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Multituberculata (commonly known as multituberculates, named for the multiple
tubercle In anatomy, a tubercle (literally 'small tuber', Latin for 'lump') is any round nodule, small eminence, or warty outgrowth found on external or internal organs of a plant or an animal. In plants A tubercle is generally a wart-like projection ...
s of their teeth) is an extinct
order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of ...
of rodent-like
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur ...
s with a fossil record spanning over 130 million years. They first appeared in the Middle
Jurassic The Jurassic ( ) is a geologic period and stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The Jurassic constitutes the middle period of ...
, and reached a peak diversity during the Late
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of ...
and
Paleocene The Paleocene, ( ) or Palaeocene, is a geological epoch that lasted from about 66 to 56 million years ago (mya). It is the first epoch of the Paleogene Period in the modern Cenozoic Era. The name is a combination of the Ancient Greek ''pala ...
. They eventually declined from the mid
Paleocene The Paleocene, ( ) or Palaeocene, is a geological epoch that lasted from about 66 to 56 million years ago (mya). It is the first epoch of the Paleogene Period in the modern Cenozoic Era. The name is a combination of the Ancient Greek ''pala ...
onwards, disappearing from the known fossil record in the late
Eocene The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', ...
. They are the most diverse order of
Mesozoic The Mesozoic Era ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last era of Earth's geological history, lasting from about , comprising the Triassic, Jurassic and Cretace ...
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 Jerboas (from ar, جربوع ') are hopping desert rodents found throughout North Africa and Asia, and are members of the family Dipodidae. They tend to live in hot deserts. When chased, jerboas can run at up to . Some species are preyed on b ...
-like hoppers. Multituberculates are usually placed as
crown A crown is a traditional form of head adornment, or hat, worn by monarchs as a symbol of their power and dignity. A crown is often, by extension, a symbol of the monarch's government or items endorsed by it. The word itself is used, partic ...
mammals outside either of the two main groups of living mammals—
Theria Theria (; Greek: , wild beast) is a subclass of mammals amongst the Theriiformes. Theria includes the eutherians (including the placental mammals) and the metatherians (including the marsupials) but excludes the egg-laying monotremes. ...
, including
placental Placental mammals (infraclass Placentalia ) 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 distinguishe ...
s and
marsupial Marsupials are any members of the mammalian infraclass Marsupialia. All extant marsupials are endemic to Australasia, Wallacea and the Americas. A distinctive characteristic common to most of these species is that the young are carried in ...
s, and MonotremataAgustí-Antón 2002, pp 3-4—but usually as closer to Theria than to monotremes. They are considered to be closely related to Euharamiyida and
Gondwanatheria Gondwanatheria is an extinct group of mammaliaforms that lived in parts of Gondwana, including Madagascar, India, South America, Africa and Antarctica during the Upper Cretaceous through the Paleogene (and possibly much earlier, if '' Allostaf ...
as part of
Allotheria Allotheria (meaning "other beasts", from the Greek , '–other and , '–wild animal) is an extinct branch of successful Mesozoic mammals. The most important characteristic was the presence of lower molariform teeth equipped with two longitudin ...
.


Description

The multituberculates had a cranial and dental anatomy superficially similar to rodents such as mice and rats, with cheek-teeth separated from the chisel-like front teeth by a wide tooth-less gap (the diasteme). Each cheek-tooth displayed several rows of small cusps (or
tubercle In anatomy, a tubercle (literally 'small tuber', Latin for 'lump') is any round nodule, small eminence, or warty outgrowth found on external or internal organs of a plant or an animal. In plants A tubercle is generally a wart-like projection ...
s, hence the name) that operated against similar rows in the teeth of the jaw; the exact homology of these cusps to
theria Theria (; Greek: , wild beast) is a subclass of mammals amongst the Theriiformes. Theria includes the eutherians (including the placental mammals) and the metatherians (including the marsupials) but excludes the egg-laying monotremes. ...
n ones is still a matter of debate. Unlike rodents, which have ever-growing teeth, multituberculates underwent dental replacement patterns typical to most mammals (though in at least some species the lower incisors continued to erupt long after the root's closure). Kielan-Jaworowska, Zofia, Richard L. Cifelli, and Zhe-Xi Luo (2005). ''Mammals from the Age of Dinosaurs: Origins, Evolution, and Structure ''
p. 299
/ref> Multituberculates are notable for the presence of a massive fourth lower premolar, the plagiaulacoid; other mammals, like
Plesiadapiformes Plesiadapiformes ("Adapid-like" or "near Adapiformes") is a group of Primates, a sister of the Dermoptera. While none of the groups normally directly assigned to this group survived, the group appears actually not to be literally extinct (in t ...
and
diprotodontia Diprotodontia (, from Greek "two forward teeth") is the largest extant order of marsupials, with about 155 species, including the kangaroos, wallabies, possums, koala, wombats, and many others. Extinct diprotodonts include the hippopotamus-sized ...
n
marsupials Marsupials are any members of the mammalian infraclass Marsupialia. All extant marsupials are endemic to Australasia, Wallacea and the Americas. A distinctive characteristic common to most of these species is that the young are carried in a po ...
, also have similar premolars in both upper and lower jaws, but in multituberculates this tooth is massive and the upper premolars aren't modified this way. In basal multituberculates all three lower premolars were plagiaulacoids, increasing in size posteriorly, but in
Cimolodonta Cimolodonta is a taxon of extinct mammals that lived from the Cretaceous to the Eocene. They were some of the more derived members of the extinct order Multituberculata. They probably lived something of a rodent-like existence until their ...
only the fourth lower premolar remained, with the third one remaining only as a vestigial peg-like tooth, and in several taxa like
taeniolabidoidea Taeniolabidoidea is a group of extinct mammals known from North America and Asia. They were the largest members of the extinct order Multituberculata, as well as the largest non-therian mammals. ''Lambdopsalis'' even provides direct fossil eviden ...
ns, the plagiaulacoid disappeared entirely or was reconverted into a molariform tooth. Unlike rodents and similar therians, multituberculates had a palinal jaw stroke (front-to-back), instead of a propalinal (back-to-front) or transverse (side-to-side) one; as a consequence, their jaw musculature and cusp orientation is radically different. Palinal jaw strokes are almost entirely absent in modern mammals (with the possible exception of the
dugong The dugong (; ''Dugong dugon'') is a marine mammal. It is one of four living species of the order Sirenia, which also includes three species of manatees. It is the only living representative of the once-diverse family Dugongidae; its closest m ...
), but are also present in
haramiyida Haramiyida ("thief" from Arabic الحرامية (al ḥarāmiyah), "thief, bandit") is a possibly polyphyletic order of mammaliaform cynodonts or mammals of controversial taxonomic affinites. Their teeth, which are by far the most common remai ...
ns, argyrolagoideans and
tritylodontid Tritylodontidae ("three-knob teeth", named after the shape of their cheek teeth) is an extinct family of small to medium-sized, highly specialized mammal-like cynodonts, bearing several mammalian traits like erect limbs, endothermy and details ...
s, the former historically united with multituberculates on that basis. Multituberculate mastication is thought to have operated in a two stroke cycle: first, food held in place by the last upper premolar was sliced by the bladelike lower pre-molars as the dentary moved orthally (upward). Then the lower jaw moved palinally, grinding the food between the molar cusp rows. The structure of the
pelvis The pelvis (plural pelves or pelvises) is the lower part of the trunk, between the abdomen and the thighs (sometimes also called pelvic region), together with its embedded skeleton (sometimes also called bony pelvis, or pelvic skeleton). The ...
in the Multituberculata suggests that they gave birth to tiny helpless, underdeveloped young, similar to modern
marsupial Marsupials are any members of the mammalian infraclass Marsupialia. All extant marsupials are endemic to Australasia, Wallacea and the Americas. A distinctive characteristic common to most of these species is that the young are carried in ...
s, such as kangaroos. However, a 2022 study reveals that they might actually have had long gestation periods like placentals. At least two lineages developed
hypsodonty Hypsodont is a pattern of dentition with high-crowned teeth and enamel extending past the gum line, providing extra material for wear and tear. Some examples of animals with hypsodont dentition are cows and horses; all animals that feed on gritty ...
, in which tooth enamel extends beyond the gumline: lambdopsalid
taeniolabidoidea Taeniolabidoidea is a group of extinct mammals known from North America and Asia. They were the largest members of the extinct order Multituberculata, as well as the largest non-therian mammals. ''Lambdopsalis'' even provides direct fossil eviden ...
ns and
sudamericid Sudamericidae is a family of gondwanathere mammals that lived during the late Cretaceous to Miocene. Its members include ''Lavanify'' and ''Vintana'' from the Cretaceous of Madagascar, ''Bharattherium'' (=''Dakshina'') from the Cretaceous of In ...
gondwanatheres. Studies published in 2018 demonstrated that multituberculates had relatively complex brains, some braincase regions even absent in therian mammals.


