Haramiyida
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Haramiyida is a possibly
paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
order of
mammaliaform Mammaliaformes ("mammalian forms") is a clade of synapsid tetrapods that includes the crown group mammals and their closest extinct relatives; the group radiated from earlier probainognathian cynodonts during the Late Triassic. It is defined a ...
cynodont Cynodontia () is a clade of eutheriodont therapsids that first appeared in the Late Permian (approximately 260 Megaannum, mya), and extensively diversified after the Permian–Triassic extinction event. Mammals are cynodonts, as are their extin ...
s or
mammal A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...
s of controversial taxonomic affinites. Their teeth, which are by far the most common remains, resemble those of the
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. They first appeared in the Middle Jurassic, a ...
s. However, based on '' Haramiyavia'', the jaw is less derived; and at the level of evolution of earlier basal mammals like ''
Morganucodon ''Morganucodon'' ("Glamorgan tooth") is an early mammaliaform genus that lived from the Late Triassic to the Middle Jurassic. It first appeared about 205 million years ago. Unlike many other early mammaliaforms, ''Morganucodon'' is well represent ...
'' and '' Kuehneotherium'', with a groove for ear ossicles on the dentary. Some authors have placed them in a clade with Multituberculata dubbed
Allotheria Allotheria (meaning "other beasts", from the Ancient Greek language, Greek , '–other and , '–wild animal) is an extinct clade of mammals known from the Mesozoic and early Cenozoic. Shared characteristics of the group are the presence of lower ...
within Mammalia. Other studies have disputed this and suggested the Haramiyida were not
crown group In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor ...
mammals, but were part of an earlier offshoot of Mammaliaformes instead, either closely related or unrelated to Multituberculates. It is also disputed whether the Late Triassic species are closely related to the Jurassic and Cretaceous members belonging to Euharamiyida/Eleutherodontida, as some phylogenetic studies recover the two groups as unrelated, recovering the Triassic haramiyidians as non-mammalian cynodonts, while recovering the Euharamiyida as crown-group mammals closely related to multituberculates.


Taxonomy

Order †HaramiyidaMikko's Phylogeny Archiv

Hahn, Sigogneau-Russell & Wouters, 1989 aramiyoidea Hahn, 1973 sensu McKenna & Bell, 1997* †'' Kirtlingtonia'' Butler & Hooker, 2005 *Family † Haramiyaviidae Butler, 2000 ** †'' Haramiyavia'' Jenkins ''et al.'', 1997 *Family † Theroteinidae Sigogneau-Russell, Frank & Hammerle, 1986 ** †'' Theroteinus nikolai'' Sigogneau-Russell, Frank & Hammerle, 1986 ** †'' Theroteinus rosieriensis'' Sigogneau-Russell, 2016 *Family † Haramiyidae Poche, 1908 aramiyidae Simpson, 1947 sensu Jenkins ''et al.'', 1997; Microlestidae Murry, 1866; Microcleptidae Simpson, 1928** †'' Eoraetica'' **'' Hypsiprymnopsis rhaeticus'' Dawkins, 1864 'Microlestes rhaeticus'' Dawkins, 1864** †'' Avashishta bacharamensis'' Anantharaman ''et al.'', 2006 ** ?†'' Allostaffia aenigmatica'' (Heinrich, 1999) Heinrich 2004 'Staffia'' Heinrich, 1999 non Schubert, 1911; ''Staffia aenigmatica'' Heinrich, 1999 possible, gondwanathere instead. ** †'' Thomasia'' Poche, 1908 'Haramiya'' Simpson, 1947; ''Microlestes'' Plieninger, 1847 non Schmidt-Goebel, 1846; ''Microcleptes'' Simpson, 1928 non Newman, 1840; ''Plieningeria'' Krausse, 1919; ''Stathmodon'' Henning, 1911] *** †''Thomasia (animal), T. woutersi'' Butler & MacIntyre, 1994 *** †''Thomasia (animal), T. hahni'' Butler & MacIntyre, 1994 *** †''Haramiya, T. moorei'' (Owen 1871) Butler & MacIntyre, 1994 'Haramiya moorei'' (Owen, 1871) Simpson, 1947; ''Microleptes moorei'' Owen, 1871; ''Microcleptes moorei'' (Owen, 1871) Simpson, 1928*** †'' T. antiqua'' (Plieninger, 1847) Poche 1908 [''Microlestes antiquus'' Plieninger, 1847; ''Haramiya antiqua'' (Plieninger, 1847); ''Microleptes fissurae'' Simpson, 1928; ''Haramiya fissurae'' (Simpson 1928); ''Haramiya butleri'' Sigogneau-Russell, 1990; ''Thomasia anglica'' Simpson, 1928] *†Hahnodontidae Sigogneau-Russell, 1991 **†''Cifelliodon wahkarmoosuch'' Huttenlocker ''et al.'', 2018Huttenlocker, Adam K.; Grossnickle, David M.; Kirkland, James, I.; Schultz, Julia A.; Luo, Zhe-Xi (23 May 2018). "Late-surviving stem mammal links the lowermost Cretaceous of North America and Gondwana". ''Nature''. 558 (7708): 108–112. doi:10.1038/s41586-018-0126-y. Retrieved 23 May 2018. **† '' Denisodon'' **† '' Hahnodon taqueti'' Sigogneau-Russell, 1991 *† Euharamiyida Bi et al., 1994 *?† Gondwanatheria Mones, 1987


