Docofossor Temporal range: | |
---|---|
Life restoration of D. brachydactylus | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Synapsida |
Clade: | Therapsida |
Clade: | Cynodontia |
Clade: | Mammaliaformes |
Order: | † Docodonta |
Family: | † Docodontidae |
Genus: | † Docofossor Luo et al., 2015 |
Species: | †D. brachydactylus |
Binomial name | |
†Docofossor brachydactylus Luo et al., 2015 | |
Docofossor is an extinct mammaliaform (a docodont) from the Jurassic period. Its remains have been recovered in China from 160 million years old rocks. It appears to have been the earliest-known subterranean mammaliaform, with adaptations remarkably similar to the modern Chrysochloridae, the golden moles. [1]
The fossil of Docofossor brachydactylus, holotype BMNH 131735, along with that of Agilodocodon scansorius , was originally found by farmers near Nanshimen in the province of Hebei in a layer of the Chinese Tiaojishan Formation (Oxfordian) and acquired by the Beijing Museum of Natural History. The holotype consists of a compressed skeleton with skull and lower jaws, preserved on a plate and counterplate, along with soft-tissue remnants. The shoulder girdle area and the tail have been damaged. The type species Docofossor brachydactylus was named and described by Zhe-Xi Luo, Meng Qingjin, Ji Qiang, Liu Di, Zhang Yuguang, and April I. Neander in the journal Science in 2015. The generic name refers to the membership of the Docodonta and a burrowing lifestyle, fossor meaning "digger" in Latin. The specific name is derived from Greek βραχύς, "short", and δάκτυλος, "finger", referring to the reduction of the finger phalanges. [2]
Docofossor was at least nine centimetres long, exempting the tail, and weighed at least nine grams, perhaps sixteen. It had a skeletal structure and body proportions strikingly similar to the modern day African golden mole. It had shovel-like fingers for digging, short and wide upper molars typical of mammals that forage underground, and a sprawling posture indicative of subterranean movement. The sprawling is proven by a short hindlimb of just twenty-three millimetres, a massive olecranon as an adaptation for digging and a projecting parafibula forcing the knee joint into a bent position. Its snout point was blunt and slightly overhanging.
Docofossor shows two unique derived traits or autapomorphies. The upper molars have a grooved (prestylar) shelf at the front. The fourth molar has a single root.
Docofossor had a reduced number of phalanges in its fingers. The phalangeal formula was 2-2-2-2-2 instead of the ancestral 2-3-3-3-3. This led to shortened but wide digits. Furthermore, the claw-bearing sections were enlarged and the upper phalanges shortened. African golden moles possess almost exactly the same adaptation, which provides an evolutionary advantage for digging mammals. This characteristic is due to the fusion of the bone joints between the upper and middle phalanges during embryonic development – a process influenced by the genes BMP and GDF-5. Because of the many anatomical similarities, the researchers hypothesize that this genetic mechanism may have played a comparable role in early mammaliaform evolution, as in the case of Docofossor. [3]
The spines and ribs of Docofossor also show evidence for the influence of genes seen in modern mammals, since they feature a gradual thoracic to lumbular vertebrae transition. These shifting patterns of thoracic-lumbular transition have been seen in modern mammals and are known to be regulated by the genes Hox 9-10 and Myf 5-6. That these ancient mammaliaforms had similar developmental patterns is evidence that these gene networks could have functioned in a similar way long before true mammals evolved.
Docofossor was placed in the Docodontidae together with Docodon and Haldanodon . Docodontidae are basal Mammaliaformes outside of the crown group Mammalia.
Early mammaliaforms were once thought to have limited ecological opportunities to diversify during the dinosaur-dominated Mesozoic era. However, Docofossor and numerous other fossils – including Castorocauda , a (related) swimming, fish-eating mammaliaform – provide strong evidence that forms ancestral to the true mammals adapted to wide-ranging environments despite competition from dinosaurs. [4]
Zhangheotherium is an extinct genus of "symmetrodont" mammal from the Early Cretaceous of China. A single species is known, Zhangheotherium quinquecuspidens from Jianshangou Beds of the Yixian Formation. Zhangheotherium was the first "symmetrodont" known from a nearly complete skeleton, expanding knowledge of the group beyond isolated teeth and jaws. The genus name honors Zhang He, who collected the holotype fossil from Liaoning Province prior to its 1997 description. The specific name is Latin for "five-cusped teeth".
Akidolestes is an extinct genus of mammals of the family Spalacotheriidae, a group of mammals related to therians.
Castorocauda is an extinct, semi-aquatic, superficially otter-like genus of docodont mammaliaforms with one species, C. lutrasimilis. It is part of the Yanliao Biota, found in the Daohugou Beds of Inner Mongolia, China dating to the Middle to Late Jurassic. It was part of an explosive Middle Jurassic radiation of Mammaliaformes moving into diverse habitats and niches. Its discovery in 2006, along with the discovery of other unusual mammaliaforms, disproves the previous hypothesis of Mammaliaformes remaining evolutionarily stagnant until the extinction of the non-avian dinosaurs.