Evolution

Multituberculates first appear in the fossil record during the
Jurassic The Jurassic ( ) is a geologic period and stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The Jurassic constitutes the middle period of ...
period, and then survived and even dominated for over one hundred million years, longer than any other order of
mammaliforms Mammaliaformes ("mammalian forms") is a clade that contains the crown group mammals and their closest extinct relatives; the group radiated from earlier probainognathian cynodonts. It is defined as the clade originating from the most recent c ...
, including placental mammals. The earliest known multituberculates are from the
Middle Jurassic The Middle Jurassic is the second Epoch (geology), epoch of the Jurassic Period (geology), Period. It lasted from about 174.1 to 163.5 million years ago. Fossils of land-dwelling animals, such as dinosaurs, from the Middle Jurassic are relatively ...
(
Bathonian In the geologic timescale the Bathonian is an age and stage of the Middle Jurassic. It lasted from approximately 168.3 Ma to around 166.1 Ma (million years ago). The Bathonian Age succeeds the Bajocian Age and precedes the Callovian Age. ...
~166-168 million years ago) of England and Russia, including '' Hahnotherium'' ''and
Kermackodon ''Kermackodon'' is an extinct genus of allotherian mammal, known from the Middle Jurassic of England. It combines features of multituberculates with those of euharamyidans. The remains of type species, ''K. multicuspis'' were collected from Kirtl ...
'' from the
Forest Marble Formation The Forest Marble is a geological formation in England. Part of the Great Oolite Group, it dates to the late Bathonian stage of the Middle Jurassic.Weishampel, David B; et al. (2004). "Dinosaur distribution (Middle Jurassic, Europe)." In: Weisha ...
of England, and '' Tashtykia'' and '' Tagaria'' from the
Itat Formation The Itat Formation ( Russian: итатская свита) is a geologic formation in western Siberia. It was deposited in the Bajocian to Bathonian ages of the Middle Jurassic. Dinosaur remains are among the fossils that have been recovered from ...
of Russia. These forms are only known from isolated teeth, which bear close similarity to those of euharamyidans, which they are suspected to be closely related. During the Late Jurassic and Early Cretaceous, primitive multituberculates, collectively grouped into the
paraphyletic In taxonomy, a group is paraphyletic if it consists of the group's last common ancestor and most of its descendants, excluding a few monophyletic subgroups. The group is said to be paraphyletic ''with respect to'' the excluded subgroups. In ...
"
Plagiaulacida Plagiaulacida is a group of extinct multituberculate mammals. Multituberculates were among the most common mammals of the Mesozoic, "the age of the dinosaurs". Plagiaulacids are a paraphyletic grouping, containing all multituberculates that lie ...
" were abundant and widespread across
Laurasia Laurasia () was the more northern of two large landmasses that formed part of the Pangaea supercontinent from around ( Mya), the other being Gondwana. It separated from Gondwana (beginning in the late Triassic period) during the breakup of Pa ...
(including Europe, Asia and North America). During the
Aptian The Aptian is an age in the geologic timescale or a stage in the stratigraphic column. It is a subdivision of the Early or Lower Cretaceous Epoch or Series and encompasses the time from 121.4 ± 1.0 Ma to 113.0 ± 1.0 Ma (million years ag ...
stage of the Early Cretaceous, the advanced subgroup
Cimolodonta Cimolodonta is a taxon of extinct mammals that lived from the Cretaceous to the Eocene. They were some of the more derived members of the extinct order Multituberculata. They probably lived something of a rodent-like existence until their ...
appeared in North America, characterised by a reduced number of lower premolars, with a blade-like lower fourth premolar. By the early Late Cretaceous (
Cenomanian The Cenomanian is, in the ICS' geological timescale, the oldest or earliest age of the Late Cretaceous Epoch or the lowest stage of the Upper Cretaceous Series. An age is a unit of geochronology; it is a unit of time; the stage is a unit in ...
) Cimolodonta had replaced all other multituberculate lineages. During the Late Cretaceous, multituberculates experienced an
adaptive radiation In evolutionary biology, adaptive radiation is a process in which organisms diversify rapidly from an ancestral species into a multitude of new forms, particularly when a change in the environment makes new resources available, alters biotic in ...
, corresponding with a shift towards herbivory. Multituberculates reached their peak diversity during the early
Paleocene The Paleocene, ( ) or Palaeocene, is a geological epoch that lasted from about 66 to 56 million years ago (mya). It is the first epoch of the Paleogene Period in the modern Cenozoic Era. The name is a combination of the Ancient Greek ''pala ...
, shortly after the
Cretaceous–Paleogene extinction event The Cretaceous–Paleogene (K–Pg) extinction event (also known as the Cretaceous–Tertiary extinction) was a sudden mass extinction of three-quarters of the plant and animal species on Earth, approximately 66 million years ago. With the ...
, but declined from the mid Paleocene onwards, likely due to competition with placental mammals such as
rodent Rodents (from Latin , 'to gnaw') are mammals of the order Rodentia (), which are characterized by a single pair of continuously growing incisors in each of the upper and lower jaws. About 40% of all mammal species are rodents. They are n ...
s and
ungulate Ungulates ( ) are members of the diverse clade Ungulata which primarily consists of large mammals with hooves. These include odd-toed ungulates such as horses, rhinoceroses, and tapirs; and even-toed ungulates such as cattle, pigs, giraffes, ...
s, the group finally became extinct in the Late
Eocene The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', ...
. There are some isolated records of multituberculates from the Southern Hemisphere, including the cimolodontan ''
Corriebaatar ''Corriebaatar'' is the type and only genus of Corriebaataridae, a family of multituberculate mammals. It contains the single species ''Corriebaatar marywaltersae'' and represents the first evidence of Australian multituberculates Fossils found ...
'' from the Early Cretaceous of Australia, and fragmentary remains from the Late Cretaceous Maevarano Formation of Madagascar. The family
Ferugliotheriidae Ferugliotheriidae is one of three known families in the order Gondwanatheria, an enigmatic group of extinct mammals. Gondwanatheres have been classified as a group of uncertain affinities or as members of Multituberculata, a major extinct mammal ...
from the Late Cretaceous of South America, traditionally considered gondwanatherians, may actually be cimolodontan multituberculates. During the Late Cretaceous and Paleocene the multituberculates radiated into a wide variety of
morphotype In biology, polymorphism is the occurrence of two or more clearly different morphs or forms, also referred to as alternative '' phenotypes'', in the population of a species. To be classified as such, morphs must occupy the same habitat at the ...
s, including the squirrel-like arboreal ptilodonts. The peculiar shape of their last lower
premolar The premolars, also called premolar teeth, or bicuspids, are transitional teeth located between the canine and molar teeth. In humans, there are two premolars per quadrant in the permanent set of teeth, making eight premolars total in the mouth ...
is their most outstanding feature. These teeth were larger and more elongated than the other cheek-teeth and had an
occlusive In phonetics, an occlusive, sometimes known as a stop, is a consonant sound produced by occluding (i.e. blocking) airflow in the vocal tract, but not necessarily in the nasal tract. The duration of the block is the ''occlusion'' of the consonan ...
surface forming a serrated slicing blade. Though it can be assumed that this was used for crushing seeds and nuts, it is believed that most small multituberculates also supplemented their diet with insects, worms, and fruits. Tooth marks attributed to multituberculates are known on ''
Champsosaurus ''Champsosaurus'' is an extinct genus of crocodile-like choristodere reptile, known from the Late Cretaceous and early Paleogene periods of North America and Europe (Campanian-Paleocene). The name ''Champsosaurus'' is thought to come from , () s ...
'' fossils, indicating that at least some of these mammals were
scavenger Scavengers are animals that consume dead organisms that have died from causes other than predation or have been killed by other predators. While scavenging generally refers to carnivores feeding on carrion, it is also a herbivorous feedin ...
s. A ptilodont that thrived in North America was ''
Ptilodus ''Ptilodus'' (meaning "soft-haired") is a genus of mammals from the extinct order of Multituberculata, and lived during the Paleocene in North America. ''Ptilodus'' was a relatively large multituberculate of in length, which is about the same s ...
''. Thanks to the well-preserved ''Ptilodus'' specimens found in the
Bighorn Basin The Bighorn Basin is a plateau region and intermontane basin, approximately 100 miles (160 km) wide, in north-central Wyoming in the United States. It is bounded by the Absaroka Range on the west, the Pryor Mountains on the north, the Bigho ...
,
Wyoming Wyoming () is a state in the Mountain West subregion of the Western United States. It is bordered by Montana to the north and northwest, South Dakota and Nebraska to the east, Idaho to the west, Utah to the southwest, and Colorado to t ...
, we know that these multituberculates were able to abduct and adduct their big toes, and thus that their foot mobility was similar to that of modern squirrels, which descend trees head first. Another group of multituberculates, the taeniolabids, were heavier and more massively built, indicating that they lived a fully terrestrial life. The largest specimens weighted probably as much as 100 kg, making them comparable in size to large rodents like ''
Castoroides ''Castoroides'' (Latin: "beaver" (castor), "like" (oides)), or giant beaver, is an extinct genus of enormous, bear-sized beavers that lived in North America during the Pleistocene. Two species are currently recognized, ''C. dilophidus'' in the S ...
''. They reached their highest diversity in Asia during the late Cretaceous and Paleocene, which suggests that they originated from there.


Classification

Multituberculate is generally placed in the
Allotheria Allotheria (meaning "other beasts", from the Greek , '–other and , '–wild animal) is an extinct branch of successful Mesozoic mammals. The most important characteristic was the presence of lower molariform teeth equipped with two longitudin ...
alongside Euharamiyida, a clade of mammals known from the Middle Jurassic to Early Cretaceous of the Asia and possibly Europe that several morphological similarities to multituberculates.
Gondwanatheria Gondwanatheria is an extinct group of mammaliaforms that lived in parts of Gondwana, including Madagascar, India, South America, Africa and Antarctica during the Upper Cretaceous through the Paleogene (and possibly much earlier, if '' Allostaf ...
is a monophyletic group of allotherians that was diverse in the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', ...
of South America, India,
Madagascar Madagascar (; mg, Madagasikara, ), officially the Republic of Madagascar ( mg, Repoblikan'i Madagasikara, links=no, ; french: République de Madagascar), is an island country in the Indian Ocean, approximately off the coast of East Afric ...
and possibly Africa and occurs onwards into the
Paleogene The Paleogene ( ; also spelled Palaeogene or Palæogene; informally Lower Tertiary or Early Tertiary) is a geologic period and system that spans 43 million years from the end of the Cretaceous Period million years ago ( Mya) to the beginning o ...
of South America and
Antarctica Antarctica () is Earth's southernmost and least-populated continent. Situated almost entirely south of the Antarctic Circle and surrounded by the Southern Ocean, it contains the geographic South Pole. Antarctica is the fifth-largest cont ...
. Their placement within Allotheria is highly controversial, with some phylogenies recovering the group as deeply nested within multituberculates, while others recover them as a distinct branch of allotherians separate from multituberculates. In their 2001 study, Kielan-Jaworowska and
Hurum Hurum was a municipality in Buskerud county, Norway. As of 1 January 2020 Hurum has merged with the municipalities of Røyken and Asker to form the new Asker Municipality located in the newly formed Viken county. The administrative centre of the ...
found that most multituberculates could be referred to two suborders: "
Plagiaulacida Plagiaulacida is a group of extinct multituberculate mammals. Multituberculates were among the most common mammals of the Mesozoic, "the age of the dinosaurs". Plagiaulacids are a paraphyletic grouping, containing all multituberculates that lie ...
" and
Cimolodonta Cimolodonta is a taxon of extinct mammals that lived from the Cretaceous to the Eocene. They were some of the more derived members of the extinct order Multituberculata. They probably lived something of a rodent-like existence until their ...
. The exception is the genus ''
Arginbaatar ''Arginbaatar'' is a genus of extinct mammal from the Lower Cretaceous of Mongolia. It was a member of the Multituberculata, an order which is also extinct. It belongs to the family Arginbaataridae (Hahn & Hahn 1983). The genus ''Arginbaatar'' w ...
'', which shares characteristics with both groups. "Plagiaulacida" is
paraphyletic In taxonomy, a group is paraphyletic if it consists of the group's last common ancestor and most of its descendants, excluding a few monophyletic subgroups. The group is said to be paraphyletic ''with respect to'' the excluded subgroups. In ...
, representing the more primitive evolutionary grade. Its members are the more basal Multituberculata. Chronologically, they ranged from perhaps the Middle Jurassic until the mid-
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of ...
. This group is further subdivided into three informal groupings: the allodontid line, the paulchoffatiid line, and the plagiaulacid line.
Cimolodonta Cimolodonta is a taxon of extinct mammals that lived from the Cretaceous to the Eocene. They were some of the more derived members of the extinct order Multituberculata. They probably lived something of a rodent-like existence until their ...
is, apparently, a natural (
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gr ...
) suborder. This includes the more derived Multituberculata, which have been identified from the lower Cretaceous to the
Eocene The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', ...
. The superfamilies
Djadochtatherioidea Djadochtatherioidea is a group of extinct mammals known from the upper Cretaceous of Central Asia. They were members of an also extinct order called Multituberculata. These were generally somewhat rodent-like creatures, who scurried around dur ...
,
Taeniolabidoidea Taeniolabidoidea is a group of extinct mammals known from North America and Asia. They were the largest members of the extinct order Multituberculata, as well as the largest non-therian mammals. ''Lambdopsalis'' even provides direct fossil eviden ...
,
Ptilodontoidea Ptilodontoidea is a group of extinct mammals from the Northern Hemisphere. They were generally small, somewhat rodent-like creatures of the extinct order Multituberculata. Some of these genera boast a great many species, though remains are gen ...
are recognized, as is the
Paracimexomys group ''Paracimexomys'' is a genus of extinct mammals in the also extinct Multituberculata order. ''Paracimexomys'' lived during the Cretaceous period. The few fossils remains come from North America. Some Romanian fossils were also tentatively assign ...
. Additionally, there are the families
Cimolomyidae Cimolomyidae is a family of fossil mammal within the extinct order Multituberculata. Representatives are known from the Upper Cretaceous and the Paleocene of North America and perhaps Mongolia. The family is part of the suborder Cimolodonta. Oth ...
, Boffiidae, Eucosmodontidae,
Kogaionidae Kogaionidae is a family of fossil mammals within the extinct order Multituberculata. Representatives are known from the Upper Cretaceous and the Paleocene of Europe. Having started as island endemics on Hateg Island during the Upper Cretaceous, ...
, Microcosmodontidae and the two genera ''
Uzbekbaatar ''Uzbekbaatar'' is a genus of extinct mammal from the Upper Cretaceous of Uzbekistan. It was a member of the extinct order Multituberculata within the suborder Cimolodonta, though its further affinities are unclear. The genus was named by Kiel ...
'' and ''
Viridomys ''Viridomys'' is a genus of extinct mammal from the Upper Cretaceous of North America. It was a member of the extinct order of Multituberculata, and lived during the Mesozoic, also known as the "age of the dinosaurs." It's within the suborder o ...
''. More precise placement of these types awaits further discoveries and analysis.Dykes ''Multituberculata (Cope 1884)''