Relationships

The relationships of haramiydans to other mammals and mammaliaform cynodonts are controversial and have been subject to numerous conflicting phylogenetic analysis results. Major unresolved questions are whether or not haramiyidans are more closely related to
marsupial Marsupials are a diverse group of mammals belonging to the infraclass Marsupialia. They are natively found in Australasia, Wallacea, and the Americas. One of marsupials' unique features is their reproductive strategy: the young are born in a r ...
s and
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 distinguished ...
mammals (
Theria Theria ( or ; ) is a scientific classification, subclass of mammals amongst the Theriiformes. Theria includes the eutherians (including the Placentalia, placental mammals) and the metatherians (including the marsupials) but excludes the egg-lay ...
) than they are to
monotremes Monotremes () are mammals of the order Monotremata. They are the only group of living mammals that lay eggs, rather than bearing live young. The extant monotreme species are the platypus and the four species of echidnas. Monotremes are typified ...
(and thus inside the
crown group In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor ...
of
Mammalia A mammal () is a vertebrate animal of the class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three middle ear bon ...
), or whether all living mammals (including therians and monotremes) are more closely related to each other than to haramiyidans (and thus placing Haramiyida outside crown Mammalia) and whether or not haramiyidans are closely related to multituberculates, an important of group of Mesozoic and early Cenozoic mammaliaforms typically regarded as crown group mammals, as part of the group
Allotheria Allotheria (meaning "other beasts", from the Ancient Greek language, Greek , '–other and , '–wild animal) is an extinct clade of mammals known from the Mesozoic and early Cenozoic. Shared characteristics of the group are the presence of lower ...
. While many studies recover Triassic haramiyidans and Jurassic euharamiyidans as closely related, some phylogenetic studies have recovered them as unrelated, find the Triassic haramiyidians as non-mammalian cynodonts, while recovering the Euharamiyida as crown-group mammals closely related to multituberculates. Cladogram from Luo et al 2017, showing a monophyletic Haramiyida outside of crown Mammalia unrelated to Multituberculata: Cladogram from Han et al. 2017, showing a paraphyletic Haramiyida within crown Mammalia as ancestral to Multituberculata: Cladogram from Hoffmann et al. 2020, showing a diphyletic Haramiyida. Simplified cladogram from Mao et al. 2024, showing a paraphyletic Haramiyida closely related to Multituberculata outside of crown Mammalia:


