Zhelestidae Temporal range: Late Cretaceous [1] | |
---|---|
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Clade: | Eutheria |
Family: | † Zhelestidae Nessov (1985) |
Genera | |
Zhelestidae is a lineage of extinct eutherian mammals. Occurring in the Late Cretaceous from the Turonian to the Maastrichtian, they were an extremely successful group, with representatives present in Europe, Asia, India (and subsequently in Madagascar), Africa and North America, ostensibly rendering them a cosmopolitan clade. They were specialised towards an herbivorous lifestyle and were in fact initially considered stem-ungulates, [2] but the presence of epipubics and "archaic" dental characters render them as non-placental eutherians. [3] [4] [5] [6] [7] [8]
The earliest zhelestid remains occur in the Cenomanian of Central Asia. By the Campanian, however, they are present in Europe and North America, and by the Maastrichtian in India and in Madagascar (UA 8699). In Europe, then an island continent, taxa usually accepted to be zhelestids are the most common Late Cretaceous mammal remains, aside from Hateg Island where kogaionid multituberculates are more common instead, though there is a single possible zhelestid tooth there as well. [9]
Zhelestids are well specialised towards an herbivorous diet, their teeth suited for a lateral chewing mechanism similar to that of modern ungulates. An exception to this might be forms like Oxlestes and Khuduklestes , which are normally interpreted as carnivorous, and perhaps a few forms like Borisodon which may have been insectivorous. Ranging from mouse sized forms to ones comparable to small ungulates in size, zhelestids occupied a massive variety of ecological guilds.
In contrast to their dietary divergence, the few available limb remains suggest that they weren't very specialised locomotion wise, in contrast to some other basal eutherians like for instance Zalambdalestes , which was long-limbed and had several adaptations for cursoriality and hopping.
There is evidence that zhelestids were in direct competition with another clade of Mesozoic herbivorous mammals, the multituberculates, being rarer in places where these were most common and vice versa. [10] However, multituberculates eventually outlived zhelestids, surviving the K-Pg extinction event everywhere but Asia, where the niches taken by both clades in the Cretaceous would be taken by early rodents. [11]
Below is a phylogeny from Gheerbrant & Teodori (2021): [8]
Multituberculata is an extinct order of rodent-like mammals with a fossil record spanning over 130 million years. They first appeared in the Middle Jurassic, and reached a peak diversity during the Late Cretaceous and Paleocene. They eventually declined from the mid-Paleocene onwards, disappearing from the known fossil record in the late Eocene. They are the most diverse order of Mesozoic mammals with more than 200 species known, ranging from mouse-sized to beaver-sized. These species occupied a diversity of ecological niches, ranging from burrow-dwelling to squirrel-like arborealism to jerboa-like hoppers. Multituberculates are usually placed as crown mammals outside either of the two main groups of living mammals—Theria, including placentals and marsupials, and Monotremata—but usually as closer to Theria than to monotremes. They are considered to be closely related to Euharamiyida and Gondwanatheria as part of Allotheria.
Placental mammals are one of the three extant subdivisions of the class Mammalia, the other two being Monotremata and Marsupialia. Placentalia contains the vast majority of extant mammals, which are partly distinguished from monotremes and marsupials in that the fetus is carried in the uterus of its mother to a relatively late stage of development. The name is something of a misnomer considering that marsupials also nourish their fetuses via a placenta, though for a relatively briefer period, giving birth to less developed young which are then nurtured for a period inside the mother's pouch. Placentalia represents the only living group within Eutheria, which contains all mammals more closely related to placentals than to marsupials.
Eomaia is a genus of extinct fossil mammals containing the single species Eomaia scansoria, discovered in rocks that were found in the Yixian Formation, Liaoning Province, China, and dated to the Barremian Age of the Lower Cretaceous about 125 million years ago. The single fossil specimen of this species is 10 centimetres (3.9 in) in length and virtually complete. An estimate of the body weight is 20–25 grams (0.71–0.88 oz). It is exceptionally well-preserved for a 125-million-year-old specimen. Although the fossil's skull is squashed flat, its teeth, tiny foot bones, cartilages and even its fur are visible.
Eutheria, also called Pan-Placentalia, is the clade consisting of placental mammals and all therian mammals that are more closely related to placentals than to marsupials.
Condylarthra is an informal group – previously considered an order – of extinct placental mammals, known primarily from the Paleocene and Eocene epochs. They are considered early, primitive ungulates. It is now largely considered to be a wastebasket taxon, having served as a dumping ground for classifying ungulates which had not been clearly established as part of either Perissodactyla or Artiodactyla, being composed thus of several unrelated lineages.
Zalambdalestes is an extinct genus of eutherian mammal known from the Upper Cretaceous in Mongolia.