Taxonomy

Based on the combined works of Mikko's Phylogeny ArchiveMikko's Phylogeny Archive
and Paleofile.com. Suborder †
Plagiaulacida Plagiaulacida is a group of extinct multituberculate mammals. Multituberculates were among the most common mammals of the Mesozoic, "the age of the dinosaurs". Plagiaulacids are a paraphyletic grouping, containing all multituberculates that lie ...
Simpson 1925 * Genus ?†'' Argillomys'' Cifelli, Gordon & Lipka 2013 ** Species †'' Argillomys marylandensis'' Cifelli, Gordon & Lipka 2013 * Genus ?†''
Janumys ''Janumys'' is a genus of extinct mammal of the middle Cretaceous. It was a member of the order of Multituberculata (also extinct). It lived in North America during the Mesozoic era, also known as the "age of the reptiles." It has been provisio ...
'' Eaton & Cifelli 2001 ** Species †''
Janumys erebos ''Janumys'' is a genus of extinct mammal of the middle Cretaceous. It was a member of the order of Multituberculata (also extinct). It lived in North America during the Mesozoic era, also known as the "age of the reptiles." It has been provisio ...
'' Eaton & Cifelli 2001 * Super family †Allodontoidea Marsh 1889 ** Genus †?''
Glirodon ''Glirodon'' is a genus of extinct mammal from the Upper Jurassic. It was a relatively early member of the also-extinct order of Multituberculata, suborder "Plagiaulacida". These mammals lived in North America during the Mesozoic, also known ...
'' Engelmann & Callison, 2001 *** Species †'' G. grandis'' Engelmann & Callison, 2001 ** Family † Arginbaataridae Hahn & Hahn, 1983 *** Genus †''
Arginbaatar ''Arginbaatar'' is a genus of extinct mammal from the Lower Cretaceous of Mongolia. It was a member of the Multituberculata, an order which is also extinct. It belongs to the family Arginbaataridae (Hahn & Hahn 1983). The genus ''Arginbaatar'' w ...
'' Trofimov, 1980 **** Species †'' A. dmitrievae'' Trofimov, 1980 ** Family † Zofiabaataridae Bakker, 1992 *** Genus †''
Zofiabaatar ''Zofiabaatar'' is a genus of extinct mammal from the Upper Jurassic period. It was a relatively early member of the extinct order Multituberculata within the suborder "Plagiaulacida". It lived in North America along with dinosaurs such ...
'' Bakker & Carpenter, 1990 **** Species †'' Z. pulcher'' Bakker & Carpenter, 1990 ** Family † Allodontidae Marsh, 1889 *** Genus †'' Passumys'' Cifelli, Davis & Sames 2014 **** Species †'' Passumys angelli'' Cifelli, Davis & Sames 2014 *** Genus †'' Ctenacodon'' Marsh, 1879 **** Species †'' C. serratus'' Marsh, 1879 **** Species †'' C. nanus'' Marsh, 1881 **** Species †'' C. laticeps'' (Marsh, 1881) 'Allodon laticeps'' Marsh 1881 **** Species †'' C. scindens'' Simpson, 1928 *** Genus †'' Psalodon'' Simpson, 1926 **** Species †'' P. potens'' (Marsh, 1887) 'Ctenacodon potens'' Marsh 1887**** Species †'' P. fortis'' (Marsh, 1887) Simpson 1929 'Allodon fortis'' Marsh 1887**** Species †'' P. marshi'' Simpson, 1929 * Super family † Paulchoffatioidea Hahn 1969 sensu Hahn & Hahn 2003 ** Genus ?†''
Mojo Mojo may refer to: *Mojo (African-American culture), a magical charm bag used in voodoo Arts, entertainment and media Film and television * MOJO HD, an American television network * ''Mojo'' (play), by Jez Butterworth, made into a 1997 film * '' ...
'' Hahn, LePage & Wouters 1987 *** Species †''
Mojo usuratus Mojo may refer to: *Mojo (African-American culture), a magical charm bag used in voodoo Arts, entertainment and media Film and television * MOJO HD, an American television network * ''Mojo'' (play), by Jez Butterworth, made into a 1997 film * '' ...
'' Hahn, LePage & Wouters 1987 ** Genus ?†'' Rugosodon'' Yuan et al., 2013 *** Species †''
Rugosodon eurasiaticus ''Rugosodon'' is an extinct genus of multituberculate (rodent-like) mammals from eastern China that lived 160 million years ago during the Jurassic period. The discovery of its type species and currently only known species ''Rugosodon eurasiat ...
'' Yuan et al., 2013 ** Family †
Pinheirodontidae Pinheirodontidae is a poorly known family of fossil mammals which belong to the informal suborder "Plagiaulacida" within the order Multituberculata. Remains are known from the Late Jurassic to Early Cretaceous of Europe, (predominantly Portuga ...
Hahn & Hahn, 1999 *** Genus †''
Bernardodon ''Bernardodon'' was a small, Lower Cretaceous mammal from Portugal. It is part of the extinct order Multituberculata, living at the same time as the dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first ...
'' Hahn & Hahn, 1999 **** Species †'' B. atlanticus'' Hahn & Hahn, 1999 **** Species †''B. sp.'' Hahn & Hahn, 1999 *** Genus †'' Cantalera'' Badiola, Canudo & Cuenca-Bescos, 2008 **** Species †'' Cantalera abadi'' Badiola, Canudo & Cuenca-Bescos, 2008 *** Genus †''
Ecprepaulax ''Ecprepaulax'' is a Lower Cretaceous mammal from Portugal. It was a member of the also extinct order Multituberculata and shared the world with dinosaurs. It lies within the suborder "Plagiaulacida" and family Pinheirodontidae. The genus ''Ecp ...
'' Hahn & Hahn, 1999 **** Species †'' E. anomala'' Hahn & Hahn, 1999 *** Genus †''
Gerhardodon ''Gerhardodon'' is an extinct genus of mammal from the Lower Cretaceous of southern England. It was a member of the also extinct order of Multituberculata, and lived with such dinosaurs as ''Iguanodon''. It lies within the suborder "Plagiaulacid ...
'' Kielan-Jaworowska & Ensom, 1992 **** Species †'' G. purbeckensis'' Kielan-Jaworowska & Ensom, 1992 *** Genus †''
Iberodon ''Iberodon'' is a small, extinct mammal of the Lower Cretaceous from Portugal. It was a member of the also extinct order Multituberculata, and led its obscure and plant-eating existence in the company of dinosaurs. It lies within the suborder " ...
'' Hahn & Hahn, 1999 **** Species †'' I. quadrituberculatus'' Hahn & Hahn, 1999 *** Genus †''
Lavocatia ''Lavocatia'' is a genus of extinct mammal from the Lower Cretaceous of Spain. It was a member of the also extinct order Multituberculata, and lived alongside of dinosaurs. Like most Mesozoic mammals, it was a shrewish-sized animal. It's in the ...
'' Canudo & Cuenca-Bescós, 1996 **** Species †'' L. alfambrensis'' Canudo & Cuenca-Bescós, 1996 *** Genus †''
Pinheirodon ''Pinheirodon'' is a genus of extinct mammal from Portugal. It is a member of the also extinct order of Multituberculata, and shared the world with dinosaurs. It is placed in the suborder "Plagiaulacida" and family Pinheirodontidae. The species ...
'' Hahn & Hahn, 1999 **** Species †'' P. pygmaeus'' Hahn & Hahn, 1999 **** Species †'' P. vastus'' Hahn & Hahn, 1999 ** Family †
Paulchoffatiidae Paulchoffatiidae is a family of extinct mammals that lived predominantly during the Upper Jurassic period, though a couple of genera are known from the Early Cretaceous. Fossils have been reported from Europe (Portugal, Spain, Germany and Engla ...
Hahn, 1969 *** Genus ?†''
Galveodon ''Galveodon'' is an extinct mammal of the Lower Cretaceous. It was a relatively early representative of the also extinct order of Multituberculata. It lived during 'the age of the dinosaurs'. It's within the suborder "Plagiaulacida" and famil ...
'' Hahn & Hahn, 1992 **** Species †'' G. nannothus'' Hahn & Hahn, 1992 *** Genus ?†''
Sunnyodon ''Sunnyodon'' is a genus of tiny, extinct mammal, probably of the Lower Cretaceous. Found in what is now southern England and Denmark, it was a relatively early member of the extinct order of Multituberculata. It is part of the suborder Pl ...
'' Kielan-Jaworowska & Ensom, 1992 **** Species †'' S. notleyi'' Kielan-Jaworowska & Ensom, 1992 *** subfamily †
Paulchoffatiinae Paulchoffatiidae is a family of extinct mammals that lived predominantly during the Upper Jurassic period, though a couple of genera are known from the Early Cretaceous. Fossils have been reported from Europe (Portugal, Spain, Germany and Englan ...
Hahn, 1971 **** Genus †''
Paulchoffatia ''Paulchoffatia'' is a genus of extinct mammal of the Upper Jurassic - Lower Cretaceous. It was a relatively early member of the also extinct order Multituberculata, within the suborder "Plagiaulacida" and family Paulchoffatiidae. It lived i ...
'' Kühne, 1961 ***** Species †'' P. delgador'' Kühne, 1961 **** Genus †''
Pseudobolodon ''Pseudobolodon'' was a relatively early member of the also extinct order of mammals, Multituberculata. It lived in Portugal during the Upper Jurassic, part of the "age of the dinosaurs." It lies within the suborder "Plagiaulacida" and fami ...
'' Hahn, 1977 ***** Species †'' P. oreas'' Hahn, 1977 ***** Species †'' P. krebsi'' Hahn & Hahn, 1994 **** Genus †''
Henkelodon ''Henkelodon'' was a small mammal of the Upper Jurassic. It was a relatively early member of the extinct order Multituberculata. ''Henkelodon'' was a European herbivore that lived during the "age of the dinosaurs". It lies within the suborder " ...
'' Hahn, 1987 ***** Species †'' H. naias'' Hahn, 1987 **** Genus †''
Guimarotodon ''Guimarotodon'' is an extinct mammal of the Upper Jurassic. It was a relatively early member of the also extinct order of Multituberculata. It made its living nibbling plants as great big, and small, dinosaurs roamed the world. (For the te ...
'' Hahn, 1969 ***** Species †'' G. leiriensis'' Hahn, 1969 **** Genus †''
Meketibolodon ''Meketibolodon'' is a genus of extinct mammal from the Kimmeridgian (Upper Jurassic) Camadas de Guimarota of Guimarota, Portugal. It was a relatively early member of the also extinct order Multituberculata, suborder Plagiaulacida, fam ...
'' (Hahn, 1978) Hahn, 1993 ***** Species †'' M. robustus'' (Hahn, 1978) Hahn, 1993 'Pseudobolodon robusutus'' Hahn 1978**** Genus †''
Plesiochoffatia ''Plesiochoffatia'' is an extinct mammal of the Upper Jurassic. It was a relatively early member of the also extinct order Multituberculata. It was a resident of Portugal during the "age of the dinosaurs." It's in the suborder "Plagiaulacid ...
'' Hahn & Hahn, 1999 'Parachoffatia'' Hahn & Hahn 1998 non Mangold 1970***** Species †'' P. thoas'' (Hahn & Hahn, 1998) Hahn & Hahn 1999 'Parachoffatia thoa'' Hahn & Hahn 1998***** Species †'' P. peparethos'' (Hahn & Hahn, 1998) Hahn & Hahn 1999 'Parachoffatia peparethos'' Hahn & Hahn 1998***** Species †'' P. staphylos'' (Hahn & Hahn, 1998) Hahn & Hahn 1999 'Parachoffatia staphylos'' Hahn & Hahn 1998**** Genus †''
Xenachoffatia ''Xenachoffatia'' is a small Jurassic mammal from Portugal. It was a relatively early member of the also extinct order of Multituberculata. It lived during "the age of the dinosaurs" and belongs to the suborder Plagiaulacida, family Paulcho ...
'' Hahn & Hahn, 1998 ***** Species †'' X. oinopion'' Hahn & Hahn, 1998 **** Genus †''
Bathmochoffatia ''Bathmochoffatia'' is an extinct mammal of the Upper Jurassic. It was a relatively early member of the also extinct order Multituberculata. It lived in Portugal at about the same time as the far more famous dinosaur, Allosaurus. It is in the ...
'' Hahn & Hahn, 1998 ***** Species †'' B. hapax'' Hahn & Hahn, 1998 **** Genus †''
Kielanodon ''Kielanodon'' is an extinct mammal of the Portuguese Upper Jurassic. It was a relatively early member of the also extinct order of Multituberculata. It eked out a living during the Mesozoic era, also known as the "Age of the Dinosaurs." It is ...
'' Hahn, 1987 ***** Species †'' K. hopsoni'' Hahn, 1987 **** Genus †''
Meketichoffatia ''Meketichoffatia'' was a small mammal from the Upper Jurassic of Portugal. It was a relatively early member of the extinct order Multituberculata. It lived at the same time as dinosaurs such as Allosaurus. It's within the suborder "Plagia ...
'' Hahn, 1993 ***** Species †'' M. krausei'' Hahn, 1993 **** Genus †'' Renatodon'' Hahn, 2001 ***** Species †'' Renatodon amalthea'' Hahn, 2001 *** Subfamily † Kuehneodontinae Hahn, 1971 **** Genus †''