Lifestyle

Haramiyids seem to have generally been herbivorous or omnivorous, possibly the first mammalian herbivores; however, the sole haramiyid tested in a study involving Mesozoic mammal dietary habits, '' Haramiyavia'', ranks among insectivorous species. At least some species were very good climbers and were similar to modern day squirrels; and several others have more recently been reassessed as possibly arboreal. General arboreal habits might explain their rarity in the fossil record. Several euharamiyidans, ''
Maiopatagium ''Maiopatagium'' is an extinct genus of gliding euharamiyida, euharamiyids which existed in Asia during the Jurassic period. It possessed a patagium between its limbs and presumably had similar lifestyle to living Flying squirrel, flying squirrel ...
'', '' Xianshou'', ''
Vilevolodon ''Vilevolodon'' is an extinct, monotypic genus of volant, arboreal euharamiyids from the Oxfordian age of the Late Jurassic of China. The type species is ''Vilevolodon diplomylos''. The genus name ''Vilevolodon'' references its gliding capabili ...
'' and ''
Arboroharamiya ''Arboroharamiya'' is an extinct genus of mammaliaform from the Late Jurassic Tiaojishan Formation of Inner Mongolia, China. ''Arboroharamiya'' belongs to a group of mammaliaforms called Haramiyida. The genus contains three species: ''A. jenkins ...
'', took it one step further and developed the ability to glide, having extensive membranes similar to those of modern colugos. In many of these taxa, the coracoid bones (absent in modern
theria Theria ( or ; ) is a scientific classification, subclass of mammals amongst the Theriiformes. Theria includes the eutherians (including the Placentalia, placental mammals) and the metatherians (including the marsupials) but excludes the egg-lay ...
ns but present in many other mammal groups, albeit highly reduced) are remarkably large and similar to those of birds and pterosaurs, presumably due to impact stresses at landing. Mammalian tooth marks on dinosaur bones may belong to '' Sineleutherus'', suggesting that some haramiyidans scavenged on dinosaur remains.


Range

The fossils of Late Triassic Haramiyids are primarily known from Europe and Greenland, while the fossils of Euharamiyids are primarily known from the Middle to Late Jurassic of Asia. Remains of eleutherodontids from Europe are only known from isolated teeth. The youngest haramiyid fossil genus has been considered to be possibly be '' Avashishta bacharamensis'' from the
Maastrichtian The Maastrichtian ( ) is, in the International Commission on Stratigraphy (ICS) geologic timescale, the latest age (geology), age (uppermost stage (stratigraphy), stage) of the Late Cretaceous epoch (geology), Epoch or Upper Cretaceous series (s ...
of India, however, this has not been robustly assessed by phylogenetics. The youngest definitive euharamiyidan is '' Cryoharamiya'' from the Early Cretaceous Batylykh Formation of
Yakutia Sakha, officially the Republic of Sakha (Yakutia), is a republics of Russia, republic of Russia, and the largest federal subject of Russia by area. It is located in the Russian Far East, along the Arctic Ocean, with a population of one million ...
, Russia.


Notes


References

* Zofia Kielan-Jaworowska, Richard L. Cifelli, and Zhe-Xi Luo, ''Mammals from the Age of Dinosaurs: Origins, Evolution, and Structure'' (New York:
Columbia University Press Columbia University Press is a university press based in New York City New York, often called New York City (NYC), is the most populous city in the United States, located at the southern tip of New York State on one of the world's la ...
, 2004), 249–260.


External links


Palaeos, Mammaliaformes: Allotheria

John H Burkitt, Mammals, A World Listing of Living and Extinct SpeciesArchived
from the original on 21 December 2004. Retrieved 2015-06-18.

{{Taxonbar, from=Q137933 Late Triassic synapsids Jurassic synapsids Transitional fossils Norian first appearances Middle Jurassic extinctions Taxa named by Gerhard Hahn (palaeontologist) Taxa named by Denise Sigogneau-Russell