Haramiyida is a possibly
polyphyletic order of
mammaliaformcynodonts or
mammals of controversial taxonomic affinites.[2] 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.[3] Some authors have placed them in a clade with Multituberculata dubbed
Allotheria within Mammalia.[4][5] Other studies have disputed this and suggested the Haramiyida were not crown mammals, but were part of an earlier offshoot of mammaliaformes instead.[6] 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.[7]
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,[8] although some studies still unite haramiyids (or at least
euharamiyids) and multituberculates in the
Allotheria hypothesis.[9]
In a 2018 study, haramiyidans have been found to be a monophyletic group of non-mammalian
Mammaliaformes. In this study,
gondwanatheres – usually interpreted as mammals, and derived
multituberculates in particular – have been found to be deeply nested among them.[10]
Taxonomy
Order †Haramiyida[11][12] Hahn, Sigogneau-Russell & Wouters, 1989 [Haramiyoidea Hahn, 1973 sensu McKenna & Bell, 1997]
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,[7] 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.[7]
^MUSSER, A.M., LAMANNA, M.C., MARTINELLI, A.G., SALISBURY, S.W.,
AHYONG, S. & JONES, R., 2019. The first non-mammalian
cynodonts from Australia and the unusual nature of Australian
Cretaceous terrestrial tetrapod faunas. Journal of Vertebrate
Paleontology 39, Society of Vertebrate Paleontology Annual Meeting
Abstracts, 157.
^
abHuttenlocker, 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.
^Mikko's Phylogeny Archive
[1] Haaramo, Mikko (2007).
"†Haramiyida". Retrieved 30 December 2015.
^Nicholas Chimento, Frederico Agnolin, Agustin Martinelli, Mesozoic Mammals from South America: Implications for understanding early mammalian faunas from Gondwana, May 2016
^David M. Grossnickle, P. David Polly, Mammal disparity decreases during the Cretaceous angiosperm radiation, Published 2 October 2013.DOI: 10.1098/rspb.2013.2110
^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
^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
^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.
doi:
10.1671/0272-4634(2006)26[488:aplchf]2.0.co;2.
S2CID41722902.
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, 2004), 249–260.