Kuehneodon ''Kuehneodon'' is a genus of extinct mammal of the Upper Jurassic - Lower Cretaceous of Europe. It was a relatively early member of the also extinct order of Multituberculata. Members of this genus lived alongside such dinosaurs as Allosaur ...
'' Hahn, 1969 ***** Species †'' K. dietrichi'' Hahn, 1969 ***** Species †'' K. barcasensis'' Hahn & Hahn, 2001 ***** Species †'' K. dryas'' Hahn, 1977 ***** Species †'' K. guimarotensis'' Hahn, 1969 ***** Species †'' K. hahni'' Antunes, 1988 ***** Species †'' K. simpsoni'' Hahn, 1969 ***** Species †'' K. uniradiculatus'' Hahn, 1978 * Super family †Plagiaulacoidea Ameghino, 1894 ** Family †
Plagiaulacidae Plagiaulacidae is a family of fossil mammals within the order Multituberculata. Remains are known from the Upper Jurassic and earliest Cretaceous of North America and Europe. They were among the more derived representatives of the informal subo ...
Gill, 1872 sensu Kielan-Jaworowska & Hurum, 2001 olodontidae Osborn 1887 *** Genus ?†''
Morrisonodon ''Morrisonodon brentbaatar'' is an extinct multituberculate mammal from the Upper Jurassic Morrison Formation of North America. ''M. brentbaatar'' was described by Robert T. Bakker in 1998, who originally placed it in the genus ''Ctenacodon''. ...
'' Hahn & Hahn, 2004 **** Species †''
Morrisonodon brentbaatar ''Morrisonodon brentbaatar'' is an extinct multituberculate mammal from the Upper Jurassic Morrison Formation of North America. ''M. brentbaatar'' was described by Robert T. Bakker in 1998, who originally placed it in the genus ''Ctenacodon''. ...
'' (Bakker, 1998) Hahn & Hahn, 2004 'Ctenacodon brentbaatar'' Bakker, 1998*** Genus †''
Plagiaulax ''Plagiaulax'' is a genus of mammal from the Lower Cretaceous of Europe. It was a member of the also extinct order Multituberculata, and shared the world with dinosaurs. It is of the suborder "Plagiaulacida" and family Plagiaulacidae. The genus ...
'' Falconer, 1857 **** Species †'' P. becklesii'' Falconer, 1857 **** Species †'' P. dawsoni'' Woodward, 1891 'Plioprion dawsoni'' Woodward, 1891; ''Loxaulax dawsoni'' (Woodward, 1891) Sloan, 1979*** Genus †''
Bolodon ''Bolodon'' ("lump tooth" from Ancient Greek βῶλος (bôlos), “clod, lump” + ὀδών (odṓn), "tooth") is a genus of extinct mammal from the Lower Cretaceous of Europe and North America. It was a member of the extinct order of Multi ...
'' Owen, 1871 'Plioprion''_Cope,_1884.html" ;"title="Plioprion.html" ;"title="' 'Plioprion''_Cope,_1884">Plioprion.html"_;"title="'Plioprion">'Plioprion''_Cope,_1884****_Species_†'' 'Plioprion''_Cope,_1884">Plioprion.html"_;"title="'Plioprion">'Plioprion''_Cope,_1884****_Species_†''Bolodon_crassidens">B._crassidens''_Owen,_1871 ****_Species_†''Bolodon_falconeri.html" ;"title="Bolodon_crassidens.html" ;"title="Plioprion">'Plioprion'' Cope, 1884">Plioprion.html" ;"title="'Plioprion">'Plioprion'' Cope, 1884**** Species †''Bolodon crassidens">B. crassidens'' Owen, 1871 **** Species †''Bolodon falconeri">B. falconeri'' Owen, 1871 [''Pligiaulax falconeri'' Owen, 1871; ''Plioprion falconeri'' (Owen, 1871)] **** Species †''Bolodon hydei, B. hydei'' Cifelli, Davis & Sames, 2014 **** Species †''Bolodon minor, B. minor'' Falconer, 1857 [''Pligiaulax minor'' Falconer, 1857; ''Plioprion minor'' (Falconer, 1857)] **** Species †'' Bolodon osborni, B. osborni'' Simpson, 1928 'Plioprion osborni'' (Simpson, 1928); ''Ctenacodon osborni'' Simpson, 1928**** Species ?†'' B. elongatus'' Simpson, 1928 * Family †
Eobaataridae Eobaataridae is a family of fossil mammals Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their youn ...
Kielan-Jaworowska, Dashzeveg & Trofimov, 1987 ** Genus †''
Eobaatar ''Eobaatar'' is a genus of extinct mammal from the Lower Cretaceous of Mongolia, Spain and England. A member of the also extinct order Multituberculata, it lies within the suborder Plagiaulacida and family Eobaataridae. The genus ''Eobaatar'' wa ...
'' Kielan-Jaworowska, Dashzeveg & Trofimov, 1987 *** Species †'' E. clemensi'' Sweetman, 2009 *** Species †'' E. hispanicus'' Hahn & Hahn, 1992 *** Species †'' E. magnus'' Kielan-Jaworowska, Dashzeveg & Trofimov, 1987 *** Species †'' E. minor'' Kielan-Jaworowska, Dashzeveg & Trofimov, 1987 *** Species †'' E. pajaronensis'' Hahn & Hahn, 2001 ** Genus †''
Hakusanobaatar ''Hakusanobaatar'' is an extinct genus of eobaatarid multituberculate Multituberculata (commonly known as multituberculates, named for the multiple tubercles of their teeth) is an extinct order of rodent-like mammals with a fossil record spa ...
'' Kusuhashi et al., 2008 *** Species †'' H. matsuoi'' Kusuhashi et al., 2008 ** Genus †''
Heishanobaatar ''Heishanobaatar'' ("Heishan hero" from Chinese 黑山 (Hēishān), " Heishan" + Mongolian baatar, "hero") is an extinct genus of eobaatarid multituberculate which existed in Shahai and Fuxin formations, northeastern China, during the early C ...
'' Kusuhashi et al., 2010 *** Species †'' H. triangulus'' Kusuhashi et al., 2010 ** Genus †'' Iberica'' Badiola et al., 2011 *** Species †''
Iberica hahni ''Iberica'' is an extinct genus of eobaatarid or a possible plagiaulacid multituberculate which existed in what is now Galve, Spain, during the early Cretaceous (late Hauterivian-early Barremian age). It was first named by Ainara Badiola, ...
'' Badiola et al., 2011 ** Genus †''
Liaobaatar ''Liaobaatar changi'' is a multituberculate Multituberculata (commonly known as multituberculates, named for the multiple tubercles of their teeth) is an extinct order of rodent-like mammals with a fossil record spanning over 130 million years ...
'' Kusuhashi et al., 2009 *** Species †'' L. changi'' Kusuhashi et al., 2009 ** Genus †'' Loxaulax'' Simpson, 1928 'Parendotherium''_Crusafont_Pairó_&_Adrover,_1966.html" ;"title="Parendotherium.html" ;"title="' 'Parendotherium''_Crusafont_Pairó_&_Adrover,_1966">Parendotherium.html"_;"title="'Parendotherium">'Parendotherium''_Crusafont_Pairó_&_Adrover,_1966***_Species_†''Loxaulax.html" ;"title="Parendotherium">'Parendotherium'' Crusafont Pairó & Adrover, 1966">Parendotherium.html" ;"title="'Parendotherium">'Parendotherium'' Crusafont Pairó & Adrover, 1966*** Species †''Loxaulax">L. valdensis'' (Woodward, 1911) Simpson, 1928[''Dipriodon valdensis'' Woodward, 1911] *** Species †''Loxaulax, L. herreroi'' (Crusafont Pairó & Adrover, 1966) [''Parendotherium herreroi'' Crusafont Pairó & Adrover 1966] ** Genus †''Monobaatar'' Kielan-Jaworowska, Dashzeveg & Trofimov, 1987 *** Species †'' M. mimicus'' Kielan-Jaworowska, Dashzeveg & Trofimov, 1987 ** Genus †''
Sinobaatar ''Sinobaatar'' is a genus of extinct mammal from the Lower Cretaceous of China. It is categorized within the also extinct order Multituberculata and among these it belongs to the plagiaulacid lineage (a possible infraorder). ''Sinobaatar'' was ...
'' Hu & Wang, 2002 *** Species †'' S. lingyuanensis'' Hu & Wang, 2002 *** Species †'' S. xiei'' Kusuhashi et al., 2009 *** Species †'' S. fuxinensis'' Kusuhashi et al., 2009 ** Genus †'' Tedoribaatar'' Kusuhashi et al., 2008 *** Species †'' T. reini'' Kusuhashi et al., 2008 ** Genus †''
Teutonodon Pinheirodontidae is a poorly known family of fossil mammals which belong to the informal suborder "Plagiaulacida" within the order Multituberculata. Remains are known from the Late Jurassic to Early Cretaceous of Europe, (predominantly Portuga ...
'' Martin et al., 2016 *** Species †''
Teutonodon langenbergensis Pinheirodontidae is a poorly known family of fossil mammals which belong to the informal suborder "Plagiaulacida" within the order Multituberculata. Remains are known from the Late Jurassic to Early Cretaceous of Europe, (predominantly Portuga ...
'' Martin et al. 2016 * Family † Albionbaataridae Kielan-Jaworowska & Ensom, 1994 ** Genus †'' Albionbaatar'' Kielan-Jaworowska & Ensom, 1994 *** Species †'' A. denisae'' Kielan-Jaworowska & Ensom, 1994 ** Genus †'' Kielanobaatar'' Kusuhashi et al., 2010 *** Species †'' K. badaohaoensis'' ''Kusuhashi et al.'', 2010 ** Genus †'' Proalbionbaatar'' Hahn & Hahn, 1998 *** Species †'' P. plagiocyrtus'' Hahn & Hahn, 1998 * Suborder †
Gondwanatheria Gondwanatheria is an extinct group of mammaliaforms that lived in parts of Gondwana, including Madagascar, India, South America, Africa and Antarctica during the Upper Cretaceous through the Paleogene (and possibly much earlier, if '' Allostaf ...
McKenna 1971 ondwanatheroidea Krause & Bonaparte 1993** Family † Groeberiidae Patterson, 1952 *** Genus †'' Groeberia'' Patterson 1952 **** Species †'' G. minoprioi'' Ryan Patterson, 1952 **** Species †'' G. pattersoni'' G. G. Simpson, 1970 *** Genus †'' Klohnia'' Flynn & Wyss 1999 **** Species †'' K. charrieri'' Flynn & Wyss 1999 **** Species †'' K. major'' Goin et al., 2010 *** Genus ?†'' Epiklohnia'' Goin et al., 2010 **** Species †'' Epiklohnia verticalis'' Goin et al., 2010 *** Genus ?†'' Praedens'' Goin et al., 2010 **** Species †'' Praedens aberrans'' Goin et al., 2010 ** Family †
Ferugliotheriidae Ferugliotheriidae is one of three known families in the order Gondwanatheria, an enigmatic group of extinct mammals. Gondwanatheres have been classified as a group of uncertain affinities or as members of Multituberculata, a major extinct mammal ...
Bonaparte, 1986 *** Genus †'' Ferugliotherium'' Bonaparte, 1986a 'Vucetichia''_Bonaparte,_1990.html" ;"title="Vucetichia.html" ;"title="' 'Vucetichia''_Bonaparte,_1990">Vucetichia.html"_;"title="'Vucetichia">'Vucetichia''_Bonaparte,_1990****_†''Ferugliotherium_windhauseni.html" ;"title="Vucetichia">'Vucetichia'' Bonaparte, 1990">Vucetichia.html" ;"title="'Vucetichia">'Vucetichia'' Bonaparte, 1990**** †''Ferugliotherium windhauseni">Vucetichia">'Vucetichia'' Bonaparte, 1990">Vucetichia.html" ;"title="'Vucetichia">'Vucetichia'' Bonaparte, 1990**** †''Ferugliotherium windhauseni'' Bonaparte, 1986a [''Vucetichia gracilis'' Bonaparte, 1990] *** Genus †''Trapalcotherium'' Rougier et al., 2008 **** †''Trapalcotherium matuastensis'' Rougier et al., 2008 ** Family †Sudamericidae Scillato-Yané & Pascual, 1984 ondwanatheridae Bonaparte, 1986; Patagonidae Pascual & Carlini, 1987*** Genus †''
Greniodon ''Greniodon'' is a genus of extinct gondwanatherian mammal known from the Early to Middle Eocene ( Lutetian age, Mustersan to Divisaderan in the SALMA classification) of Argentina. A single species, ''Greniodon sylvaticus'', is known, describ ...
'' Goin et al., 2012 **** †'' Greniodon sylvanicus'' Goin et al., 2012 *** Genus †''
Vintana ''Vintana sertichi'' (from Malagasy vintana, "luck" ) is an early groundhog-like mammal dating from the Late Cretaceous, approximately 66 million years ago. Scientists found the lone fossil, a skull, on Madagascar's west coast in the Maastri ...
'' Krause et al., 2014 **** †''
Vintana sertichi ''Vintana sertichi'' (from Malagasy vintana, "luck" ) is an early groundhog-like mammal dating from the Late Cretaceous, approximately 66 million years ago. Scientists found the lone fossil, a skull, on Madagascar's west coast in the Maastri ...
'' Krause et al., 2014 *** Genus †'' Dakshina'' Wilson, Das Sarama & Anantharaman, 2007 **** †''
Dakshina jederi ''Bharattherium'' is a mammal that lived in India during the Maastrichtian (latest Cretaceous) and possibly the Paleocene. The genus has a single species, ''Bharattherium bonapartei''. It is part of the gondwanathere family Sudamericidae, wh ...
'' Wilson, Das Sarama & Anantharaman, 2007 *** Genus †''
Gondwanatherium ''Gondwanatherium'' is a genus of stem-mammal from the extinct suborder Gondwanatheria that lived in Patagonia, South America during the "Age of Dinosaurs", specifically during the Late Cretaceous (Campanian to Maastrichtian epochs). Descript ...
'' Bonaparte, 1986 **** †'' Gondwanatherium patagonicum'' Bonaparte, 1986 *** Genus †'' Sudamerica'' Scillato-Yané & Pascual, 1984 **** †''
