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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 projectio ...
s of their teeth) is an extinct order 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), fu ...
s with a fossil record spanning over 130 million years. They first appeared in the Middle Jurassic, and reached a peak diversity during the Late
Cretaceous 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 ''palai ...
. 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 ''palai ...
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'', "da ...
. 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 Cretaceo ...
mammals with more than 200 species known, ranging from mouse-sized to beaver-sized. These species occupied a diversity of ecological niches, ranging from burrow-dwelling to squirrel-like arborealism to jerboa-like hoppers. Multituberculates are usually placed as
crown 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 a p ...
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 Euharamiyida also known as Eleutherodontida, is clade of early mammals or mammal-like cynodonts from the Middle Jurassic to Early Cretaceous of Eurasia and possibly North America. The group is sometimes considered a sister group to Multitubercula ...
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 '' Allostaffi ...
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 projectio ...
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 A plagiaulacoid is a type of blade-like, most often serrated, tooth present in various mammal groups, usually a premolar. Among modern species it is present chiefly on diprotodontian marsupials (specifically, Potoroidae, '' Bettongia'' and '' Burram ...
; other mammals, like Plesiadapiformes and diprotodontian
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 p ...
, 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 only the fourth lower premolar remained, with the third one remaining only as a vestigial peg-like tooth, and in several taxa like taeniolabidoideans, 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 ...
), 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 remains ...
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 o ...
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 a p ...
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, in which tooth enamel extends beyond the gumline: lambdopsalid taeniolabidoideans and sudamericid
gondwanatheres 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 '' Allostaffia ...
. 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 period, and then survived and even dominated for over one hundred million years, longer than any other order of mammaliforms, including placental mammals. The earliest known multituberculates are from the
Middle Jurassic The Middle Jurassic is the second epoch of the Jurassic 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 rare, but geological formations c ...
( Bathonian ~166-168 million years ago) of England and Russia, including '' Hahnotherium'' ''and Kermackodon'' 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" were abundant and widespread across Laurasia (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 ago) ...
stage of the Early Cretaceous, the advanced subgroup Cimolodonta 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 int ...
, 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 ''palai ...
, 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 th ...
, 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 ...
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, c ...
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'', "da ...
. There are some isolated records of multituberculates from the Southern Hemisphere, including the cimolodontan '' Corriebaatar'' from the Early Cretaceous of Australia, and fragmentary remains from the Late Cretaceous
Maevarano Formation The Maevarano Formation is a Late Cretaceous sedimentary rock formation found in the Mahajanga Province of northwestern Madagascar. It is most likely Maastrichtian in age, and records a seasonal, semiarid environment with rivers that had greatly ...
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 ...
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 mou ...
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 consona ...
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 feeding ...
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 Bi ...
, Wyoming, 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 Euharamiyida also known as Eleutherodontida, is clade of early mammals or mammal-like cynodonts from the Middle Jurassic to Early Cretaceous of Eurasia and possibly North America. The group is sometimes considered a sister group to Multitubercula ...
, 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 '' Allostaffi ...
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'', t ...
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 th ...
found that most multituberculates could be referred to two suborders: " Plagiaulacida" and Cimolodonta. The exception is the genus '' Arginbaatar'', 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 A grade is a taxon united by a level of morphological or physiological complexity. The term was coined by British biologist Julian Huxley, to contrast with clade, a strictly phylogenetic unit. Definition An evolutionary grade is a group of sp ...
. 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 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 grou ...
) 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'', "da ...
. The superfamilies Djadochtatherioidea, Taeniolabidoidea, Ptilodontoidea 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 assigne ...
. 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. Othe ...
