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Gobiconodonta is an order of extinct
mammals 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 ...
known from the
Early Jurassic The Early Jurassic Epoch (geology), Epoch (in chronostratigraphy corresponding to the Lower Jurassic series (stratigraphy), Series) is the earliest of three epochs of the Jurassic Period. The Early Jurassic starts immediately after the Triassic-J ...
(such as '' Huasteconodon'') to early
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', the ...
. They are generally held to be part of
Eutriconodonta Eutriconodonta is an order of early mammals. Eutriconodonts existed in Asia, Africa, Europe, North and South America during the Jurassic and the Cretaceous periods. The order was named by Kermack ''et al.'' in 1973 as a replacement name for the ...
.


Biology

Like many other non-
theria Theria (; Greek: , wild beast) is a subclass of mammals amongst the Theriiformes. Theria includes the eutherians (including the placental mammals) and the metatherians (including the marsupials) but excludes the egg-laying monotremes. Ch ...
n mammals, gobiconodonts retained classical mammalian synapomorphies like
epipubic Epipubic bones are a pair of bones projecting forward from the pelvic bones of modern marsupials, monotremes and fossil mammals like multituberculates, and even basal eutherians (the ancestors of placental mammals, who lack them). They first occ ...
bones (and likely the associated reproductive constrictions), venomous spurs and sprawling limbs. However, the forelimb and shoulder anatomy of at least some species like ''
Jeholodens ''Jeholodens'' was a primitive mammal belonging to the order Eutriconodonta, and which lived in present-day China during the Middle Cretaceous about 125 million years ago. Only one specimen has been formally described. This specimen (the holotyp ...
'' are similar to those of therian mammals, though the hindlimbs remain more conservative. Gobiconodonts had a modern ear anatomy, the main difference from
theria Theria (; Greek: , wild beast) is a subclass of mammals amongst the Theriiformes. Theria includes the eutherians (including the placental mammals) and the metatherians (including the marsupials) but excludes the egg-laying monotremes. Ch ...
ns being that the ear ossicles were still somewhat connected to the jaw via the
Meckel's cartilage In humans, the cartilaginous bar of the mandibular arch is formed by what are known as Meckel's cartilages (right and left) also known as Meckelian cartilages; above this the incus and malleus are developed. Meckel's cartilage arises from the first ...
. Some information on gobiconodont soft-tissues can be seen in ''
Spinolestes ''Spinolestes'' is an extinct mammal genus from the Early Cretaceous of Spain. A gobiconodontid eutriconodont, it is notable for the remarkable degree of preservation, offering profound insights to the biology of non-therian mammals.Thomas Marti ...
'', which was very well preserved, showing evidence of fur and internal organs. ''Spinolestes'' shows hair similar to that of modern mammals, with compound
hair follicle The hair follicle is an organ found in mammalian skin. It resides in the dermal layer of the skin and is made up of 20 different cell types, each with distinct functions. The hair follicle regulates hair growth via a complex interaction between h ...
s with primary and secondary hair, even preserving traces of a pore infection. It also possesses a clear
thoracic diaphragm The thoracic diaphragm, or simply the diaphragm ( grc, διάφραγμα, diáphragma, partition), is a sheet of internal Skeletal striated muscle, skeletal muscle in humans and other mammals that extends across the bottom of the thoracic cavit ...
like modern mammals, as well as spines, dermal
scutes A scute or scutum (Latin: ''scutum''; plural: ''scuta'' "shield") is a bony external plate or scale overlaid with horn, as on the shell of a turtle, the skin of crocodilians, and the feet of birds. The term is also used to describe the anterior po ...
and an ossified
Meckel's cartilage In humans, the cartilaginous bar of the mandibular arch is formed by what are known as Meckel's cartilages (right and left) also known as Meckelian cartilages; above this the incus and malleus are developed. Meckel's cartilage arises from the first ...
. The gobiconodont triconodont dentition has no analogue among living mammals, so comparisons are difficult. However, its clear that most if not all gobiconodonts were primarily carnivorous, given the presence of long, sharp canines and fang-like incisors, premolars with trenchant main cusps that were well suited to grasp and pierce prey, strong development of the mandibular abductor musculature, bone crushing ability in at least some species and several other features. Gobiconodonts are often among the largest mammals in Mesozoic faunal assemblages, displaying a broad size range from small
shrew Shrews (family Soricidae) are small mole-like mammals classified in the order Eulipotyphla. True shrews are not to be confused with treeshrews, otter shrews, elephant shrews, West Indies shrews, or marsupial shrews, which belong to different fa ...
-like insectivores to large forms like ''
Repenomamus ''Repenomamus'' (Latin: "reptile" (reptilis), "mammal" (mammalis)) is a genus of opossum- to badger-sized gobiconodontid mammal containing two species, ''Repenomamus robustus'' and ''Repenomamus giganticus''. Both species are known from fossils ...
'' and ''
Gobiconodon ''Gobiconodon'' is an extinct genus of carnivorous mammal from the Middle Jurassic to Early Cretaceous. It weighed , had a skull of in length, and had in presacral body length. It was one of the largest mammals known from the Mesozoic. Like o ...
''. They were among the first mammals to be specialised for vertebrate prey, and likely occupied the highest
trophic Trophic, from Ancient Greek τροφικός (''trophikos'') "pertaining to food or nourishment", may refer to: * Trophic cascade * Trophic coherence * Trophic egg * Trophic function * Trophic hormone * Trophic level index * Trophic level * Trop ...
levels among mammals in their faunal communities. Several forms like ''Gobiconodon'' and ''Repenomamus'' show evidence of scavenging, being among the few Mesozoic mammals to have significantly exploited that. At least in carnivorous niches, gobiconodonts were probably replaced by
deltatheroidean Deltatheroida is an extinct group of basal metatherians that were distantly related to modern marsupials. The majority of known members of the group lived in the Cretaceous; one species, '' Gurbanodelta kara'', is known from the late Paleocene ( ...
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 wel ...
ns, which are the dominant carnivorous mammals in
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', the ...
faunal assemblages. Competition between both groups is unattested, but in Asia the Early Cretaceous gobiconodontid diversity is replaced entirely by a deltatheroidean one, while in North America ''
Nanocuris ''Nanocuris'' is an extinct genus of Deltatheridiidae from the Cretaceous of Canada (Saskatchewan) and United States (Wyoming - Lance Formation).R. C. Fox, C. S. Scott, and H. N. Bryant. 2007. A new, unusual therian mammal from the Upper Cretac ...
'' appears after the absence of ''
Gobiconodon ''Gobiconodon'' is an extinct genus of carnivorous mammal from the Middle Jurassic to Early Cretaceous. It weighed , had a skull of in length, and had in presacral body length. It was one of the largest mammals known from the Mesozoic. Like o ...
'' and other larger eutriconodonts. Given that all insectivorous and carnivorous mammals groups suffered heavy losses during the mid-Cretaceous, it seems likely these metatherians simply occupied niches left after the extinction of eutriconodonts. At least ''Spinolestes'' had xenarthrous vertebrae and osseous scutes, convergent to those of modern
xenarthrans Xenarthra (; from Ancient Greek wikt:ξένος, ξένος, xénos, "foreign, alien" + wikt:ἄρθρον, ἄρθρον, árthron, "joint") is a major clade of Placentalia, placental mammals native to the Americas. There are 31 living species: ...
and to a lesser extent the
hero shrew The hero shrew (''Scutisorex somereni''), also known as the armored shrew, is a large shrew native to the Congo Basin of Africa. Its features are typical of a white-toothed shrew − short legs, slender snout, dense fur − except for a highly u ...
. Uniquely among crown-group mammals, gobiconodontids replaced their molariform teeth by successors of similar complexity, while in other mammals less complex replacements are the norm.