Sudamerica ameghinoi ''Sudamerica'', literally " South America" in Spanish, is a genus of mammal from the extinct suborder Gondwanatheria that lived in Patagonia, Argentina (Salamanca Formation) and Antarctica (La Meseta Formation) from the Middle Paleocene (Peligra ...
'' Scillato-Yané & Pascual, 1984 *** Genus †'' Lavanify'' Krause et al., 1997 **** †''
Lavanify miolaka ''Lavanify'' is a mammalian genus from the late Cretaceous (probably Maastrichtian, about 71 to 66 million years ago) of Madagascar. The only species, ''L. miolaka'', is known from two isolated teeth, one of which is damaged. The teeth w ...
'' Krause et al., 1997 *** Genus †'' Bharattherium'' Prasad et al., 2007 **** †'' Bharattherium bonapartei'' Prasad et al.,, 2007 *** Genus †''
Patagonia Patagonia () refers to a geographical region that encompasses the southern end of South America, governed by Argentina and Chile. The region comprises the southern section of the Andes Mountains with lakes, fjords, temperate rainforests, and g ...
'' Pascual & Carlini' 1987 **** †''
Patagonia peregrina ''Patagonia'' is an extinct genus of non- placental mammal from the Miocene of Argentina. Traditionally considered a metatherian ''incertae sedis'', one analysis suggested it to be a gondwanathere. However, this has been rejected by other auth ...
'' Pascual & Carlini' 1987 * Suborder †
Cimolodonta Cimolodonta is a taxon of extinct mammals that lived from the Cretaceous to the Eocene. They were some of the more derived members of the extinct order Multituberculata. They probably lived something of a rodent-like existence until their ...
McKenna, 1975 ** Genus ?†''Allocodon'' non Marsh, 1881 *** Species †'' A. fortis'' Marsh, 1889 *** Species †'' A. lentus'' Marsh, 1892 'Cimolomys lentus''*** Species †'' A. pumilis'' Marsh, 1892 'Cimolomys pumilus''*** Species †'' A. rarus'' Marsh, 1889 ** Genus ?†''
Ameribaatar ''Ameribaatar'' is an extinct mammal of the Late Cretaceous. It was a member of the also extinct order of Multituberculata. It lived in North America during the Mesozoic, also known as the "age of the dinosaurs". Whether it belongs to Plagiaulac ...
'' Eaton & Cifelli, 2001 *** Species †'' A. zofiae'' Eaton & Cifelli, 2001 ** Genus ?†'' Bubodens'' Wilson, 1987 *** Species †'' Bubodens magnus'' Wilson, 1987 ** Genus ?†'' Clemensodon'' Krause, 1992 *** Species †''
Clemensodon megaloba ''Clemensodon'' is a genus of extinct mammal from the Upper Cretaceous of North America. It lived during the end of the Mesozoic, also known as the "age of the dinosaurs". It was a member of the extinct order of Multituberculata within the ...
'' Krause, 1992 'Kimbetohia cambi'', in partim** Genus ?†'' Fractinus'' Higgins 2003 *** Species †'' Fractinus palmorum'' Higgins, 2003 ** Genus ?†''
Uzbekbaatar ''Uzbekbaatar'' is a genus of extinct mammal from the Upper Cretaceous of Uzbekistan. It was a member of the extinct order Multituberculata within the suborder Cimolodonta, though its further affinities are unclear. The genus was named by Kiel ...
'' Kielan-Jaworowska & Nesov, 1992 *** Species †''
Uzbekbaatar kizylkumensis ''Uzbekbaatar'' is a genus of extinct mammal from the Upper Cretaceous of Uzbekistan. It was a member of the extinct order Multituberculata within the suborder Cimolodonta, though its further affinities are unclear. The genus was named by Kiel ...
'' Kielan-Jaworowska & Nesov, 1992 ** Genus ?†''
Viridomys ''Viridomys'' is a genus of extinct mammal from the Upper Cretaceous of North America. It was a member of the extinct order of Multituberculata, and lived during the Mesozoic, also known as the "age of the dinosaurs." It's within the suborder o ...
'' Fox 1971 *** Species †''
Viridomys orbatus ''Viridomys'' is a genus of extinct mammal from the Upper Cretaceous of North America. It was a member of the extinct order of Multituberculata, and lived during the Mesozoic, also known as the "age of the dinosaurs." It's within the subo ...
'' Fox 1971 ** Family †
Corriebaataridae ''Corriebaatar'' is the type and only genus of Corriebaataridae, a family of multituberculate mammals. It contains the single species ''Corriebaatar marywaltersae'' and represents the first evidence of Australian multituberculates Fossils fou ...
Rich et al., 2009 *** Genus ?†''
Corriebaatar ''Corriebaatar'' is the type and only genus of Corriebaataridae, a family of multituberculate mammals. It contains the single species ''Corriebaatar marywaltersae'' and represents the first evidence of Australian multituberculates Fossils found ...
'' Rich et al., 2009 **** Species †''
Corriebaatar marywaltersae ''Corriebaatar'' is the type and only genus of Corriebaataridae, a family of multituberculate mammals. It contains the single species ''Corriebaatar marywaltersae'' and represents the first evidence of Australian multituberculates Fossils fou ...
'' Rich et al., 2009 ** '' Paracimexomys'' group *** Genus '' Paracimexomys'' Archibald, 1982 **** Species? †'' P. crossi'' Cifelli, 1997 **** Species? †'' P. dacicus'' Grigorescu & Hahn, 1989 **** Species? †'' P. oardaensis'' (Codrea et al., 2014) 'Barbatodon oardaensis'' Codrea et al., 2014**** Species †'' P. magnus'' (Sahni, 1972) Archibald, 1982 'Cimexomys magnus'' Sahni, 1972**** Species †'' P. magister'' (Fox, 1971) Archibald, 1982 'Cimexomys magister'' Fox, 1971**** Species †'' P. perplexus'' Eaton & Cifelli, 2001 **** Species †'' P. robisoni'' Eaton & Nelson, 1991 **** Species †'' P. priscus'' (Lillegraven, 1969) Archibald, 1982 'Cimexomys priscus'' Lillegraven, 1969; genotype Paracimexomys sensu Eaton & Cifelli, 2001**** Species †'' P. propriscus'' Hunter, Heinrich & Weishampel 2010 *** Genus ''
Cimexomys ''Cimexomys'' is an extinct North American mammal that lived from the Upper Cretaceous to the Paleocene. For a while, it shared the world with dinosaurs, but outlived them. It was a member of the extinct order Multituberculata and lies withi ...
'' Sloan & Van Valen, 1965 **** Species †'' C. antiquus'' Fox, 1971 **** Species †'' C. gregoryi'' Eaton, 1993 **** Species †'' C. judithae'' Sahni, 1972 'Paracimexomys judithae'' (Sahni, 1972) Archibald, 1982**** Species †'' C. arapahoensis'' Middleton & Dewar, 2004 **** Species †'' C. minor'' Sloan & Van Valen, 1965 **** Species? †'' C. gratus'' (Jepson, 1930) Lofgren, 1995 'Cimexomys hausoi'' Archibald, 1983; ''Eucosmodon gratus'' Jepson, 1930; ''Mesodma ambigua''? Jepson, 1940; ''Stygimus gratus'' Jepson, 1930*** Genus †''
Bryceomys ''Bryceomys'' is an extinct mammal that lived during the late Cretaceous period (between 100 and 66 million years ago) and thus shared the world with dinosaurs. It was a member of the also extinct order of Multituberculata. It was within the subo ...
'' Eaton, 1995 **** Species †'' B. fumosus'' Eaton, 1995 **** Species †'' B. hadrosus'' Eaton, 1995 **** Species †'' B. intermedius'' Eaton & Cifelli, 2001 *** Genus †'' Cedaromys'' Eaton & Cifelli, 2001 **** Species †'' C. bestia'' (Eaton & Nelson, 1991) Eaton & Cifelli, 2001 'Paracimexomys bestia'' Eaton & Nelson, 1991/small> **** Species †'' C. hutchisoni'' Eaton 2002 **** Species †'' C. minimus'' Eaton 2009 **** Species †'' C. parvus'' Eaton & Cifelli, 2001 *** Genus †''
Dakotamys ''Dakotamys'' is a genus of extinct mammal that lived during the Upper Cretaceous. It shared the world with dinosaurs. This small creature was a member of the also extinct order Multituberculata within the suborder Cimolodonta and was a memb ...
'' Eaton, 1995 **** Species? †''D. sp.'' Eaton, 1995 **** Species †'' D. malcolmi'' Eaton, 1995 **** Species †'' D. shakespeari'' Eaton 2013 ** Family † Boffidae Hahn & Hahn, 1983 sensu Kielan-Jaworowska & Hurum 2001 *** Genus †''
Boffius ''Boffius'' is a genus of mammal from the Paleocene of Europe, which was named by Vianey-Liaud M. in 1979. It is a member of the extinct order of Multituberculata. Boffius lies within the suborder Cimolodonta and is the only known member of ...
'' Vianey-Liaud, 1979 **** Species †''
Boffius splendidus ''Boffius'' is a genus of mammal from the Paleocene of Europe, which was named by Vianey-Liaud M. in 1979. It is a member of the extinct order of Multituberculata. Boffius lies within the suborder Cimolodonta and is the only known member o ...
'' Vianey-Liaud, 1979 offiidae Hahn & Hahn, 1983 sensu Kielan-Jaworowska & Hurum, 2001** Family †
Cimolomyidae Cimolomyidae is a family of fossil mammal within the extinct order Multituberculata. Representatives are known from the Upper Cretaceous and the Paleocene of North America and perhaps Mongolia. The family is part of the suborder Cimolodonta. Oth ...
Marsh, 1889 sensu Kielan-Jaworowska & Hurum, 2001 *** Genus †'' Paressodon'' Wilson, Dechense & Anderson, 2010 **** Species †'' Paressodon nelsoni'' Wilson, Dechense & Anderson, 2010 *** Genus †'' Cimolomys'' Marsh, 1889 ''Allacodon''_Marsh,_1889;_''Selenacodon.html" ;"title="Allacodon.html" ;"title="''Allacodon">''Allacodon'' Marsh, 1889; ''Selenacodon">Allacodon.html" ;"title="''Allacodon">''Allacodon'' Marsh, 1889; ''Selenacodon'' Marsh, 1889] **** Species †''Cimolomys clarki, C. clarki'' Sahni, 1972 **** Species †''Cimolomys gracilis, C. gracilis'' Marsh, 1889 [''Cimolomys digona'' Marsh, 1889; ''Meniscoessus brevis''; ''Ptilodus gracilis'' Osborn, 1893 non Gidley 1909; ''Selenacodon brevis'' Marsh, 1889] **** Species †'' Cimolomys trochuus, C. trochuus'' Lillegraven, 1969 **** Species †'' C. milliensis'' Eaton, 1993a **** Species ?†'' C. bellus'' Marsh, 1889 *** Genus ?†'' Essonodon'' Simpson, 1927 **** Species †'' E. browni'' Simpson, 1927 imolodontidae? Kielan-Jaworowska & Hurum 2001*** Genus ?†'' Buginbaatar'' Kielan-Jaworowska & Sochava, 1969 **** Species †''
Buginbaatar transaltaiensis ''Buginbaatar'' is an extinct genus of mammal from the Upper Cretaceous of Mongolia. It is a member of the extinct order Multituberculata, within the suborder Cimolodonta and family Cimolomyidae. It lived towards the end of the Mesozoic era. ...
'' Kielan-Jaworowska & Sochava, 1969 *** Genus ?†''
Meniscoessus ''Meniscoessus'' is a genus of extinct mammal from the Upper Cretaceous Period of what is now North America. It was a member of the extinct order Multituberculata, lying within the suborder Cimolodonta and family Cimolomyidae. Taxonomy The ge ...
'' Cope, 1882 'Dipriodon''_Marsh,_1889;_''Tripriodon.html" ;"title="Dipriodon.html" ;"title="'Dipriodon">'Dipriodon'' Marsh, 1889; ''Tripriodon">Dipriodon.html" ;"title="'Dipriodon">'Dipriodon'' Marsh, 1889; ''Tripriodon'' Marsh, 1889 ''nomen dubium''; ''Triprotodon'' Chure & McIntosh, 1989 ''nomen dubium''; ''Selenacodon'' Marsh, 1889, ''Halodon'' Marsh, 1889, ''Oracodon'' Marsh, 1889] **** Species †'' M. caperatus'' Marsh, 1889 **** Species †'' M. collomensis'' Lillegraven, 1987 **** Species †'' M. conquistus'' Cope 1882 **** Species †'' M. ferox'' Fox, 1971a **** Species †'' M. intermedius'' Fox, 1976b **** Species †'' M. major'' (Russell, 1936) 'Cimolomys major'' Russell 1937**** Species †'' M. robustus'' (Marsh, 1889) [''Dipriodon robustus'' Marsh 1889; ''Dipriodon lacunatus'' Marsh, 1889; ''Tripriodon coelatus'' Marsh, 1889; ''Meniscoessus coelatus'' Marsh, 1889; ''Selenacodon fragilis'' Marsh, 1889; ''Meniscoessus fragilis'' Marsh, 1889; ''Halodon sculptus'' (Marsh, 1889); ''Cimolomys sculptus'' Marsh, 1889; ''Meniscoessus sculptus'' Marsh, 1889; ''Oracodon anceps'' Marsh, 1889; ''Oracodon conulus'' Marsh, 1892; ''Meniscoessus borealis'' Simpson, 1927c; ''Meniscoessus greeni'' Wilson, 1987] **** Species †''Meniscoessus seminoensis, M. seminoensis'' Eberle & Lillegraven, 1998a ** Family †
Kogaionidae Kogaionidae is a family of fossil mammals within the extinct order Multituberculata. Representatives are known from the Upper Cretaceous and the Paleocene of Europe. Having started as island endemics on Hateg Island during the Upper Cretaceous, ...
Rãdulescu & Samson, 1996 *** Genus †'' Kogaionon'' Rãdulescu & Samson, 1996 **** Species †'' K. ungureanui'' Rãdulescu & Samson, 1996 *** Genus †''
