Haramiyida ("thief" from Arabic الحرامية (al ḥarāmiyah), "thief, bandit") is a possibly
polyphyletic order of
mammaliaform cynodont
The cynodonts () (clade Cynodontia) are a clade of eutheriodont therapsids that first appeared in the Late Permian (approximately 260 mya), and extensively diversified after the Permian–Triassic extinction event. Cynodonts had a wide varie ...
s or
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 of controversial taxonomic affinites. Their teeth, which are by far the most common remains, resemble those of the
multituberculates. However, based on ''
Haramiyavia
''Haramiyavia'' ("thief grandmother" from Arabic الحرامية (al ḥarāmiyah), "thief, '' Haramiya''" + Latin avia, "grandmother") is a genus of synapsid in the clade Haramiyida that existed about 200 million years ago in the Rhaetian st ...
'', the jaw is less derived; and at the level of evolution of earlier basal mammals like ''
Morganucodon'' and ''
Kuehneotherium'', with a groove for
ear ossicles on the dentary. If they are early multituberculates, they would be the longest lived mammalian clade of all time. However, a more recent study in November 2015 may dispute this and suggested the Haramiyida were not crown mammals, but were part of an earlier offshoot of mammaliaformes instead.
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.
Relationships
Haramiyids show certain similarities to multituberculates, a group of mammals that survived until about 40 million years ago. It is possible that haramiyids are ancestral to multituberculates, although the available evidence is insufficient to be conclusive. Certain characteristics of the teeth seem to rule out a special relationship between the two groups, although some studies still unite haramiyids (or at least
euharamiyids) and multituberculates in the
Allotheria hypothesis.
In a 2018 study, haramiyidans have been found to be a monophyletic group of non-mammalian
Mammaliaformes
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 ...
. In this study,
gondwanatheres – usually interpreted as mammals, and derived
multituberculates in particular – have been found to be deeply nested among them.
[Huttenlocker, 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.]
Taxonomy
Order †Haramiyida
[Mikko's Phylogeny Archiv]
Hahn, Sigogneau-Russell & Wouters, 1989 Hahn, 1973 sensu McKenna & Bell, 1997">aramiyoidea Hahn, 1973 sensu McKenna & Bell, 1997* †''
Kirtlingtonia''
Butler & Hooker, 2005
*Family †
Haramiyaviidae Butler, 2000
** †''
Haramiyavia
''Haramiyavia'' ("thief grandmother" from Arabic الحرامية (al ḥarāmiyah), "thief, '' Haramiya''" + Latin avia, "grandmother") is a genus of synapsid in the clade Haramiyida that existed about 200 million years ago in the Rhaetian st ...
''
Jenkins ''et al.'', 1997
*Family †
Theroteinidae Sigogneau-Russell, Frank & Hammerle, 1986
** †''
Theroteinus nikolai
''Theroteinus'' is an extinct genus of haramiyidan mammaliaforms from the Late Triassic of France and Britain. It contains three species: ''T. nikolai,'' ''T. rosieriensis'' and ''T. jenkinsi'', the former two of which are known exclusively from ...
''
Sigogneau-Russell, Frank & Hammerle, 1986
** †''
Theroteinus rosieriensis
''Theroteinus'' is an extinct genus of haramiyidan mammaliaforms from the Late Triassic of France and Britain. It contains three species: ''T. nikolai,'' ''T. rosieriensis'' and ''T. jenkinsi'', the former two of which are known exclusively from ...
''
Sigogneau-Russell, 2016
*Family †
Haramiyidae Poche, 1908 Simpson, 1947 sensu Jenkins ''et al.'', 1997; Microlestidae Murry, 1866; Microcleptidae Simpson, 1928">aramiyidae Simpson, 1947 sensu Jenkins ''et al.'', 1997; Microlestidae Murry, 1866; Microcleptidae Simpson, 1928** †''
Eoraetica''
**''
Hypsiprymnopsis rhaeticus''
Dawkins, 1864 Dawkins, 1864">'Microlestes rhaeticus'' Dawkins, 1864** †''
Avashishta bacharamensis''
Anantharaman ''et al.'', 2006
** ?†''
Allostaffia aenigmatica''
(Heinrich, 1999) Heinrich 2004 Heinrich, 1999 non Schubert, 1911; ''Staffia aenigmatica'' Heinrich, 1999">'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 (Owen, 1871) Simpson, 1947; ''Microleptes moorei'' Owen, 1871; ''Microcleptes moorei'' (Owen, 1871) Simpson, 1928">'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.'', 2018
**† ''Denisodon''
**† ''
Hahnodon taqueti
''Hahnodon'' ("Hahn's tooth") is an extinct genus of mammaliaforms from the Early Cretaceous Ksar Metlili Formation in Morocco. Although originally considered to be a relatively early member of the extinct clade Multituberculata, recent studie ...
''
Sigogneau-Russell, 1991
*†
Euharamiyida Bi et al., 1994
*?†
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 ...
Mones, 1987
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
''Haramiyavia'' ("thief grandmother" from Arabic الحرامية (al ḥarāmiyah), "thief, '' Haramiya''" + Latin avia, "grandmother") is a genus of synapsid in the clade Haramiyida that existed about 200 million years ago in the Rhaetian st ...
'', 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 haramiyidans, ''
Maiopatagium'', ''
Xianshou'', ''
Vilevolodon'' and ''
Arboroharamiya
''Arboroharamiya'' ("tree thief" from Latin arbor, "tree" + Arabic الحرامية (al ḥarāmiyah), "thief, '' Haramiya''") is an extinct genus of early mammal (or possibly a non-mammalian mammaliaform) from the Middle Jurassic Tiaojishan Form ...
'', took it one step further and developed the ability to
glide, having extensive membranes similar to those of modern
colugos
Colugos () are arboreal gliding mammals that are native to Southeast Asia. Their closest evolutionary relatives are primates. There are just two living species of colugos: the Sunda flying lemur (''Galeopterus variegatus'') and the Philippine fly ...
. In many of these taxa, the coracoid bones (absent in modern
therians 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 ICS geologic timescale, the latest age (uppermost stage) of the Late Cretaceous Epoch or Upper Cretaceous Series, the Cretaceous Period or System, and of the Mesozoic Era or Erathem. It spanned the inte ...
of India,
however, this has not been robustly assessed by phylogenetics.
The youngest definitive euharamiyidan is ''
Cryoharamiya'' from the Early Creatceous
Batylykh Formation
The Batylykh Formation is a geological formation in Yakutia, Russia. It is of an uncertain Early Cretaceous age, probably dating between the Berriasian and the Barremian. It is the oldest unit of the thick Sangar Series within the Vilyuy synec ...
of
Yakutia, 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, and affiliated with Columbia University. It is currently directed by Jennifer Crewe (2014–present) and publishes titles in the humanities and sciences, including the fie ...
, 2004), 249–260.
External links
Palaeos, Mammaliaformes: AllotheriaJohn H Burkitt, Mammals, A World Listing of Living and Extinct SpeciesArchivedfrom 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