''Tholodus'' 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 basal

ichthyopterygia Ichthyopterygia ("fish flippers") was a designation introduced by Richard Owen, Sir Richard Owen in 1840 to designate the Jurassic ichthyosaurs that were known at the time, but the term is now used more often for both true Ichthyosauria and their ...

n known from the

Middle Triassic In the geologic timescale, the Middle Triassic is the second of three epoch (geology), epochs of the Triassic period (geology), period or the middle of three series (stratigraphy), series in which the Triassic system (stratigraphy), system is di ...

(mid-late

Anisian In the geologic timescale, the Anisian is the lower stage (stratigraphy), stage or earliest geologic age, age of the Middle Triassic series (stratigraphy), series or geologic epoch, epoch and lasted from million years ago until million years ag ...

to late

Ladinian The Ladinian is a stage and age in the Middle Triassic series or epoch. It spans the time between Ma and ~237 Ma (million years ago). The Ladinian was preceded by the Anisian and succeeded by the Carnian (part of the Upper or Late Triassic ...

stage) of

Germany Germany, officially the Federal Republic of Germany, is a country in Central Europe. It lies between the Baltic Sea and the North Sea to the north and the Alps to the south. Its sixteen States of Germany, constituent states have a total popu ...

, northeastern

Italy Italy, officially the Italian Republic, is a country in Southern Europe, Southern and Western Europe, Western Europe. It consists of Italian Peninsula, a peninsula that extends into the Mediterranean Sea, with the Alps on its northern land b ...

and possibly

China China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...

. It was first named by

Christian Erich Hermann von Meyer Christian Erich Hermann von Meyer (3 September 1801 – 2 April 1869), known as Hermann von Meyer, was a German palaeontologist. He was awarded the 1858 Wollaston medal by the Geological Society of London. Life He was born in Frankfurt am ...

1851 Events January–March * January 11 – Hong Xiuquan officially begins the Taiping Rebellion in China, one of the bloodiest revolts that would lead to 20 million deaths. * January 15 – Christian Female College, modern-d ...

and the

type species In International_Code_of_Zoological_Nomenclature, 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 spe ...

is ''Tholodus schmidi''. It is known from many disarticulated and fragmentary remains, mainly teeth and

jaw The jaws are a pair of opposable articulated structures at the entrance of the mouth, typically used for grasping and manipulating food. The term ''jaws'' is also broadly applied to the whole of the structures constituting the vault of the mouth ...

fragments. Most specimens were collected from various localities across the Ladinian-aged

Muschelkalk The Muschelkalk (German for "shell-bearing limestone"; ) is a sequence of sedimentary rock, sedimentary rock strata (a lithostratigraphy, lithostratigraphic unit) in the geology of central and western Europe. It has a Middle Triassic (240 to 230 m ...

, Germany, mainly from the Jena Formation of the upper Lower Muschelkalk, where 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 ...

was found. Dalla Vecchia (2004) recently described two additional specimens, a mandibular ramus and a

maxilla In vertebrates, the maxilla (: maxillae ) 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 two maxil ...

, both bearing teeth and nearly uncrushed, and some

postcrania The postcranium ("behind the cranium"; plural: postcrania) or postcranial skeleton in zoology and vertebrate paleontology is the skeleton apart from the skull. The postcranium encompasses the axial skeleton, which includes the entirety of the verte ...

l remains, from a single late Anisian outcrop, from the southern

Alps The Alps () are some of the highest and most extensive mountain ranges in Europe, stretching approximately across eight Alpine countries (from west to east): Monaco, France, Switzerland, Italy, Liechtenstein, Germany, Austria and Slovenia. ...

of Italy. The

humerus The humerus (; : humeri) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius (bone), radius and ulna, and consists of three sections. The humeral upper extrem ...

resembled that of immature individuals of the Asian genus ''

Chaohusaurus ''Chaohusaurus'' is an extinct genus of basal ichthyosauriform, depending on definition possibly ichthyosaur, from the Early Triassic of Chaohu and Yuanan, China. Discovery and naming The type species ''Chaohusaurus geishanensis'' was name ...

'', suggesting possible affinities to

Grippidia Ichthyopterygia ("fish flippers") was a designation introduced by Sir Richard Owen in 1840 to designate the Jurassic ichthyosaurs that were known at the time, but the term is now used more often for both true Ichthyosauria and their more primitiv ...

Jiang ''et al.'' (2008) described and named ''

Xinminosaurus ''Xinminosaurus'' is an extinct genus of cymbospondylid ichthyosaur known from the Middle Triassic (mid-late Anisian stage) of Guizhou Province, China. Etymology The generic name is derived from ''Xinmin'', the district where the fossil was fo ...

'' from the mid-late Anisian

Guanling Formation The Guanling Formation is a Middle Triassic (Anisian or Pelsonian in the regional chronostratigraphy) geologic Formation (geology), formation in southwestern China. Geology The formation encompasses two members. The first member is primarily c ...

Guizhou ) , image_skyline = , image_caption = , image_map = Guizhou in China (+all claims hatched).svg , mapsize = 275px , map_alt = Map showing the location of Guizhou Province , map_caption = Map s ...

, China. Maisch (2010) suggested that ''Xinminosaurus'' might be a subjective

junior synonym In taxonomy, the scientific classification of living organisms, a synonym is an alternative scientific name for the accepted scientific name of a taxon. The botanical and zoological codes of nomenclature treat the concept of synonymy differently. ...

of ''Tholodus''. Although Jiang ''et al.'' (2008) regarded ''Tholodus'' to be a possible ''

nomen dubium In binomial nomenclature, a ''nomen dubium'' (Latin for "doubtful name", plural ''nomina dubia'') is a scientific name that is of unknown or doubtful application. Zoology In case of a ''nomen dubium,'' it may be impossible to determine whether a ...

'', Maisch (2010) rejected this opinion, stating that ''Tholodus'' is easily recognized and characterized by unequivocal dental

autapomorphies In phylogenetics, an autapomorphy is a distinctive feature, known as a derived trait, that is unique to a given taxon. That is, it is found only in one taxon, but not found in any others or outgroup taxa, not even those most closely related to ...

, so that even jaw and tooth fragments are diagnostic, and it is thus a valid taxon. Furthermore, he noted that ''Tholodus'' is clearly distinguishable from all other known marine reptiles except ''Xinminosaurus''. The only difference between the taxa, according to Maisch (2010), is that ''Tholodus'' specimens are at average twice as large as ''Xinminosaurus'' holotype.

References

Fossil taxa described in 1851 Late Triassic ichthyosaurs Middle Triassic reptiles of Europe Anisian life Ichthyosauromorph genera {{triassic-reptile-stub