Hainina ''Hainina'' is an extinct mammal genus from the latest Cretaceous to the Paleocene of Europe. Genus The genus ''Hainina'' ("from Hainin") was named by Vianey-Liaud M. in 1979. This genus was originally referred to as Cimolomyidae. "We assign ...
'' Vianey-Liaud, 1979 **** Species †'' H. belgica'' Vianey-Liaud, 1979 **** Species †'' H. godfriauxi'' Vianey-Liaud, 1979 **** Species †'' H. pyrenaica'' Peláez-Campomanes, López-Martínez, Álvarez-Sierra & Daams, 2000 **** Species †'' H. vianeyae'' Peláez-Campomanes, López-Martínez, Álvarez-Sierra & Daams, 2000 *** Genus †''
Barbatodon ''Barbatodon'' is a mammal genus from the Upper Cretaceous period. It lived in Transylvania at the same time as some of the last dinosaurs and was a member of the extinct order of Multituberculata. It is within the suborder of Cimolodonta, ...
'' Rãdulescu & Samson, 1986 **** Species †'' B. transylvanicum'' Rãdulescu & Samson, 1986 ** Family † Eucosmodontidae Jepsen, 1940 sensu Kielan-Jaworowska & Hurum, 2001 ucosmodontidae: Eucosmodontinae Jepsen, 1940 sensu McKenna & Bell, 1997*** Genus †''
Eucosmodon ''Eucosmodon'' is a genus of extinct mammal from the Paleocene of North America. It is a member of the extinct order of Multituberculata within the suborder of Cimolodonta, and the family Eucosmodontidae. This genus has partly also been known ...
'' Matthew & Granger, 1921 **** Species †'' E. primus'' Granger & Simpson, 1929 **** Species †'' E. americanus'' Cope, 1885 **** Species †'' E. molestus'' Cope, 1869 'Neoplagiaulax molestus'' Cope, 1869*** Genus †''
Stygimys ''Stygimys'' is an extinct mammal genus from the Upper Cretaceous and Paleocene of North America. It was a member of the extinct order Multituberculata within the suborder Cimolodonta, family Eucosmodontidae. The genus ''Stygimys'' ("Styx mous ...
'' Sloan & Van Valen, 1965 **** Species †'' S. camptorhiza'' Johnston & Fox, 1984 **** Species †'' S. cupressus'' Fox, 1981 **** Species †'' S. kuszmauli'' 'Eucosmodon kuszmauli''**** Species †'' S. jepseni'' Simpson, 1935 **** Species †'' S. teilhardi'' Granger & Simpson, 1929 ** Family † Microcosmodontidae Holtzman & Wolberg, 1977 ucosmodontidae: Microcosmodontinae Holtzman & Wolberg, 1977 sensu McKenna & Bell, 1997*** Genus †''
Pentacosmodon ''Pentacosmodon'' is a mammal genus from the Paleocene of North America, so it lived somewhat after the "age of the dinosaurs". It was a member of the extinct order Multituberculata. It's within the suborder Cimolodonta and family Microcosmodonti ...
''Jepsen, 1940 **** Species †'' P. pronus'' Jepsen, 1940 jadochtatheroid? (Kielan-Jaworowska & Hurum, 2001)*** Genus †''
Acheronodon ''Acheronodon'' is a genus of herbivorous arboreal mammal which belongs to the family Microcosmodontidae and which was endemic to North America during the Early Paleocene subepoch (66—56.8 mya) and in existence for approximately . It is a mem ...
'' Archibald, 1982 **** Species †'' A. garbani'' Archibald, 1982 *** Genus †''
Microcosmodon ''Microcosmodon'' is a mammal genus from the Paleocene of North America. It was a member of the extinct order Multituberculata, and lies within the suborder Cimolodonta and family Microcosmodontidae. The genus ''Microcosmodon'' was named by G.L. ...
'' Jepsen, 1930 **** Species †'' M. conus'' Jepsen, 1930 **** Species †'' M. rosei'' Krause, 1980 **** Species †'' M. arcuatus'' Johnston & Fox, 1984 **** Species †'' M. woodi'' Holtzman & Wolberg, 1977 ucosmodontine?**** Species †'' M. harleyi'' Weil, 1998 ** Superfamily †
Ptilodontoidea Ptilodontoidea is a group of extinct mammals from the Northern Hemisphere. They were generally small, somewhat rodent-like creatures of the extinct order Multituberculata. Some of these genera boast a great many species, though remains are gen ...
Cope, 1887 sensu McKenna & Bell, 1997 e Kielan-Jaworowska & Hurum, 2001 *** Family †
Cimolodontidae Cimolodontidae is a family of fossil mammals within the extinct order Multituberculata. Representatives are known from the Upper Cretaceous and Paleocene of North America. The family Cimolodontidae was named by Othniel Charles Marsh in 18 ...
Marsh, 1889 sensu Kielan-Jaworowska & Hurum, 2001 **** Genus †''
Liotomus ''Liotomus'' is a genus of extinct mammal from the Paleocene epoch (early Cenozoic era). It lived in Europe and North America, and was a member of the extinct order Multituberculata, lying within the suborder Cimolodonta and possibly the f ...
'' Lemoine, 1882 'Neoctenacodon'' Lemoine 1891 ***** Species? †'' L. marshi'' (Lemoine, 1882) Cope, 1884 'Neoctenacodon marshi'' Lemoine, 1882; ''Neoplagiaulax marshi'' (Lemoine 1882); ''Plagiaulax marshi'' (Lemoine 1882) ucosmodontidae? McKenna & Bell, 1997/small> **** Genus †''
Yubaatar ''Yubaatar'' is a genus of multituberculate, an extinct order of rodent-like mammals, which lived in what is now China during the Late Cretaceous. The first specimen was discovered in the Qiupa Formation of Luanchuan County, in the Henan Provinc ...
'' Xu et al., 2015 ***** Species †''
Yubaatar zhongyuanensis ''Yubaatar'' is a genus of multituberculate, an extinct order of rodent-like mammals, which lived in what is now China during the Late Cretaceous. The first specimen was discovered in the Qiupa Formation of Luanchuan County, in the Henan Pro ...
'' Xu et al., 2015 **** Genus †''
Anconodon ''Anconodon'' is an extinct genus of mammal from the Paleocene of North America, and thus lived just after the " age of the dinosaurs". It was a member of the extinct order Multituberculata within the suborder Cimolodonta and possibly the fa ...
'' Jepsen, 1940 ***** Species? †'' A. lewisi'' (Simpson 1935) Sloan, 1987 ***** Species †'' A. gibleyi'' (Simpson, 1935) 'Ptilodus gidleyi'' Simpson, 1935***** Species †'' A. cochranensis'' (Russell, 1929) 'Liotomus russelli'' (Simpson, 1935); ''Anconodon russelli'' (Simpson, 1935) Sloan, 1987; ''Ectopodon cochranensis'' (Russell, 1967)**** Genus †''
Cimolodon ''Cimolodon'' is a genus of the extinct mammal order of Multituberculata within the suborder Cimolodonta and the family Cimolodontidae. Specimens are known from the Late Cretaceous of North America. Systematics The genus ''Cimolodon'' was nam ...
'' Marsh, 1889 'Nanomys''_Marsh,_1889,_''Nanomyops.html" ;"title="Nanomys.html" ;"title="'Nanomys">'Nanomys'' Marsh, 1889, ''Nanomyops">Nanomys.html" ;"title="'Nanomys">'Nanomys'' Marsh, 1889, ''Nanomyops'' Marsh, 1892] ***** Species †''Cimolodon agilis, C. agilis'' Marsh, 1889 ***** Species †''Cimolodon foxi, C. foxi'' Eaton, 2002 ***** Species †''Cimolodon gracilis, C. gracilis'' Marsh, 1889 ***** Species †'' Cimolodon electus, C. electus'' Fox, 1971 ***** Species †'' C. nitidus'' Marsh, 1889 [''Allacodon rarus'' Marsh, 1892 sensu Clemens, 1964a; ''Nanomys minutus'' Marsh, 1889; ''Nanomyops minutus'' (Marsh, 1889) Marsh, 1892; ''Halodon serratus'' Marsh, 1889; ''Ptilodus serratus'' (Marsh, 1889) Gidley 1909] ***** Species †''Cimolodon parvus, C. parvus'' Marsh, 1889 ***** Species †''Cimolodon peregrinus, C. peregrinus'' Donohue, Wilson & Breithaupt, 2013 ***** Species †'' C. similis'' Fox, 1971 ***** Species †'' C. wardi'' Eaton, 2006 *** Family ''
Incertae sedis ' () or ''problematica'' is a term used for a taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertain ...
'' **** Genus ''
Neoliotomus ''Neoliotomus'' is a genus of North American mammal from the Paleocene. It existed in the age immediately following the extinction of the last dinosaurs and was a member of the extinct order Multituberculata. It lies within the suborder Cimo ...
'' Jepsen, 1930 ***** Species †'' N. conventus'' Jepsen, 1930 ***** Species †'' N. ultimus'' (Granger & Simpson, 1928) *** Family †
Neoplagiaulacidae Neoplagiaulacidae is a family of mammal within the extinct order Multituberculata. Fossil remains are known from the Upper Cretaceous through to the latest Eocene/early Oligocene. Representatives have been found in North America, Europe and Asi ...
Ameghino, 1890 tilodontidae: Neoplagiaulacinae Ameghino, 1890 sensu McKenna & Bell, 1997**** Genus †''
Mesodma ''Mesodma'' is an extinct genus of mammal, a member of the extinct order Multituberculata within the suborder Cimolodonta, family Neoplagiaulacidae. It lived during the upper Cretaceous and Paleocene Periods of what is now North America. The e ...
'' Jepsen, 1940 ***** Species? †'' M. hensleighi'' Lillegraven, 1969 ***** Species? †'' M. senecta'' Fox, 1971 ***** Species †'' M. ambigua'' Jepsen, 1940 ***** Species? †'' M. pygmaea'' Sloan, 1987 ***** Species †'' M. formosa'' (Marsh, 1889) alodon formosus Marsh, 1889/small> ***** Species †'' M. primaeva'' (Lambe, 1902) ***** Species †'' M. thompsoni'' Clemens, 1964 **** Genus '' Ectypodus'' Matthew & Cranger, 1921 harlesmooria Kühne, 1969 /small> ***** Species †'' E. aphronorus'' Sloan, 1981 ***** Species? †'' E. childei'' Kühne, 1969 ***** Species? †'' E. elaphus'' Scott, 2005 ***** Species? †'' E. lovei'' (Sloan, 1966) Krishtlaka & Black, 1975 ***** Species †'' E. musculus'' Matthew & Granger, 1921 ***** Species †'' E. powelli'' Jepsen, 1940 ***** Species? †'' E. simpsoni'' Jepsen, 1930 ***** Species †'' E. szalayi'' Sloan, 1981 ***** Species †'' E. tardus'' Jepsen, 1930 **** Genus †''
Mimetodon ''Mimetodon'' is a small mammal from the Paleocene of North America and perhaps Europe. It was a member of the extinct order Multituberculata within the suborder Cimolodonta and family Neoplagiaulacidae. The genus ''Mimetodon'' was named by ...
'' Jepsen, 1940 ***** Species †'' M. krausei'' Sloan, 1981 ***** Species †'' M. nanophus'' Holtzman, 1978 'Neoplagiaulax nanophus'' Holtzman, 1978/small> ***** Species †'' M. siberlingi''(Simpson, 1935) Schiebout, 1974 ***** Species †'' M. churchilli'' Jepsen, 1940 **** Genus †''
Neoplagiaulax ''Neoplagiaulax'' is a mammal genus from the Paleocene of Europe and North America. In the case of the latter continent, there may possibly be some slightly earlier, Upper Cretaceous material too. It existed in the age immediately following the ...
'' Lemoine, 1882 ***** Species †'' N. annae'' Vianey-Liaud, 1986 ***** Species? †'' N. burgessi'' Archibald, 1982 ***** Species †'' N. cimolodontoides'' Scott, 2005 ***** Species †'' N. copei'' Lemoine, 1885 ***** Species †'' N. donaldorum'' Scott & Krause, 2006 ***** Species †'' N. eocaenus'' Lemoine, 1880 ***** Species †'' N. grangeri'' Simpson, 1935 ***** Species †'' N. hazeni'' Jepsen, 1940 ***** Species †'' N. hunteri'' Krishtalka, 1973 ***** Species †'' N. jepi'' Sloan, 1987 ***** Species †'' N. kremnus'' Johnston & Fox, 1984 ***** Species †'' N. macintyrei'' Slaon, 1981 ***** Species †'' N. macrotomeus'' Wilson, 1956 ***** Species †'' N. mckennai'' Sloan, 1987 ***** Species †'' N. nelsoni'' Sloan, 1987 ***** Species †'' N. nicolai'' Vianey-Liaud, 1986 ***** Species †'' N. paskapooensis'' Scott, 2005 ***** Species? †'' N. serrator'' Scott, 2005 ***** Species †'' N. sylvani'' Vianey-Liaud, 1986 **** Genus †''
Parectypodus ''Parectypodus'' (meaning "besides '' Ectypodus''") is an extinct genus of mammals that lived from Late Cretaceous (Maastrichtian) to Eocene time in North America. It is a member of the extinct order of Multituberculata, suborder Cimolodonta, fam ...
'' Jepsen, 1930 ***** Species †'' P. armstrongi'' Johnston & Fox, 1984 ***** Species? †'' P. corystes'' Scott, 2003 ***** Species? †'' P. foxi'' Storer, 1991 ***** Species †'' P. laytoni'' Jepsen, 1940 ***** Species †'' P. lunatus'' Krause, 1982 'P. childei'' Kühne, 1969***** Species †'' P. simpsoni'' Jepsen, 1940 ***** Species †'' P. sinclairi'' Simpson, 1935 ***** Species †'' P. sloani'' Schiebout, 1974 ***** Species †'' P. trovessartianus'' Cope, 1882 'P. trouessarti''; ''Ptilodus''; ''Mimetodon''; ''Neoplagiaulax''***** Species †'' P. sylviae'' Rigsby, 1980 ctypodus sylviae Rigby, 1980***** Species? †'' P. vanvaleni'' Sloan, 1981 **** Genus †''