, Boffiidae, Eucosmodontidae, Kogaionidae, Microcosmodontidae and the two genera '' Uzbekbaatar'' and '' Viridomys''. 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 Simpson 1925 * Genus ?†'' Argillomys'' Cifelli, Gordon & Lipka 2013 ** Species †'' Argillomys marylandensis'' Cifelli, Gordon & Lipka 2013 * Genus ?†'' Janumys'' Eaton & Cifelli 2001 ** Species †'' Janumys erebos'' Eaton & Cifelli 2001 * Super family †Allodontoidea Marsh 1889 ** Genus †?'' Glirodon'' Engelmann & Callison, 2001 *** Species †'' G. grandis'' Engelmann & Callison, 2001 ** Family † Arginbaataridae Hahn & Hahn, 1983 *** Genus †'' Arginbaatar'' Trofimov, 1980 **** Species †'' A. dmitrievae'' Trofimov, 1980 ** Family † Zofiabaataridae Bakker, 1992 *** Genus †'' Zofiabaatar'' 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'' Hahn, LePage & Wouters 1987 ** Genus ?†'' Rugosodon'' Yuan et al., 2013 *** Species †'' Rugosodon eurasiaticus'' Yuan et al., 2013 ** Family † Pinheirodontidae Hahn & Hahn, 1999 *** Genus †'' Bernardodon'' 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'' Hahn & Hahn, 1999 **** Species †'' E. anomala'' Hahn & Hahn, 1999 *** Genus †'' Gerhardodon'' Kielan-Jaworowska & Ensom, 1992 **** Species †'' G. purbeckensis'' Kielan-Jaworowska & Ensom, 1992 *** Genus †'' Iberodon'' Hahn & Hahn, 1999 **** Species †'' I. quadrituberculatus'' Hahn & Hahn, 1999 *** Genus †'' Lavocatia'' Canudo & Cuenca-Bescós, 1996 **** Species †'' L. alfambrensis'' Canudo & Cuenca-Bescós, 1996 *** Genus †'' Pinheirodon'' Hahn & Hahn, 1999 **** Species †'' P. pygmaeus'' Hahn & Hahn, 1999 **** Species †'' P. vastus'' Hahn & Hahn, 1999 ** Family † Paulchoffatiidae Hahn, 1969 *** Genus ?†'' Galveodon'' Hahn & Hahn, 1992 **** Species †'' G. nannothus'' Hahn & Hahn, 1992 *** Genus ?†'' Sunnyodon'' Kielan-Jaworowska & Ensom, 1992 **** Species †'' S. notleyi'' Kielan-Jaworowska & Ensom, 1992 *** subfamily † Paulchoffatiinae Hahn, 1971 **** Genus †'' Paulchoffatia'' Kühne, 1961 ***** Species †'' P. delgador'' Kühne, 1961 **** Genus †'' Pseudobolodon'' Hahn, 1977 ***** Species †'' P. oreas'' Hahn, 1977 ***** Species †'' P. krebsi'' Hahn & Hahn, 1994 **** Genus †'' Henkelodon'' Hahn, 1987 ***** Species †'' H. naias'' Hahn, 1987 **** Genus †'' Guimarotodon'' Hahn, 1969 ***** Species †'' G. leiriensis'' Hahn, 1969 **** Genus †'' Meketibolodon'' (Hahn, 1978) Hahn, 1993 ***** Species †'' M. robustus'' (Hahn, 1978) Hahn, 1993 'Pseudobolodon robusutus'' Hahn 1978**** Genus †'' Plesiochoffatia'' 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'' Hahn & Hahn, 1998 ***** Species †'' X. oinopion'' Hahn & Hahn, 1998 **** Genus †'' Bathmochoffatia'' Hahn & Hahn, 1998 ***** Species †'' B. hapax'' Hahn & Hahn, 1998 **** Genus †'' Kielanodon'' Hahn, 1987 ***** Species †'' K. hopsoni'' Hahn, 1987 **** Genus †'' Meketichoffatia'' Hahn, 1993 ***** Species †'' M. krausei'' Hahn, 1993 **** Genus †'' Renatodon'' Hahn, 2001 ***** Species †'' Renatodon amalthea'' Hahn, 2001 *** Subfamily † Kuehneodontinae Hahn, 1971 **** Genus †'' Kuehneodon'' 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 Gill, 1872 sensu Kielan-Jaworowska & Hurum, 2001 olodontidae Osborn 1887 *** Genus ?†'' Morrisonodon'' Hahn & Hahn, 2004 **** Species †'' Morrisonodon brentbaatar'' (Bakker, 1998) Hahn & Hahn, 2004 'Ctenacodon brentbaatar'' Bakker, 1998*** Genus †'' Plagiaulax'' Falconer, 1857 **** Species †'' P. becklesii'' Falconer, 1857 **** Species †'' P. dawsoni'' Woodward, 1891 'Plioprion dawsoni'' Woodward, 1891; ''Loxaulax dawsoni'' (Woodward, 1891) Sloan, 1979*** Genus †'' Bolodon'' 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 Kielan-Jaworowska, Dashzeveg & Trofimov, 1987 ** Genus †'' Eobaatar'' 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'' Kusuhashi et al., 2008 *** Species †'' H. matsuoi'' Kusuhashi et al., 2008 ** Genus †'' Heishanobaatar'' Kusuhashi et al., 2010 *** Species †'' H. triangulus'' Kusuhashi et al., 2010 ** Genus †'' Iberica'' Badiola et al., 2011 *** Species †'' Iberica hahni'' Badiola et al., 2011 ** Genus †'' Liaobaatar'' 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'' 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'' Martin et al., 2016 *** Species †'' Teutonodon langenbergensis'' 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 '' Allostaffi ...