Docodonta is an order of extinct Mesozoic mammaliaforms. They were among the most common mammaliaforms of their time, persisting from the Middle Jurassic to the Early Cretaceous across the continent of Laurasia. They are distinguished from other early mammaliaforms by their relatively complex molar teeth. Docodont teeth have been described as "pseudotribosphenic": a cusp on the inner half of the upper molar grinds into a basin on the front half of the lower molar, like a mortar-and-pestle. This is a case of convergent evolution with the tribosphenic teeth of therian mammals. There is much uncertainty for how docodont teeth developed from their simpler ancestors. Their closest relatives may have been certain Triassic "symmetrodonts", namely Woutersia, Delsatia.
Mammaliaformes is a clade that contains the crown group mammals and their closest extinct relatives; the group radiated from earlier probainognathian cynodonts. It is defined as the clade originating from the most recent common ancestor of Morganucodonta and the crown group mammals; the latter is the clade originating with the most recent common ancestor of extant Monotremata, Marsupialia, and Placentalia. Besides Morganucodonta and the crown group mammals, Mammaliaformes includes Docodonta and Hadrocodium.
Volaticotherium antiquum is an extinct, gliding, insectivorous mammal that lived in Asia during the Jurassic period, around 164 mya. It is the only member of the genus Volaticotherium.
Volaticotherini is a clade of eutriconodont mammals from the Mesozoic. In addition to the type genus Volaticotherium, it includes the genera Argentoconodon, Ichthyoconodon, and potentially Triconolestes.
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 paraphyletic Triconodonta.
Yanoconodon is a monotypic genus of extinct early mammal whose representative species Yanoconodon allini lived during the Mesozoic in what is now China. The holotype fossil of Yanoconodon was excavated in the Yan Mountains about 300 kilometres from Beijing in the Qiaotou member of the Huajiying Formation of Hebei Province, China, and is therefore of uncertain age. The Qiaotou Member may correlate with the more well-known Early Cretaceous Yixian Formation, and so probably dates to around 122 Ma ago.
Haramiyida is a possibly polyphyletic order of mammaliaform cynodonts or mammals of controversial taxonomic affinites. 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. Some authors have placed them in a clade with Multituberculata dubbed Allotheria within Mammalia. Other studies have disputed this and suggested the Haramiyida were not crown mammals, but were part of an earlier offshoot of mammaliaformes instead. 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.
Shuotherium is a fossil mammaliaform known from Middle-Late Jurassic of the Forest Marble Formation of England, and the Shaximiao Formation of Sichuan, China.
Juramaia is an extinct genus of a therian mammal, possibly a very basal eutherian mammal, known from the Late Jurassic or Early Cretaceous deposits of western Liaoning, China. It is a small shrew-like mammal weighing around 15–17 grams (0.53–0.60 oz).
Haldanodon is an extinct docodont mammaliaform which lived in the Upper Jurassic. Its fossil remains have been found in Portugal, in the well-known fossil locality of Guimarota, which is in the Alcobaça Formation. It may have been a semi-aquatic burrowing insectivore, similar in habits to desmans and the platypus. Several specimens are known, include a partial skeleton and well-preserved skulls.
Shenshou is a genus of haramiyidan dating from the Oxfordian stage of the Late Jurassic, approximately 160 million years ago. Fossils were recovered from the Tiaojishan Formation in the Liaoning province of China.
Xianshou is a genus of gliding haramiyidan synapsid known from the Oxfordian stage of the Jurassic period, approximately 160 million years ago. Two species, X. linglong and X. songae, are known from fossils of the Tiaojishan Formation in the Liaoning province of China.
Agilodocodon was a genus of shrew-sized docodont from the Middle Jurassic, believed to be the earliest known tree-climbing mammaliaform. It contains one species, A. scansorius.
Vilevolodon is an extinct, monotypic genus of volant, arboreal euharamiyids from the Oxfordian age of the Late Jurassic of China. The type species is Vilevolodon diplomylos. The genus name Vilevolodon references its gliding capabilities, Vilevol, while don is a common suffix for mammalian taxon titles. The species name diplomylos refers to the dual mortar-and-pestle occlusion of upper and lower molars observed in the holotype; diplo, mylos.
Maiopatagium is an extinct genus of gliding euharamiyids which existed in Asia during the Jurassic period. It possessed a patagium between its limbs and presumably had similar lifestyle to living flying squirrels and colugos. The type species is Maiopatagium furculiferum, which was described from the Tiaojishan Formation by Zhe-Xi Luo in 2017; it lived in what is now the Liaoning region of China during the late Jurassic .Maiopatagium and Vilevolodon, described concurrently, offer clues to the ways various synapsids have taken to the skies over evolutionary time scales. A second species, M. sibiricum, was described from the Bathonian aged Itat Formation in western Siberia, Russia in 2019
Microdocodon is a genus of docodontan mammaliaform from the Late Jurassic Yanliao Biota. It contains only a single species, Microdocodon gracilis, known from the Daohugou locality. It is unique for preserving the hyoid bone, which is almost unknown in the early mammal fossil record.
Zhe-Xi Luo is an American paleontologist of Chinese origin, specializing in vertebrate paleontology, particularly mammal evolution, morphology, and systematics.