Volaticotherium
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''Volaticotherium antiquum'' (meaning "ancient gliding beast") is an extinct, gliding,
insectivorous A robber fly eating a hoverfly An insectivore is a carnivorous animal or plant that eats insects. An alternative term is entomophage, which can also refer to the human practice of eating insects. The first vertebrate insectivores were ...
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 or ...
that lived in
Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an area ...
during the
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The J ...
period, around 164 mya. It is the only member of the genus ''Volaticotherium''. The discovery of ''Volaticotherium'' provided the earliest-known record of a gliding mammal (70 million years older than the next oldest example), until the discovery of the contemporary
haramiyida Haramiyida ("thief" from Arabic الحرامية (al ḥarāmiyah), "thief, bandit") is a possibly polyphyletic order of mammaliaform cynodonts or mammals of controversial taxonomic affinites. Their teeth, which are by far the most common remains ...
ns ''
Maiopatagium ''Maiopatagium'' ("greater patagium" from Latin maius, “greater, larger” + Modern Latin patagium, “ patagium”) is an extinct genus of gliding euharamiyids which existed in Asia during the Jurassic period. It possessed a patagium between ...
'', ''
Vilevolodon ''Vilevolodon'' is an extinct, monotypic genus of volant, Arboreal locomotion, arboreal Euharamiyida, euharamiyids from the Oxfordian (stage), Oxfordian age of the Late Jurassic of China. The type species is ''Vilevolodon diplomylos''. The genus ...
'' and ''
Xianshou ''Xianshou'' is a genus of glidingQing-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. . haramiyidan synapsid known from the ...
'', and provided further evidence of mammalian diversity during the
Mesozoic The Mesozoic Era ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last era of Earth's geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceo ...
Era. The closely related and significantly older ''
Argentoconodon ''Argentoconodon'' (meaning "Argentina cone tooth") is an extinct genus of theriimorph mammal from the Cañadón Asfalto Formation of the Cañadón Asfalto Basin in Patagonia. When originally described, it was known only from a single molarifo ...
'' shows similar post-cranial adaptations for aerial locomotion also seen in ''Volaticotherium''.


Discovery

The only known fossil of ''Volaticotherium'' was recovered from the
Daohugou Beds The Haifanggou Formation (), also known as the Jiulongshan Formation (), is a fossil-bearing rock deposit located near Daohugou () village of Ningcheng County, in Inner Mongolia, northeastern China. The formation consists of coarse conglomerates ...
of Ningcheng County,
Inner Mongolia Inner Mongolia, officially the Inner Mongolia Autonomous Region, is an autonomous region of the People's Republic of China. Its border includes most of the length of China's border with the country of Mongolia. Inner Mongolia also accounts for a ...
,
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...
. The age of the Daohugou Beds is currently uncertain and the subject of debate, but most studies suggest an age of around 164 plus or minus 4 million years ago. The description was published in an issue of the journal ''
Nature Nature, in the broadest sense, is the physics, physical world or universe. "Nature" can refer to the phenomenon, phenomena of the physical world, and also to life in general. The study of nature is a large, if not the only, part of science. ...
''.


Description

''Volaticotherium'' had a
gliding Gliding is a recreational activity and competitive air sport in which pilots fly unpowered aircraft known as gliders or sailplanes using naturally occurring currents of rising air in the atmosphere to remain airborne. The word ''soaring'' is al ...
membrane, sometimes compared to modern-day
flying squirrel Flying squirrels (scientifically known as Pteromyini or Petauristini) are a tribe of 50 species of squirrels in the family Sciuridae. Despite their name, they are not in fact capable of full flight in the same way as birds or bats, but they ar ...
, that extended not just between the limbs and at least the tail base, but also the digits, "sandwiching" them. It was densely covered by
fur Fur is a thick growth of hair that covers the skin of mammals. It consists of a combination of oily guard hair on top and thick underfur beneath. The guard hair keeps moisture from reaching the skin; the underfur acts as an insulating blanket t ...
.Meng, J.; Hu, Y.-M.; Wang, Y.-Q.; Wang, X.-L.; Li, C.-K. (2007). "Corrigendum: A Mesozoic gliding mammal from northeastern China". Nature 446 (7131): 102. . . The tail was flat, increasing the airfoil, and the limbs were proportionally long, comparable to those of modern flying and gliding mammals. The toes were grasping, as typical for arboreal mammals; the hand, however, was poorly preserved and its anatomy is therefore unclear. The teeth of ''Volaticotherium'' were highly unusual, possessing long, curved, backwards-pointing cusps, possibly used for shearing; this, combined with the long canines, indicates a carnivorous diet, which at its small size was probably composed of insects. This is supported by a study ranking it among insectivorous taxa, while the related ''
Argentoconodon ''Argentoconodon'' (meaning "Argentina cone tooth") is an extinct genus of theriimorph mammal from the Cañadón Asfalto Formation of the Cañadón Asfalto Basin in Patagonia. When originally described, it was known only from a single molarifo ...
'' ranked in carnivorous taxa. It has been noted that most gliding mammals are predominantly herbivorous, which would make volaticothere carnivory truly exceptional. In particular, ''Volaticotherium'' itself has been compared to insectivorous
bats Bats are mammals of the order Chiroptera.''cheir'', "hand" and πτερόν''pteron'', "wing". With their forelimbs adapted as wings, they are the only mammals capable of true and sustained flight. Bats are more agile in flight than most bi ...
, and its femur has unique adaptations among mammals that make it resistant to flight stresses, and render terrestrial locomotion cumbersome.


