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Lotheridium

Lotheridium is an extinct genus of deltatheroidan mammals that lived in what is now Asia during the Late Cretaceous, about 72–66 million years ago. The genus contains a single species, Lotheridium mengi, named in 2015 after paleontologist Jin Meng. It is known from a single fossil specimen—a skull with associated lower jaws—found in the Qiupa Formation of Henan Province, China and housed in the collections of the Zhejiang Museum of Natural History. The skull measures 67.3 mm (2.65 in) in length, suggesting Lotheridium was large compared to most other deltatheroidans. Though the preserved skull is almost complete, it has been flattened and the skull roof was crushed during fossilization.

Discovery and naming
with a corresponding stratigraphic chart|alt=Map of China with a closeup of Luanchuan, a stratigraphic chart and a photo of brownish rock deposits In 2015, a team of four Chinese paleontologists reported the discovery of an almost complete fossilized skull in Henan Province, China. The skull was excavated from Late Cretaceous-aged rock deposits of the Qiupa Formation in Haoping Village, Luanchuan County, and was deposited at the Zhejiang Museum of Natural History under the specimen number ZMNH M9032. After studying this fossil, the team concluded that it represented a previously unknown genus and species of prehistoric mammal which they named Lotheridium mengi. They designated the skull as the holotype (name-bearing) specimen. The generic name combines the name of Luoyang (the prefecture Luanchuan County is part of) with the Ancient Greek word theridion (meaning "small beast"). The specific name honors Jin Meng (a paleontologist who studies Mesozoic mammals). ==Description==
Description
Lotheridium is currently known from only one specimen: the holotype skull with associated lower jaws. The specimen is nearly complete, but has been flattened during the process of fossilization such that the skull roof has been crushed. This skull has a total length of , suggesting that Lotheridium was much larger than most other deltatheroidans with the exception of Deltatheroides. As all the teeth are erupted and worn, this skull likely belongs to a full-grown adult. Because the amount of wear on each molar decreases towards the back of the mouth in both the upper and lower jaws, it has been determined that the molars erupt in order from front to back. Unlike in all other deltatheroidans, the protocones (one of the cusps) on the upper molars of Lotheridium stretch further sideways and the lower molars bear small cusps with a shelf-like structure (named "cuspule f"), so these features are used as the diagnostic traits to distinguish Lotheridium from its relatives. ==Classification==
Classification
of Lotheridium|alt=Digital illustration of a mammal with reddish brown fur, a black head, orange belly and cream-coloured chin and tail tip Lotheridium belongs to the family Deltatheridiidae within an extinct order of mammals known as the Deltatheroida. The deltatheroidans are part of the larger clade Metatheria, whose only extant members are the marsupials, meaning that marsupials are the closest living relatives of Lotheridium. In the 2015 study first describing this genus, a phylogenetic analysis was carried out, recovering Lotheridium in a sister group position to a clade including Atokatheridium, Nanocuris, Deltatheridium and Deltatheroides. |label1=Deltatheroida}} The pattern of deltatheroidan dispersal has been debated among experts. Historically, the group was believed to have first evolved in Asia and later spread into North America. This was because the former continent has yielded most of their known specimens, including some of the oldest known at the time (Coniacian-aged fossils of Sulestes from Uzbekistan). However, North American fossils of Atokatheridium and Oklatheridium which predate any known Asian deltatheroidans were later reported in the 2000s, leading some authors to believe that the group originated here. Even so, others continued to believe in an Asian origin for deltatheroidans, citing how Asian genera tend to be recovered in more basal (earlier-diverging) positions in phylogenetic analyses. Being one of the geologically youngest deltatheroidans, Lotheridium does not provide new information on the origins of the group, but its discovery in central China (where deltatheroidans had not been found previously) does show that the group was more widespread in Asia than formerly known. ==Paleobiology==
Paleobiology
of the upper (B) and lower (C) jaws, with closeups of the upper (A) and lower (D) molars|right|thumb Deltatheroidans such as Lotheridium are believed to have been carnivorous animals. This is supported by the fact that two of the ridges—the postmetacrista and preprotocrista—of their upper molars form a prominent mechanism for shearing flesh. Similar shearing mechanisms formed by the postmetacrista have convergently evolved in other carnivorous mammals such as carnivorans, dasyurids, borhyaenids and stagodontids, though the preprotocrista is not involved in the shearing mechanism of marsupials and eutherian mammals. In addition, a study on jaw shapes to predict the feeding ecology of Mesozoic mammals further supports that the related Deltatheridium was a carnivore. Potential prey of deltatheroidans may have included animals up to the size of small dinosaurs, as the skull of a juvenile Archaeornithoides (a theropod dinosaur) has been found with bite marks matching a mammal of this group. Based on the amount of wear, the third premolar of both the upper and lower jaws in the only known Lotheridium specimen likely erupted far later than the first and second premolars. The wear patterns indicate that Lotheridium had the same tooth replacement pattern as marsupials, and that this pattern had already evolved in the common ancestor of marsupials and deltatheroidans. This further suggests that the two groups may have similar feeding systems and reproductive patterns. ==Paleoenvironment==
Paleoenvironment
The only known specimen of Lotheridium was collected from an exposure of the Qiupa Formation. The Qiupa Formation has been divided into three sections, and the exposure at Haoping is considered part of Section B. The fossil-bearing beds of the formation are located below the K-T boundary (which marks the end of the Mesozoic era), and are therefore believed to have formed during the Maastrichtian age of the Cretaceous period (between 72 and 66 million years ago). At this time, the area had a shallow lake and a braided river delta. This can be determined from how the fossil-bearing beds are made up of mudstones and siltstones with interbedded fine conglomerates, which form in such environments. Geochemical and pollen analyses indicates that the area was humid to semihumid and possessed a tropical to subtropical climate during the Late Cretaceous. The deposits of the Qiupa Formation have yielded fossilized remains of a wide variety of animals which likely lived alongside Lotheridium. Remains of one other mammal species, the large multituberculate Yubaatar zhongyuanensis, have been discovered at this site. Smaller theropod dinosaurs have also been named from the Qiupa Formation, including the dromaeosaurid Luanchuanraptor, the oviraptorid Yulong, the ornithomimid Qiupalong, the alvarezsaurid Qiupanykus, and the enantiornithine bird Yuornis. Partial remains of sauropods, ankylosaurs and possibly protoceratopsids are also known, but have not been attributed to any named genera or species. Aside from dinosaurs, reptiles known from the Qiupa Formation fossil assemblage include the lizards Funiusaurus, Tianyusaurus, and Zhongyuanxi, as well as unnamed turtles. ==References==
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