Cernaysia ''Cernaysia'' is an extinct genus of mammal from the Paleocene of France and the United States. It existed in the age immediately following the extinction of the last dinosaurs. This animal was a member of the extinct order Multituberculata w ...
'' Vianey-Liaud, 1986 ***** Species †'' C. manueli'' Vianey-Liaud, 1986 ***** Species †'' C. davidi'' Vianey-Liaud, 1986 **** Genus †''
Krauseia ''Krauseia'' is an extinct genus of small mammal from the Paleocene of North America. It was a member of the extinct order Multituberculata and is within the suborder of Cimolodonta, family Neoplagiaulacidae. The genus was nemed by Vianey-Li ...
'' Vianey-Liaud, 1986 ***** Species †'' K. clemensi'' Sloan, 1981 'Parectypodus clemensi'' Sloan, 1981**** Genus †''
Xyronomys ''Xyronomys'' is an extinct genus of small mammals from the Paleocene of North America, with two described species. The genus lies within the extinct order Multituberculata within the suborder Cimolodonta and family Neoplagiaulacidae. The spe ...
''Rigby, 1980 ***** Species †'' X. swainae'' Rigby, 1980 ironomys (sic); ?Eucosmodontidae**** Genus †''
Xanclomys ''Xanclomys'' is a small mammal from the Paleocene of North America. It was a genus within the extinct order Multituberculata within the suborder Cimolodonta and family Neoplagiaulacidae. The genus ''Xanclomys'', named by Rigby J.K. in 1980 ...
'' Rigby, 1980 ***** Species †'' X. mcgrewi''Rigby, 1980 **** Genus †''
Mesodmops ''Mesodmops'' is a genus of small mammal from the Eocene of China. It was a late member of the extinct order of Multituberculata. It's within the suborder of Cimolodonta and family Neoplagiaulacidae Neoplagiaulacidae is a family of mammal ...
'' Tong & Wang, 1994 ***** Species †'' M. dawsonae'' Tong & Wang, 1994 *** Family †
Ptilodontidae Ptilodontidae is a family of primitive mammals within the extinct order Multituberculata. Representatives are known from the Upper Cretaceous and Paleocene of North America. The Ptilodontidae family was originally named Ptilodontinae and class ...
Cope, 1887 tilodontidae: Ptilodontinae Cope, 1887 sensu McKenna & Bell, 1997**** Genus †'' Kimbetohia'' Simpson, 1936 ***** Species †'' K. cambi'' small>Granger, Gregory & Colbert in Matthew, 1937, or Simpson, 1936
***** Species †K. sp. cf. ''K. cambi'' **** Genus †''
Ptilodus ''Ptilodus'' (meaning "soft-haired") is a genus of mammals from the extinct order of Multituberculata, and lived during the Paleocene in North America. ''Ptilodus'' was a relatively large multituberculate of in length, which is about the same s ...
'' Cope, 1881 hirox Cope, 1884***** Species? †'' P. fractus'' ***** Species †'' P. kummae'' Krause, 1977 ***** Species †'' P. gnomus'' Scott, Fox & Youzwyshyn, 2002 f. ''Ectypodus hazeni'' (Jepsen, 1940) Gazin, 1956***** Species †'' P. mediaevus'' Cope, 1881 'Ptilodus plicatus'' (Cope, 1884); ''Chirox plicatus'' Cope, 1884 ''P. ferronensis'' Gazin, 1941***** Species †'' P. montanus'' Douglass, 1908 'P. gracilis'' Gidley, 1909; ''P. admiralis'' Hay, 1930***** Species †'' P. tsosiensis'' Sloan, 1981 ***** Species †'' P. wyomingensis'' Jepsen, 1940 **** Genus †''
Baiotomeus ''Baiotomeus'' is a genus of mammals from the extinct order of Multituberculata. It is known from the Paleocene of North America. The genus ''Baiotomeus'' was formally named by Krause in 1987 ( Krause, 1987), and has also been known as '' Mi ...
'' Krause, 1987 ***** Species †'' B. douglassi'' Simpson, 1935 'Ptilodus''; ''Mimetodon''; ''Neoplagiaulax''***** Species †'' B. lamberti'' Krause, 1987 ***** Species †'' B. russelli'' Scott, Fox & Youzwyshyn, 2002 ***** Species †'' B. rhothonion'' Scott, 2003 **** Genus †''
Prochetodon ''Prochetodon'' is a genus of mammals from the extinct order Multituberculata. It lived during the Upper Paleocene and the Lower Eocene in North America. The genus was formally named by G. L. Jepsen in 1940. ( Jepsen, 1940) ''P. cavus'' The s ...
'' Jepsen, 1940 ***** Species †'' P. cavus'' Jespen, 1940 ***** Species †'' P. foxi'' Krause, 1987 ***** Species †'' P. taxus'' Krause, 1987 ***** Species? †'' P. speirsae'' Scott, 2004 ** Superfamily †
Taeniolabidoidea Taeniolabidoidea is a group of extinct mammals known from North America and Asia. They were the largest members of the extinct order Multituberculata, as well as the largest non-therian mammals. ''Lambdopsalis'' even provides direct fossil eviden ...
Granger & Simpson, 1929 sensu Kielan-Jaworowska & Hurum, 2001 *** Genus †''
Prionessus ''Prionessus'' is a genus of extinct mammal from the Paleocene of Central Asia. It was a member of the extinct order Multituberculata within the suborder Cimolodonta and superfamily Taeniolabidoidea. The genus was named by Matthew W.D. and Gran ...
'' Matthew & Granger, 1925 **** Species †'' P. lucifer'' Matthew & Granger, 1925 *** Family †
Lambdopsalidae Lambdopsalidae is a family of extinct multituberculate mammals from the Late Paleocene of Asia. They are part of Taeniolabidoidea, a clade otherwise present in the Early Paleocene (and possibly the Late Cretaceous) of North America; lambdopsalids ...
**** Genus †''
Lambdopsalis ''Lambdopsalis bulla'' is an extinct multituberculate mammal from the Late Paleocene of China and Mongolia. It is placed within the suborder Cimolodonta and is a member of the superfamily Taeniolabidoidea. Fossil remains have been found in the ...
'' Chow & Qi, 1978 ***** Species †'' L. bulla'' Chow & Qi, 1978 **** Genus †''
Sphenopsalis ''Sphenopsalis'' is a genus of extinct mammal from the Paleocene of Central Asia. It was a member of the extinct order Multituberculata, and lies within the suborder Cimolodonta and the superfamily Taeniolabidoidea. The genus was named by Willia ...
'' Matthew, Granger & Simpson, 1928 ***** Species †'' S. nobilis'' Matthew, Granger & Simpson, 1928 *** Family †
Taeniolabididae Taeniolabididae is one of the two multituberculate clades within Taeniolabidoidea. Originally basically synonymous with Taeniolabidoidea, it has more recently been found to be a specific clade including '' Kimbetopsalis'', ''Taeniolabis'' and som ...
Granger & Simpson, 1929 **** Genus †''
Taeniolabis ''Taeniolabis'' ("banded incisor") is a genus of extinct multituberculate mammal from the Paleocene of North America. Description It is the largest known member of the extinct order Multituberculata, as well as the largest non-therian mamm ...
'' Cope, 1882 ***** Species †'' T. lamberti'' Simmons, 1987 ***** Species †'' T. taoensis'' Cope, 1882 **** Genus †'' Kimbetopsalis'' ***** Species †'' K. simmonsae'' ** Superfamily †
Djadochtatherioidea Djadochtatherioidea is a group of extinct mammals known from the upper Cretaceous of Central Asia. They were members of an also extinct order called Multituberculata. These were generally somewhat rodent-like creatures, who scurried around dur ...
Kielan-Jaworowska & Hurum, 1997 sensu Kielan-Jaworowska & Hurum, 2001 jadochtatheria Kielan-Jaworowska & Hurum, 1997*** Genus? †''
Bulganbaatar ''Bulganbaatar'' is a Central Asian mammal genus from the Upper Cretaceous. It existed in the company of dinosaurs. This animal was a member of the extinct order Multituberculata. It's within the suborder Cimolodonta and is a member of the ...
'' Kielan-Jaworowska, 1974 **** Species? †'' B. nemegtbaataroides'' Kielan-Jaworowska, 1974 *** Genus †''
Nemegtbaatar ''Nemegtbaatar'' is a genus of mammal from the Upper Cretaceous Period of what is now Mongolia. It existed in the company of much larger dinosaurs, found together in the Nemegt Basin. This creature was a member of the extinct order Multitubercu ...
'' Kielan-Jaworowska, 1974 **** Species? †'' N. gobiensis'' Kielan-Jaworowska, 1974 *** Family †
Chulsanbaataridae ''Chulsanbaatar'' is a genus of mammal from the Cretaceous of Mongolia. It lived during the Mesozoic, also known as the "age of the dinosaurs." This small creature was a member of the extinct order of Multituberculata and is within the su ...
Kielan-Jaworowska, 1974 **** Genus †''
Chulsanbaatar ''Chulsanbaatar'' is a genus of mammal from the Cretaceous of Mongolia. It lived during the Mesozoic, also known as the "age of the dinosaurs." This small creature was a member of the extinct order of Multituberculata and is within the suborder ...
'' Kielan-Jaworowska, 1974 ***** Species †'' C. vulgaris'' Kielan-Jaworowska, 1974 *** Family †
Sloanbaataridae Sloanbaataridae is a family of fossil mammals within the extinct order Multituberculata. Remains are known from the Upper Cretaceous of Mongolia. These small herbivores lived during the "age of the dinosaurs". This family is part of the suborder ...
Kielan-Jaworowska, 1974 **** Genus †''
Kamptobaatar ''Kamptobaatar'' is a Mongolian mammal genus from the Upper Cretaceous. It lived at the same time as the later dinosaurs. This animal was a member of the extinct order Multituberculata within the suborder Cimolodonta and family Sloanbaataridae. ...
'' Kielan-Jaworowska, 1970 ***** Species? †'' K. kuczynskii'' Kielan-Jaworowska, 1970 **** Genus †''
Nessovbaatar ''Nessovbaatar'' is a genus of extinct mammal from the Upper Cretaceous of Mongolia. It eked out its living in the company of Central Asian dinosaurs. This animal was a member of the extinct order Multituberculata within the suborder Cimolodo ...
'' Kielan-Jaworowska & Hurum, 1997 ***** Species †'' N. multicostatus'' Kielan-Jaworowska & Hurum, 1997 **** Genus †''
Sloanbaatar ''Sloanbaatar'' is a mammal genus that lived in Mongolia during the Upper Cretaceous. It lived at the same time as the dinosaurs. This animal was a member of the also extinct order Multituberculata within the suborder Cimolodonta and the family ...
'' Kielan-Jaworowska, 1974 ***** Species †'' S. mirabilis'' Kielan-Jaworowska, 1974 loanbaatarinae*** Family †
Djadochtatheriidae Djadochtatheriidae is a family of fossil mammals within the extinct order Multituberculata. Remains are known from the Upper Cretaceous of Central Asia. These animals lived during the Mesozoic, also known as the "age of the dinosaurs". This fam ...
Kielan-Jaworowska $ Hurum, 1997 **** Genus †''
Djadochtatherium ''Djadochtatherium'' is a mammal genus that lived in Mongolia during the Upper Cretaceous. It coexisted with some of the late dinosaurs. This animal was a member of the extinct order of Multituberculata. It is within the suborder of Cimolodonta ...
'' Simpson, 1925 ***** Species †'' D. matthewi'' Simpson, 1925 'Catopsalis matthewi'' Simpson, 1925**** Genus †'' Catopsbaatar'' Kielan-Jaworowska, 1974 ***** Species †'' C. catopsaloides'' (Kielan-Jaworowska, 1974) Kielan-Jaworowska, 1994 'Djadochtatherium catopsaloides'' Kielan-Jaworowska, 1974**** Genus †''
Tombaatar ''Tombaatar'' is a mammal genus that existed during the Mongolian Upper Cretaceous period. It co-existed with some of the late dinosaurs. This animal was a member of the extinct order Multituberculata, within the suborder Cimolodonta and f ...
'' Kielan-Jaworowska, 1974 ***** Species †'' T. sabuli'' Rougier, Novacek & Dashzeveg, 1997 **** Genus †''
Kryptobaatar ''Kryptobaatar''and also known as ''Gobibaatar'' or ''Tugrigbaatar'' is an extinct mammalian genus dating from the Upper Cretaceous Period and identified in Central Asia. This animal was a member of the extinct order of Multituberculata within ...
'' Kielan-Jaworowska, 1970 'Gobibaatar'' Kielan-Jaworowska, 1970, ''Tugrigbaatar'' Kielan-Jaworowska & Dashzeveg, 1978***** Species †'' K. saichanensis'' Kielan-Jaworowska & Dashzeveg, 1978 'Tugrigbaatar saichaenensis'' Kielan-Jaworowska & Dashzeveg, 1978??***** Species †'' K. dashzevegi'' Kielan-Jaworowska, 1970 ***** Species †'' K. mandahuensis'' Smith, Guo & Sun, 2001 ***** Species †'' K. gobiensis'' Kielan-Jaworowska, 1970 'Gobibaatar parvus'' Kielan-Jaworowska, 1970