Classification

The phylogenetic analysis conducted by the authors of the description of ''Volaticotherium antiquum'' recovered it as the
sister taxon In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and t ...
of the
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
that contained, among other taxa,
eutriconodonts Eutriconodonta is an order (biology), order of early mammals. Eutriconodonts existed in Asia, Africa, Europe, North America, North and South America during the Jurassic and the Cretaceous periods. The order was named by Kermack ''et al.'' in 1973 ...
,
multituberculates Multituberculata (commonly known as multituberculates, named for the multiple tubercles of their teeth) is an extinct order of rodent-like mammals with a fossil record spanning over 130 million years. They first appeared in the Middle Jurassic, ...
,
spalacotheriid Spalacotheriidae is a family of extinct mammals belonging to the paraphyletic group ' Symmetrodonta'. They lasted from the Early Cretaceous to the Campanian in North America, Europe, Asia and North Africa. Spalacotheriids are characterised by ha ...
and tinodontid "
symmetrodonta Symmetrodonta is a group of Mesozoic mammals and mammal-like synapsids characterized by the triangular aspect of the molars when viewed from above, and the absence of a well-developed talonid. The traditional group of 'symmetrodonts' ranges in a ...
ns", dryolestids,
metatheria Metatheria is a mammalian clade that includes all mammals more closely related to marsupials than to placentals. First proposed by Thomas Henry Huxley in 1880, it is a more inclusive group than the marsupials; it contains all marsupials as well ...
ns (including
marsupial Marsupials are any members of the mammalian infraclass Marsupialia. All extant marsupials are endemic to Australasia, Wallacea and the Americas. A distinctive characteristic common to most of these species is that the young are carried in a po ...
s) and
eutheria Eutheria (; from Greek , 'good, right' and , 'beast'; ) is the clade consisting of all therian mammals that are more closely related to placentals than to marsupials. Eutherians are distinguished from noneutherians by various phenotypic tra ...
ns (including placental mammals). As the analysis did not place ''Volaticotherium'' within any of the previously known main groups of Mesozoic mammals, the authors of its description erected a separate family Volaticotheriidae and order Volaticotheria for it. However,
Zhe-Xi Luo Zhe-Xi Luo (Chinese 骆泽喜; pinyin: Luò Zéxǐ) is an American paleontologist of Chinese origin, specializing in vertebrate paleontology, particularly mammal evolution, morphology, and systematics. Background Born in China in 1958, he ca ...
(2007) mentioned that ''Volaticotherium'' might actually be a eutriconodont. This was eventually confirmed by the phylogenetic analyses conducted by Leandro C. Gaetano and Guillermo W. Rougier (2011, 2012); these analyses recovered ''Volaticotherium antiquum'' as a eutriconodont that belonged to the family
Triconodontidae Triconodontidae is an extinct family of small, carnivorous mammals belonging to the order Eutriconodonta, endemic to what would become Asia, Europe, North America and probably also Africa and South America during the Jurassic through Cretaceous p ...
and subfamily Alticonodontinae, and was particularly closely related to the genera ''
Argentoconodon ''Argentoconodon'' (meaning "Argentina cone tooth") is an extinct genus of theriimorph mammal from the Cañadón Asfalto Formation of the Cañadón Asfalto Basin in Patagonia. When originally described, it was known only from a single molarifo ...
'' and ''
Ichthyoconodon ''Ichthyoconodon'' is an extinct genus of eutriconodont mammal from the Lower Cretaceous of Morocco. It is notable for having been found in a unique marine location, and the shape of its teeth suggests an unusual, potentially fish-eating ecologi ...
'', and possibly ''
Triconolestes ''Triconolestes'' is an extinct genus of Late Jurassic eutriconodont mammal from the Morrison Formation, present in stratigraphic zones 4.Foster, J. (2007). "Appendix." ''Jurassic West: The Dinosaurs of the Morrison Formation and Their World''. ...
''.A. O. Averianov and A. V. Lopatin. 2011. Phylogeny of Triconodonts and Symmetrodonts and the Origin of Extant Mammals. Doklady Biological Sciences 436:32-35 . Uhen/M. Uhen/ref> More recent studies keep this relationship, but move ''Volaticotherium'', ''
Argentoconodon ''Argentoconodon'' (meaning "Argentina cone tooth") is an extinct genus of theriimorph mammal from the Cañadón Asfalto Formation of the Cañadón Asfalto Basin in Patagonia. When originally described, it was known only from a single molarifo ...
'', ''
Ichthyoconodon ''Ichthyoconodon'' is an extinct genus of eutriconodont mammal from the Lower Cretaceous of Morocco. It is notable for having been found in a unique marine location, and the shape of its teeth suggests an unusual, potentially fish-eating ecologi ...
'' and ''
Jugulator ''Jugulator'' is the thirteenth studio album by English heavy metal band Judas Priest. It was released in Japan on 16 October 1997 and the rest of the world on 28 October 1997. It was their first studio album since ''Painkiller'' in 1990 an ...
'' (and, possibly by association, ''Triconolestes'') in a more basal position among triconodontids, away from alticonodontines.Thomas Martin, Jesús Marugán-Lobón, Romain Vullo, Hugo Martín-Abad, Zhe-Xi Luo & Angela D. Buscalioni (2015). A Cretaceous eutriconodont and integument evolution in early mammals. Nature 526, 380–384.


References

*


External links


Online article from ''Nature'', with illustrationBBC News: Earliest flying mammal discovered
* ttps://news.yahoo.com/s/ap/20061213/ap_on_sc/gliding_mammal AP: Mammals may have flown before birds {{Taxonbar, from=Q132474 Eutriconodonts Jurassic mammals of Asia Gliding animals Fossil taxa described in 2006 Taxa named by Jin Meng Taxa named by Yaoming Hu Taxa named by Yuanqing Wang Taxa named by Xiaolin Wang (paleontologist) Taxa named by Chuankui Li Prehistoric mammal genera