Haramiyida is a possibly Paraphyly order of Mammaliaformes or of controversial taxonomic affinites.[Butler PM, 2000] Some authors have placed them in a clade with Multituberculata dubbed Allotheria within Mammalia.
Taxonomy
Order †
Haramiyida[Mikko's Phylogeny Archive [1] ][Paleofile.com (net, info) [2]. ] Hahn, Sigogneau-Russell & Wouters, 1989 Haramiyoidea
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† Kirtlingtonia Butler & Hooker, 2005
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Family †Haramiyaviidae Butler, 2000
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Family †Theroteinidae Sigogneau-Russell, Frank & Hammerle, 1986
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Family †Haramiyidae Poche, 1908 Haramiyidae
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† Eoraetica
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Hypsiprymnopsis rhaeticus Dawkins, 1864
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† Avashishta bacharamensis Anantharaman et al., 2006
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?† Allostaffia aenigmatica (Heinrich, 1999) Heinrich 2004 Staffia; possible, gondwanathere instead.
[Nicholas Chimento, Frederico Agnolin, Agustin Martinelli, Mesozoic Mammals from South America: Implications for understanding early mammalian faunas from Gondwana, May 2016]
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† Thomasia Poche, 1908 ''
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† T. woutersi Butler & MacIntyre, 1994
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† T. hahni Butler & MacIntyre, 1994
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† Haramiya (Owen 1871) Butler & MacIntyre, 1994 ''Haramiya
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† T. antiqua (Plieninger, 1847) Poche 1908 ''Microlestes
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†Hahnodontidae Sigogneau-Russell, 1991
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† Cifelliodon wahkarmoosuch Huttenlocker et al., 2018
[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.]
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† Denisodon
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† Hahnodon Sigogneau-Russell, 1991
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
and
placental mammals (
Theria) than they are to
monotremes (and thus inside the
crown group of
Mammalia), 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. 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.
[David M. Grossnickle, P. David Polly, Mammal disparity decreases during the Cretaceous angiosperm radiation, Published 2 October 2013.DOI: 10.1098/rspb.2013.2110] 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.
[Jing Meng, Mesozoic mammals of China: implications for phylogeny and early evolution of mammals, Natl Sci Rev (December 2014) 1 (4): 521-542.
doi: 10.1093/nsr/nwu070
First published online: October 17, 2014]
Several euharamiyidans, Maiopatagium, Xianshou, Vilevolodon and Arboroharamiya, took it one step further and developed the ability to gliding flight, having extensive membranes similar to those of modern colugos. In many of these taxa, the coracoid bones (absent in modern 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.[Qing-Jin Meng; David M. Grossnickle; Di Liu; Yu-Guang Zhang; April I. Neander; Qiang Ji; Zhe-Xi Luo (2017). "New gliding mammaliaforms from the Jurassic". Nature. in press. doi:10.1038/nature23476.][A Jurassic gliding euharamiyidan mammal with an ear of five auditory bones, Nature doi:10.1038/nature24483]
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 of India,
Notes
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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, 2004), 249–260.
External links