Phylogeny


Paleoecology


Behaviour

Multituberculates are some of the earliest mammals to display complex social behaviours One species, '' Filikomys,'' from the Late Cretaceous of North America, engaged in multi-generational group nesting and burrowing.


Extinction

The extinction of multituberculates has been a topic of controversy for several decades. After at least 88 million years of dominance over most mammalian assemblies, multituberculates reached the peak of their diversity in the early
Palaeocene The Paleocene, ( ) or Palaeocene, is a geological epoch that lasted from about 66 to 56 million years ago (mya). It is the first epoch of the Paleogene Period in the modern Cenozoic Era. The name is a combination of the Ancient Greek ''pala ...
, before gradually declining across the final stages of the epoch and the
Eocene The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', ...
, finally disappearing in the early
Oligocene The Oligocene ( ) is a geologic epoch of the Paleogene Period and extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that define the epoch are well identified but t ...
. Traditionally, the extinction of multituberculates has been linked to the rise of
rodents Rodents (from Latin , 'to gnaw') are mammals of the order Rodentia (), which are characterized by a single pair of continuously growing incisors in each of the upper and lower jaws. About 40% of all mammal species are rodents. They are na ...
(and, to a lesser degree, earlier
placental Placental mammals (infraclass Placentalia ) 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 distinguishe ...
competitors like hyopsodonts and
Plesiadapiformes Plesiadapiformes ("Adapid-like" or "near Adapiformes") is a group of Primates, a sister of the Dermoptera. While none of the groups normally directly assigned to this group survived, the group appears actually not to be literally extinct (in t ...
), which supposedly competitively excluded multituberculates from most mammalian faunas. However, the idea that multituberculates were replaced by rodents and other placentals has been criticised by several authors. For one thing, it relies on the assumption that these mammals are "inferior" to more derived placentals, and ignores the fact that rodents and multituberculates had co-existed for at least 15 million years. According to some researchers, multituberculate "decline" is shaped by sharp extinction events, most notably after the
Tiffanian The Tiffanian North American Stage on the geologic timescale is the North American faunal stage according to the North American Land Mammal Ages chronology (NALMA), typically set from 60,200,000 to 56,800,000 years BP lasting . It is usually c ...
, where a sudden drop in diversity occurs. Finally, the youngest known multituberculates do not exemplify patterns of competitive exclusion; the Oligocene '' Ectypodus'' is a rather generalistic species, rather than a specialist. This combination of factors suggests that, rather than gradually declining due to pressure from rodents and similar placentals, multituberculates simply could not cope with climatic and vegetation changes, as well as the rise of new predatory eutherians, such as
miacids Miacids are extinct primitive carnivoramorphans within the family Miacidae that lived during the Paleocene and Eocene epochs, about 62–34 million years ago. Miacids existed for approximately . Miacids are thought to have evolved into the ...
. More recent studies show a mixed effect. Multituberculate faunas in North America and Europe do indeed decline in correlation to the introduction of rodents in these areas. However,
Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an are ...
n multituberculate faunas co-existed with rodents with minimal extinction events, implying that competition was not the main cause for the extinction of Asiatic multituberculates. As a whole, it seems that Asian multituberculates, unlike North American and European species, never recovered from the KT event, which allowed the evolution and propagation of rodents in the first place. A recent study seems to indeed indicate that eutherians recovered more quickly from the KT event than multituberculates. Conversely, another study has shown that placental radiation did not start significantly until after the decline of multituberculates.


References


Sources

* * * {{Taxonbar, from=Q131181 Prehistoric animal orders Prehistoric mammals Mammal orders Oligocene extinctions Kimmeridgian first appearances Taxa named by Edward Drinker Cope Fossil taxa described in 1884 Tertiary extinctions of vertebrate taxa