McKenna 1971 ondwanatheroidea Krause & Bonaparte 1993** Family †
Groeberiidae Groeberiidae is a family of strange non-placental mammals from the Eocene and Oligocene epochs of Patagonia, Argentina and Chile, South America. Originally classified as paucituberculate marsupials, they were suggested to be late representative ...
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 ''Ferugliotherium'' is a genus of fossil mammals in the family Ferugliotheriidae from the Campanian and/or Maastrichtian period (Late Cretaceous; around 70 million years ago) of Argentina. It contains a single species, ''Ferugliotherium windhaus ...
'' 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'' 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 Maastrichtia ...
'' Krause et al., 2014 **** †'' Vintana sertichi'' Krause et al., 2014 *** Genus †'' Dakshina'' Wilson, Das Sarama & Anantharaman, 2007 **** †'' Dakshina jederi'' Wilson, Das Sarama & Anantharaman, 2007 *** Genus †'' Gondwanatherium'' Bonaparte, 1986 **** †'' Gondwanatherium patagonicum'' Bonaparte, 1986 *** Genus †''
Sudamerica ''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 ( Pel ...
'' Scillato-Yané & Pascual, 1984 **** †'' Sudamerica ameghinoi'' Scillato-Yané & Pascual, 1984 *** Genus †''
Lavanify ''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 were ...
'' Krause et al., 1997 **** †'' Lavanify miolaka'' Krause et al., 1997 *** Genus †''
Bharattherium ''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, which i ...
'' 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 ...
'' Pascual & Carlini' 1987 **** †'' Patagonia peregrina'' Pascual & Carlini' 1987 * Suborder † Cimolodonta 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'' 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'' Krause, 1992 'Kimbetohia cambi'', in partim** Genus ?†'' Fractinus'' Higgins 2003 *** Species †'' Fractinus palmorum'' Higgins, 2003 ** Genus ?†'' Uzbekbaatar'' Kielan-Jaworowska & Nesov, 1992 *** Species †'' Uzbekbaatar kizylkumensis'' Kielan-Jaworowska & Nesov, 1992 ** Genus ?†'' Viridomys'' Fox 1971 *** Species †'' Viridomys orbatus'' Fox 1971 ** Family † Corriebaataridae Rich et al., 2009 *** Genus ?†'' Corriebaatar'' Rich et al., 2009 **** Species †'' Corriebaatar marywaltersae'' 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 within ...
'' 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'' 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'' 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'' 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. Othe ...
Marsh, 1889 sensu Kielan-Jaworowska & Hurum, 2001 *** Genus †'' Paressodon'' Wilson, Dechense & Anderson, 2010 **** Species †'' Paressodon nelsoni'' Wilson, Dechense & Anderson, 2010 *** Genus †''
Cimolomys ''Cimolomys'' is a mammal genus from the Upper Cretaceous of North America. It was a member of the extinct order Multituberculata within the suborder Cimolodonta and family Cimolomyidae. The genus ''Cimolomys'' was named by Othniel Charles Marsh ...
'' 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 ''Essonodon'' is a mammal genus from the Upper Cretaceous of North America. It was a member of the extinct order Multituberculata and lived towards the end of the "age of the dinosaurs." It is within the suborder Cimolodonta and perhaps the fam ...
'' Simpson, 1927 **** Species †'' E. browni'' Simpson, 1927 imolodontidae? Kielan-Jaworowska & Hurum 2001*** Genus ?†''
Buginbaatar ''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. The ...
'' Kielan-Jaworowska & Sochava, 1969 **** Species †'' Buginbaatar transaltaiensis'' Kielan-Jaworowska & Sochava, 1969 *** Genus ?†'' Meniscoessus'' 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 Rãdulescu & Samson, 1996 *** Genus †'' Kogaionon'' Rãdulescu & Samson, 1996 **** Species †'' K. ungureanui'' Rãdulescu & Samson, 1996 *** Genus †'' Hainina'' 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'' 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'' 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'' 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''Jepsen, 1940 **** Species †'' P. pronus'' Jepsen, 1940 jadochtatheroid? (Kielan-Jaworowska & Hurum, 2001)*** Genus †'' Acheronodon'' Archibald, 1982 **** Species †'' A. garbani'' Archibald, 1982 *** Genus †'' Microcosmodon'' 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 Cope, 1887 sensu McKenna & Bell, 1997 e Kielan-Jaworowska & Hurum, 2001 *** Family † Cimolodontidae Marsh, 1889 sensu Kielan-Jaworowska & Hurum, 2001 **** Genus †'' Liotomus'' 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'' Xu et al., 2015 ***** Species †'' Yubaatar zhongyuanensis'' Xu et al., 2015 **** Genus †'' Anconodon'' 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'' 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, uncertaint ...
