Haramiyida Explained

Haramiyida is a possibly paraphyletic order of mammaliaform cynodonts or mammals of controversial taxonomic affinites.[1] Their teeth, which are by far the most common remains, resemble those of the multituberculates. However, based on Haramiyavia, 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.[2] Some authors have placed them in a clade with Multituberculata dubbed Allotheria within Mammalia.[3] [4] Other studies have disputed this and suggested the Haramiyida were not crown mammals, but were part of an earlier offshoot of mammaliaformes instead.[5] 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.[6]

Taxonomy

Order †Haramiyida[7] [8] Hahn, Sigogneau-Russell & Wouters, 1989 [Haramiyoidea <small>Hahn, 1973 sensu McKenna & Bell, 1997</small>]

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 marsupials 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 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:[13] Cladogram from Han et al. 2017, showing a paraphyletic Haramiyida within crown Mammalia as ancestral to Multituberculata:[14] 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:[15]

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.[16] At least some species were very good climbers and were similar to modern day squirrels;[17] and several others have more recently been reassessed as possibly arboreal. General arboreal habits might explain their rarity in the fossil record.[18]

Several euharamiyidans, Maiopatagium, Xianshou, Vilevolodon and Arboroharamiya, 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 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.[19] [20]

Mammalian tooth marks on dinosaur bones may belong to Sineleutherus, suggesting that some haramiyidans scavenged on dinosaur remains.[21]

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.[22] 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,[23] however, this has not been robustly assessed by phylogenetics. The youngest definitive euharamiyidan is Cryoharamiya from the Early Cretaceous Batylykh Formation of Yakutia, Russia.