Embrithopoda ("heavy-footed") is an order of extinct mammals known from Asia, Africa and Eastern Europe. Most of the embrithopod genera are known exclusively from jaws and teeth dated from the late Paleocene to the late Eocene; however, the order is best known from its terminal member, the elephantine Arsinoitherium.
Protungulatum is an extinct genus of eutherian mammals within extinct family Protungulatidae, and is possibly one of the earliest known placental mammals in the fossil record, that lived in North America from the Late Cretaceous to early Paleocene.
Argiles et Grès à Reptiles Formation also known as the Argiles Rutilantes Formation is an early Maastrichtian French geologic formation in the département of Var preserving the remains of several types of dinosaurs and other extinct organisms.
Dryolestida is an extinct order of mammals, known from the Jurassic and Cretaceous. They are considered basal members of the clade Cladotheria, close to the ancestry of therian mammals. It is also believed that they developed a fully mammalian jaw and also had the three middle ear bones. Most members of the group, as with most Mesozoic mammals, are only known from fragmentary tooth and jaw remains.
UA 8699 is a fossil mammalian tooth from the Cretaceous of Madagascar. A broken lower molar about 3.5 mm (0.14 in) long, it is from the Maastrichtian of the Maevarano Formation in northwestern Madagascar. Details of its crown morphology indicate that it is a boreosphenidan, a member of the group that includes living marsupials and placental mammals. David W. Krause, who first described the tooth in 2001, interpreted it as a marsupial on the basis of five shared characters, but in 2003 Averianov and others noted that all those are shared by zhelestid placentals and favored a close relationship between UA 8699 and the Spanish zhelestid Lainodon. Krause used the tooth as evidence that marsupials were present on the southern continents (Gondwana) as early as the late Cretaceous and Averianov and colleagues proposed that the tooth represented another example of faunal exchange between Africa and Europe at the time.
Several mammals are known from the Mesozoic of Madagascar. The Bathonian Ambondro, known from a piece of jaw with three teeth, is the earliest known mammal with molars showing the modern, tribosphenic pattern that is characteristic of marsupial and placental mammals. Interpretations of its affinities have differed; one proposal places it in a group known as Australosphenida with other Mesozoic tribosphenic mammals from the southern continents (Gondwana) as well as the monotremes, while others favor closer affinities with northern (Laurasian) tribosphenic mammals or specifically with placentals. At least five species are known from the Maastrichtian, including a yet undescribed species known from a nearly complete skeleton that may represent a completely new group of mammals. The gondwanathere Lavanify, known from two teeth, is most closely related to other gondwanatheres found in India and Argentina. Two other teeth may represent another gondwanathere or a different kind of mammal. One molar fragment is one of the few known remains of a multituberculate mammal from Gondwana and another has been interpreted as either a marsupial or a placental.
Cairanoolithus is an oogenus of dinosaur egg which is found in Southwestern Europe. The eggs are large and spherical. Their outer surface is either smooth, or covered with a subdued pattern of ridges interspersed with pits and grooves. Multiple fossil egg clutches are known but the nest structure is unclear.
Oxlestes is an extinct mammal from the Late Cretaceous of Asia, more specifically from the Cenomanian of Uzbekistan. A carnivorous species of uncertain affinities, it is notable for its relatively large size, being among the largest of all Mesozoic mammals. Due to the limited amount of material, it has been considered a nomen dubium.
Zalambdalestidae is a clade of Asian eutherians occurring during the Cretaceous. Once classified as Glires, features like epipubic bones and various cranial elements have identified these animals as outside of Placentalia, representing thus a specialised clade of non-placental eutherians without any living descendants, and potentially rather different from modern placentals in at least reproductive anatomy.
Valentinella is a Late Cretaceous genus of eutherian mammal from France, Europe. It was originally known from some damaged lower and upper jaws, and was cautiously referred to the lainodontine zhelestid mammals. This identification was later questioned and even considered a nomen dubium by some researchers, and remains uncertain due to the scarcity and fragmentary nature of the fossils.
Azilestes is a genus of probable zhelestid eutherian mammal, a family consisting of small herbivores, that was discovered in the early Maastrichtian Grès de Labarre Formation of France. It is a monotypic genus, with only type species A. ragei being known. Only one specimen, the holotype described in 2021, is known. It consists of a partial dentary with teeth.
Gypsonictops is an extinct genus of leptictidan mammals of the family Gypsonictopidae, which was described in 1927 by George Gaylord Simpson. Species in this genus were small mammals and the first representatives of the order Leptictida, that appeared during the Upper Cretaceous.
Pan-Euungulata is a clade of placental mammals from grandorder Ferungulata, consisting of the taxon Euungulata [sensu stricto] and all taxa (species) more closely related to it than to any other living species.