Classification

Cladogram after Averianov & Lopatin, 2011: Gobiconodonta was named by Averianov & Lopatin (2011); according to the authors it contained two families of mammals (gobiconodontids and klameliids) that were traditionally assigned to
Eutriconodonta Eutriconodonta is an order of early mammals. Eutriconodonts existed in Asia, Africa, Europe, North and South America during the Jurassic and the Cretaceous periods. The order was named by Kermack ''et al.'' in 1973 as a replacement name for the ...
. The exact phylogenetic position of these families within
Mammaliaformes Mammaliaformes ("mammalian forms") is a clade that contains the crown group mammals and their closest Extinction, extinct relatives; the group adaptive radiation, radiated from earlier probainognathian cynodonts. It is defined as the clade origin ...
is uncertain. Some analyses using only dental and
mandibular In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
characters (the first analysis of Gao ''et al.'', 2010, Meng, Wang & Li, 2011, Averianov & Lopatin, 2011) found that gobiconodonts and eutriconodonts did not form a clade that wouldn't also include
trechnotheria Trechnotheria is a group of mammals that includes the therians and some fossil mammals from the Mesozoic Era. In the Jurassic through Cretaceous periods, the group was endemic to what would be Asia and Africa. Trechnotheria has been assigned va ...
ns; however, some other analyses of dental and mandibular characters (Gaetano and Rougier, 2011, 2012) did recover gobiconodontids as eutriconodonts. The analysis conducted by Luo ''et al.'' (2007) and the second analysis of Gao ''et al.'' (2010), involving a broader range of Mesozoic mammaliaforms and more characters (including postcranial ones) recovered gobiconodontids as eutriconodonts as well. However, Gao ''et al.'' (2010) stressed that jeholodentids and gobiconodontids are the only eutriconodonts with known postcranial skeletons; according to the authors, it remains uncertain whether the results of their second analysis represent true phylogeny or are merely "a by-product of
long branch attraction In phylogenetics, long branch attraction (LBA) is a form of systematic error whereby distantly related lineages are incorrectly inferred to be closely related. LBA arises when the amount of molecular or morphological change accumulated within a lin ...
of jeholodentids and gobiconodontids". Most recent analysis recover them as eutriconodonts, closer to
triconodontid 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 Cretaceo ...
s and their immediate outgroups than to amphilestids:


Notes


References

{{Taxonbar, from=Q5575496 Eutriconodonts Taxa named by Alexander O. Averianov