Haramiyida

Haramiyida Temporal range: Late Triassic–Late Cretaceous, 216.5–66 Ma PreꞒ Ꞓ O S D C P T J K Pg N
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Clade:	Synapsida
Clade:	Therapsida
Clade:	Cynodontia
Clade:	Mammaliaformes
Order:	†Haramiyida Hahn, Sigogneau-Russell & Wouters, 1989
Subgroups
†Kalaallitkigun †Kollikodon?[1] †Haramiyoidea †Haramiyidae †? † † †? †Thomasia †Haramiyaviidae †Haramiyavia † † †Hahnodontidae †Cifelliodon † †Hahnodon † † †Theroteinida †Theroteinidae †Theroteinus †Eleutherodontidae † †Sineleutherus †Arboroharamiya †Xianshou †Maiopatagium †Shenshouidae † † †Shenshou †Megaconus †Millsodon ?†Gondwanatheria

Haramiyidans were a long lived lineage of mammaliaform cynodonts or mammals.^[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] 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.^[4][5]

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,^[6] although some studies still unite haramiyids (or at least euharamiyids) and multituberculates in the Allotheria hypothesis.^[7]

Most recently, 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.^[8]

Taxonomy[]

Order †Haramiyida^[9][10] Hahn, Sigogneau-Russell & Wouters, 1989 [Haramiyoidea Hahn, 1973 sensu McKenna & Bell, 1997]

• † 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^[11]
• Family †Haramiyidae Poche, 1908 [Haramiyidae Simpson, 1947 sensu Jenkins et al., 1997; Microlestidae Murry, 1866; Microcleptidae Simpson, 1928]
  • †
  • † Dawkins, 1864^[12] [Microlestes rhaeticus Dawkins, 1864]
  • † Anantharaman et al., 2006
  • ?† (Heinrich, 1999) Heinrich 2004 [Staffia Heinrich, 1999 non Schubert, 1911; Staffia aenigmatica Heinrich, 1999]; possible, gondwanathere instead.^[13]
  • †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]
    • †T. woutersi Butler & MacIntyre, 1994
    • †T. hahni Butler & MacIntyre, 1994
    • †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]

• †Eleutherodontida Kermack et al., 1998
  • Family †Arboroharamiyidae Zheng et al., 2013
    • †Arboroharamiya jenkinsi Zheng et al., 2013
  • Family †Shenshouidae Mao and Meng, 2019
    • † Mao and Meng, 2019
    • †Shenshou lui Bi et al., 2014
  • Family †Eleutherodontidae Kermack et al., 1998
    • †Maiopatagium Luo et al., 2017
    • † Kermack et al., 1998
    • †Xianshou Bi et al., 2014
      • †X. linglong Bi et al., 2014
      • †X. songae Bi et al., 2014
    • †Sineleutherus Martin, Averianov & Pfretzschner, 2010
      • †S. uyguricus Martin, Averianov & Pfretzschner, 2010
      • †S. issedonicus Averianov, Lopatin & Krasnolutskii, 2011
    • †Vilevolodon Zhe-Xi Luo, 2017
      • †Vilevolodon diplomylos Zhe-Xi Luo, 2017
  • †Megaconus mammaliaformis Zhou et al., 2013
  • †Millsodon superstes Butler & Hooker, 2005 [tooth-taxa]

In May 2018, a new genus and species was named, and is believed to be an eleutherodontidan of the family Hahnodontidae:

• †Cifelliodon wahkarmoosuch Huttenlocker et al., 2018^[8]

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.^[14] At least some species were very good climbers and were similar to modern day squirrels;^[15] and several others have more recently been reassessed as possibly arboreal. General arboreal habits might explain their rarity in the fossil record.^[16]

Several haramiyidans, 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.^[17][18]

Mammalian tooth marks on dinosaur bones may belong to Sineleutherus, suggesting that some haramiyidans scavenged on dinosaur remains.^[19]

Range[]

Most fossils have been reported from Europe, but some are known from Africa and Greenland. Since 2005, the published range extended to Mongolia and China. The age of haramiyid fossils range from Upper Triassic up to Late Cretaceous. This has important implications: the fact that haramiyids were present in the Late Triassic supports the idea that crown-group mammals originated at least 208 million years ago, much earlier than some previous research suggests.

The youngest haramiyid fossil genus is from the Maastrichtian of India. Cretaceous haramiyids are previously known from the Early Cretaceous of Morocco, suggesting that these animals may have survived in gondwannan landmasses as relics; the presence of the similarly aged Cifelliodon in North America suggests that contact between both landmasses remained for a while. With a temporal range of at least 150 million years, they are the longest lived mammaliaform lineage.^[20][21]

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, 2004), 249–260.

External links[]

• Palaeos, Mammaliaformes: Allotheria
• John H Burkitt, Mammals, A World Listing of Living and Extinct Species. Archived from the original on 21 December 2004. Retrieved 2015-06-18.
• Mikko K Haaramo, Haramiyida