'' **** Genus '' Neoliotomus'' Jepsen, 1930 ***** Species †'' N. conventus'' Jepsen, 1930 ***** Species †'' N. ultimus'' (Granger & Simpson, 1928) *** Family † Neoplagiaulacidae Ameghino, 1890 tilodontidae: Neoplagiaulacinae Ameghino, 1890 sensu McKenna & Bell, 1997**** Genus †'' Mesodma'' 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'' 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'' 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'' 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'' Vianey-Liaud, 1986 ***** Species †'' C. manueli'' Vianey-Liaud, 1986 ***** Species †'' C. davidi'' Vianey-Liaud, 1986 **** Genus †'' Krauseia'' Vianey-Liaud, 1986 ***** Species †'' K. clemensi'' Sloan, 1981 'Parectypodus clemensi'' Sloan, 1981**** Genus †'' Xyronomys''Rigby, 1980 ***** Species †'' X. swainae'' Rigby, 1980 ironomys (sic); ?Eucosmodontidae**** Genus †'' Xanclomys'' Rigby, 1980 ***** Species †'' X. mcgrewi''Rigby, 1980 **** Genus †'' Mesodmops'' Tong & Wang, 1994 ***** Species †'' M. dawsonae'' Tong & Wang, 1994 *** Family † Ptilodontidae 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'' 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'' Jepsen, 1940 ***** Species †'' P. cavus'' Jespen, 1940 ***** Species †'' P. foxi'' Krause, 1987 ***** Species †'' P. taxus'' Krause, 1987 ***** Species? †'' P. speirsae'' Scott, 2004 ** Superfamily † Taeniolabidoidea Granger & Simpson, 1929 sensu Kielan-Jaworowska & Hurum, 2001 *** Genus †'' Prionessus'' Matthew & Granger, 1925 **** Species †'' P. lucifer'' Matthew & Granger, 1925 *** Family † Lambdopsalidae **** Genus †'' Lambdopsalis'' Chow & Qi, 1978 ***** Species †'' L. bulla'' Chow & Qi, 1978 **** Genus †'' Sphenopsalis'' Matthew, Granger & Simpson, 1928 ***** Species †'' S. nobilis'' Matthew, Granger & Simpson, 1928 *** Family † Taeniolabididae Granger & Simpson, 1929 **** Genus †'' Taeniolabis'' Cope, 1882 ***** Species †'' T. lamberti'' Simmons, 1987 ***** Species †'' T. taoensis'' Cope, 1882 **** Genus †'' Kimbetopsalis'' ***** Species †'' K. simmonsae'' ** Superfamily † Djadochtatherioidea Kielan-Jaworowska & Hurum, 1997 sensu Kielan-Jaworowska & Hurum, 2001 jadochtatheria Kielan-Jaworowska & Hurum, 1997*** Genus? †'' Bulganbaatar'' Kielan-Jaworowska, 1974 **** Species? †'' B. nemegtbaataroides'' Kielan-Jaworowska, 1974 *** Genus †'' Nemegtbaatar'' Kielan-Jaworowska, 1974 **** Species? †'' N. gobiensis'' Kielan-Jaworowska, 1974 *** Family † Chulsanbaataridae Kielan-Jaworowska, 1974 **** Genus †'' Chulsanbaatar'' Kielan-Jaworowska, 1974 ***** Species †'' C. vulgaris'' Kielan-Jaworowska, 1974 *** Family † Sloanbaataridae Kielan-Jaworowska, 1974 **** Genus †'' Kamptobaatar'' Kielan-Jaworowska, 1970 ***** Species? †'' K. kuczynskii'' Kielan-Jaworowska, 1970 **** Genus †'' Nessovbaatar'' Kielan-Jaworowska & Hurum, 1997 ***** Species †'' N. multicostatus'' Kielan-Jaworowska & Hurum, 1997 **** Genus †'' Sloanbaatar'' Kielan-Jaworowska, 1974 ***** Species †'' S. mirabilis'' Kielan-Jaworowska, 1974 loanbaatarinae*** Family † Djadochtatheriidae Kielan-Jaworowska $ Hurum, 1997 **** Genus †'' Djadochtatherium'' 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'' Kielan-Jaworowska, 1974 ***** Species †'' T. sabuli'' Rougier, Novacek & Dashzeveg, 1997 **** Genus †'' Kryptobaatar'' 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 ''palai ...
, 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'', "da ...
, 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 (biology), 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 roden ...
(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), 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, 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 t ...
. 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 area ...
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