References

External links

Notes and References

  1. Zheng. Xiaoting. Bi. Shundong. Wang. Xiaoli. Meng. Jin. A new arboreal haramiyid shows the diversity of crown mammals in the Jurassic period. Nature. 500. 7461. 2013. 199–202. 0028-0836. 10.1038/nature12353. 23925244. 2013Natur.500..199Z. 2164378.
  2. Butler PM, 2000
  3. Meng . Jin . Bi . Shundong . Wang . Yuanqing . Zheng . Xiaoting . Wang . Xiaoli . 2014-12-10 . Evans . Alistair R. . Dental and Mandibular Morphologies of Arboroharamiya (Haramiyida, Mammalia): A Comparison with Other Haramiyidans and Megaconus and Implications for Mammalian Evolution . PLOS ONE . en . 9 . 12 . e113847 . 10.1371/journal.pone.0113847 . 1932-6203 . 4262285 . 25494181 . free .
  4. Bi . Shundong . Wang . Yuanqing . Guan . Jian . Sheng . Xia . Meng . Jin . 2014-10-30 . Three new Jurassic euharamiyidan species reinforce early divergence of mammals . Nature . en . 514 . 7524 . 579–584 . 10.1038/nature13718 . 25209669 . 4471574 . 0028-0836.
  5. Luo . Zhe-Xi . Gates . Stephen M. . Jenkins Jr. . Farish A. . Amaral . William W. . Shubin . Neil H. . Mandibular and dental characteristics of Late Triassic mammaliaform Haramiyavia and their ramifications for basal mammal evolution . 10.1073/pnas.1519387112 . 26630008 . 16 November 2015 . . 112 . 51 . E7101–E7109. 4697399 . 2015PNAS..112E7101L . free .
  6. Hoffmann . Simone . Beck . Robin M. D. . Wible . John R. . Rougier . Guillermo W. . Krause . David W. . 2020-12-14 . Phylogenetic placement of Adalatherium hui (Mammalia, Gondwanatheria) from the Late Cretaceous of Madagascar: implications for allotherian relationships . Journal of Vertebrate Paleontology . 40 . sup1 . 213–234 . 10.1080/02724634.2020.1801706 . 230968231 . 0272-4634.
  7. Mikko's Phylogeny Archive http://www.helsinki.fi/~mhaaramo/ Web site: Haaramo . Mikko . 2007 . †Haramiyida . 30 December 2015.
  8. Paleofile.com (net, info) http://www.paleofile.com/. Web site: Taxonomic lists – Mammals . 30 December 2015.
  9. Debuysschere . Maxime . 2016 . A reappraisal of Theroteinus (Haramiyida, Mammaliaformes) from the Upper Triassic of Saint-Nicolas-de-Port (France) . PeerJ . 4 . e2592 . 10.7717/peerj.2592 . 27781174 . 5075691 . free .
  10. W.B. . Dawkins . 1864 . On the Rhaetic beds and White Lias of western and central Somerset; and on the discovery of a new fossil mammal in the grey marlstones below the bone-bed . Quarterly Journal of the Geological Society . 20 . 396–412 . 10.1144/GSL.JGS.1864.020.01-02.49 . 128992830 .
  11. Nicholas Chimento, Frederico Agnolin, Agustin Martinelli, Mesozoic Mammals from South America: Implications for understanding early mammalian faunas from Gondwana, May 2016
  12. 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.
  13. Luo . Zhe-Xi . Meng . Qing-Jin . Grossnickle . David M. . Liu . Di . Neander . April I. . Zhang . Yu-Guang . Ji . Qiang . August 2017 . New evidence for mammaliaform ear evolution and feeding adaptation in a Jurassic ecosystem . Nature . en . 548 . 7667 . 326–329 . 10.1038/nature23483 . 0028-0836.
  14. Han . Gang . Mao . Fangyuan . Bi . Shundong . Wang . Yuanqing . Meng . Jin . November 2017 . A Jurassic gliding euharamiyidan mammal with an ear of five auditory bones . Nature . en . 551 . 7681 . 451–456 . 10.1038/nature24483 . 0028-0836.
  15. Mao . Fangyuan . Li . Zhiyu . Wang . Zhili . Zhang . Chi . Rich . Thomas . Vickers-Rich . Patricia . Meng . Jin . 2024-04-18 . Jurassic shuotheriids show earliest dental diversification of mammaliaforms . Nature . en . 628 . 8008 . 569–575 . 10.1038/s41586-024-07258-7 . 0028-0836.
  16. David M. Grossnickle, P. David Polly, Mammal disparity decreases during the Cretaceous angiosperm radiation, Published 2 October 2013.DOI: 10.1098/rspb.2013.2110
  17. Web site: ScienceDaily. 2014-09-11. Three extinct squirrel-like species discovered. 2014-10-07. https://web.archive.org/web/20140918044407/https://www.sciencedaily.com/releases/2014/09/140910132522.htm. 18 September 2014. live.
  18. 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/nwu070First published online: October 17, 2014
  19. 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.
  20. A Jurassic gliding euharamiyidan mammal with an ear of five auditory bones, Nature doi:10.1038/nature24483
  21. 10.1007/s00114-020-01688-9. The smallest eating the largest: The oldest mammalian feeding traces on dinosaur bone from the Late Jurassic of the Junggar Basin (Northwestern China). 2020. Augustin. Felix J.. Matzke. Andreas T.. Maisch. Michael W.. Hinz. Juliane K.. Pfretzschner. Hans-Ulrich. The Science of Nature. 107. 4. 32. 32686025. 7369264. 2020SciNa.107...32A.
  22. Averianov . Alexander O. . Martin . Thomas . Lopatin . Alexey V. . Skutschas . Pavel P. . Schellhorn . Rico . Kolosov . Petr N. . Vitenko . Dmitry D. . 2019-11-02 . A new euharamiyidan mammaliaform from the Lower Cretaceous of Yakutia, Russia . Journal of Vertebrate Paleontology . en . 39 . 6 . e1762089 . 10.1080/02724634.2019.1762089 . 220548470 . 0272-4634.
  23. Anantharaman . S. . Wilson . G. P. . Das Sarma . D. C. . Clemens . W. A. . 2006 . A possible Late Cretaceous "haramiyidan" from India . Journal of Vertebrate Paleontology . 26 . 2. 488–490 . 10.1671/0272-4634(2006)26[488:aplchf]2.0.co;2. 41722902 .

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