.'' (2014), saurosphargids did not belong to Sauropterygia, although they were closely related to it, forming its sister taxon.

Discovery

The type species of ''Largocephalosaurus'', ''L. polycarpon'' is known only from the holotype WIGM SPC V 1009 a nearly complete and articulated skull and skeleton missing most of its tail and measuring more than in length, housed at the Wuhan Institute of Geology and Mineral Recourses. SPC V 1009 was collected from Member II of the Guanling Formation, dating to the Pelsonian substage of the mid-late Anisian stage of the early Middle Triassic, about 243 million years ago, at Luoping County of Yunnan Province.

While ''L. polycarpon'' was being studied at the WIGM, three specimens of what was later named ''L. qianensis'' were prepared at the Geological Museum of Peking University and the Institute of Vertebrate Paleontology and Paleoanthropology. These three specimens were collected in 2008 about 100 km northeast of the type locality of ''L. polycarpon'' at Xinmin District of Panxian County, southwestern Guizhou Province. They, like SPC V 1009, came from Member II of the Guanling Formation, dating to the late Anisian. Out of these specimens, IVPP V 15638, a nearly complete skeleton exposed in view from below with only the back of the tail missing, was chosen to represent the holotype of ''L. qianensis''. The other two specimen are referable to ''L. qianensis'' and include GMPKU-P-1532-A, a skull with a neck vertebra in view from above, and GMPKU-P-1532-B, an incomplete postcranial skeleton in view from above, with the back portion of the mandible preserved, but missing most of the tail.

All specimens of ''Largocephalosaurus'' came from the Lagerstätte discovered during a 2007 geological mapping project with a diverse record of marine life called Luoping Biota, that yielded, apart from ''Largocephalosaurus'', various invertebrates, fish, basal ichthyosaurs, ''Atopodentatus'', the advanced saurosphargid '' Sinosaurosphargis'', and several species of eosauropterygian, including both pachypleurosaurs and nothosaurids.

Etymology

''Largocephalosaurus'' was first described and named by Long Cheng, Xiaohong Chen, Xiongwei Zeng and Yongjian Cai in 2012 with the type species ''Largocephalosaurus polycarpon''. The generic name is said to be derived from Latin ''largus'', meaning "large", and ''cephalus'', meaning "head", and from Greek ''sauros'', meaning "lizard", a common suffix for genus names of extinct reptiles. In classical Latin, the proper word for "head" is ''caput'', while in ancient Greek ''kephalē'' (κεφαλή) was used for "head".Liddell, H.G. & Scott, R. (1940). ''A Greek-English Lexicon. revised and augmented throughout by Sir Henry Stuart Jones. with the assistance of. Roderick McKenzie.'' Oxford: Clarendon Press. ''Kephalos'' (Κέφαλος) was the first name of various Greek mythological and historical figures that was rendered in Latin as ''Cephalus''.Lewis, C.T. & Short, C. (1879). ''A Latin dictionary founded on Andrews' edition of Freund's Latin dictionary.'' Oxford: Clarendon Press.
The specific name ''polycarpon'' is said to be derived from Latin ''poly'', meaning "many", plus "carpon", meaning "ossified carpals", in reference to the eleven carpal ossifications the forelimb of ''L. polycarpon'' shows, resulting in the unique hand bone formula of 2-3-4-5-5. In classical Latin the proper word for "many" is ''multus'', while in ancient Greek they used the word ''polys'' (πολύς).

A second species ''L. qianensis'' was named by Chun Li, Da-Yong Jiang, Long Cheng, Xiao-Chun Wu and Olivier Rieppel in 2014. Its specific name is derived from ''Qian'', a shortened alternative spelling for Guizhou, plus the Latin suffix ''-ensis'' meaning "from", in reference to the different occurrence of the species - while the specimens of ''L. qianensis'' were discovered in Guizhou Province, the only known specimen of ''L. polycarpon'' came from the neighboring Yunnan Province.

Description

''Largocephalosaurus'' was a relatively large saurosphargid, with the type specimen of the second species ''L. qianensis'' measuring up to in total body length. It is a basal saurosphargid, less modified morphologically from the archetypal diapsid reptile compared to advanced saurosphargids like '' Sinosaurosphargis''. However, even in its original description that interpreted it as an eosauropterygian sauropterygian, several traits later identified as saurosphargid were noted. These include back vertebrae that show elongated, distally expanded transverse projections, and its broadened and flattened chest ribs. Yet, ''Largocephalosaurus'' unlike other saurosphargids retains an elongate body shape, a well-developed supratemporal fenestra, an oval suborbital fenestra (although much reduced compared to non-saurosphargids) and an incomplete dorsal osteoderm carapace. Additionally, the advanced saurosphargids ''Saurosphargis'' and ''Sinosaurosphargis'' have broader and flatter rib baskets, and also transverse projections of back vertebrae and proximal portions of chest ribs that are much more robust than in ''Largocephalosaurus'', being much wider than the spacing between them. Other differences include: the presence of three or four teeth on the premaxilla, and its exclusion from the external nostril outline; an elongated frontal bone back-side facing projection; the presence of a row of large osteoderms on the top of the neural spines, in the midline of the partial osteoderm shield; and similarly rounded coracoid and pubis bone with the coracoid foramen and the obturator fenestra present and open (respectively).

Due to the fact that the holotype skeleton of ''L. polycarpon'' is still only partially prepared, Li ''et al.'' (2014) refrained from providing a revised description for it. Nevertheless, at least 9 morphological differences from the second the species ''L. qianensis'' were observed. These include the paired parietal bone, the large pineal foramen, the oval eye-sockets, and the presence of 33 presacral vertebrae (9 neck and 24 back vertebrae) and two sacral vertebrae, in ''L. qianensis''. Additionally, a distinct palatal fossa exists on the upper surface of palatine bone in ''L. qianensis'', and the retroarticular process is very short. The back osteoderms of ''L. qianensis'' are elongated granular in shape, and covering the sides of body. Finally, its shoulder portion of the chest ribs is broadened in a hook-like projection, while lacking the crest on its upper surface like the one seen in ''L. polycarpon''.

Phylogeny

''Largocephalosaurus'' was first included in a phylogenetic analysis in its original description. Due to the lack of preparation of the holotype, especially of the postcranial portion, and the fact that no other saurosphargids or stem sauropterygians (like ''Eusaurosphargis'' and ''Helveticosaurus'') were included, ''L. polycarpon'' was recovered in a position most closely related to ''Wumengosaurus'' within Eosauropterygia, basal to the European pachypleurosaurs and nothosaurids. However, following additional preparation of the postcranial skeleton and the discovery of a second better known species ''L. qianensis'', Li ''et al.'' (2014) presented an analysis which includes ''Eusaurosphargis'', ''Helveticosaurus'' and all known saurosphargid species. The following cladogram is simplified after their results. The removal / inclusion of Ichthyopterygia was found to affect the topology the most - switching the positions of the ''Eusaurosphargis''+''Helveticosaurus'' and Thalattosauriformes clades, and altering the positions of several taxa within Eosauropterygia, which are not shown.

Notes

References

{{Taxonbar, from=Q6489300

Middle Triassic reptiles of Asia
Anisian life
Triassic China
Fossils of China
Paleontology in Guizhou
Guanling Formation
Fossil taxa described in 2012
Fossil taxa described in 2014
Prehistoric reptile genera