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''Taowu'' is an extinct
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
of
machairodont Machairodontinae is an extinct subfamily of carnivoran mammals of the family Felidae (true cats). They were found in Asia, Africa, North America, South America, and Europe from the Miocene to the Pleistocene, living from about 16 million until ...
s, a type of saber-toothed cat. It lived during the Early Pleistocene about 2.5 million years ago in
East Asia East Asia is the eastern region of Asia, which is defined in both Geography, geographical and culture, ethno-cultural terms. The modern State (polity), states of East Asia include China, Japan, Mongolia, North Korea, South Korea, and Taiwan. ...
. So far, only one skull is known, found in northern China. Based on this, a relatively small representative of the saber-toothed cats can be reconstructed, which only reached the size of a present-day leopard. In its dentition characteristics, it mediates between
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
older forms such as ''
Amphimachairodus ''Amphimachairodus'' is an extinct genus of large machairodonts. It is also a member of the tribe Homotherini within Machairodontinae and is most closely related to such species as '' Xenosmilus'', ''Homotherium'' itself, and ''Nimravides''. It ...
'' and younger members such as ''
Homotherium ''Homotherium'', also known as the scimitar-toothed cat or scimitar cat, is an extinct genus of machairodontine saber-toothed predator, often termed scimitar-toothed cats, that inhabited North America, South America, Eurasia, and Africa during ...
''. The genus was scientifically described in 2022, but the find material was recovered as early as the 1930s.


Description

''Taowu'' was a rather small representative of the saber-toothed cats. It reached about the size of a present-day leopards. Found so far is a nearly complete skull of 23.5 cm length and 13.4 cm width, measured across the zygomatic arches. Only the articular surface on the occipital bone is missing from this one. In top view, the skull was relatively slender, comparatively narrower than in ''
Homotherium ''Homotherium'', also known as the scimitar-toothed cat or scimitar cat, is an extinct genus of machairodontine saber-toothed predator, often termed scimitar-toothed cats, that inhabited North America, South America, Eurasia, and Africa during ...
'' or ''
Xenosmilus ''Xenosmilus hodsonae'' (from Greek, , ''xenos'', "strange" + , ''smilē'', " chisel" ) is an extinct species of the Machairodontinae, or saber-toothed cats. Description The species name ''hodsonae'' originates from Debra Hodson, the wife of ...
'', and possessed a rounded snout region. The snout did not taper behind the canine, but continuously increased in width posteriorly. Likewise, a cranial narrowing behind the
orbits In celestial mechanics, an orbit is the curved trajectory of an object such as the trajectory of a planet around a star, or of a natural satellite around a planet, or of an artificial satellite around an object or position in space such as a p ...
was only rather weakly pronounced. In lateral view, the frontal line showed a gentle uparching, and the nasal opening was recessed and concavely indented relative to the
premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammal has ...
. The
incisor Incisors (from Latin ''incidere'', "to cut") are the front teeth present in most mammals. They are located in the premaxilla above and on the mandible below. Humans have a total of eight (two on each side, top and bottom). Opossums have 18, wher ...
row thus appeared much more prominent than in, for example, ''
Amphimachairodus ''Amphimachairodus'' is an extinct genus of large machairodonts. It is also a member of the tribe Homotherini within Machairodontinae and is most closely related to such species as '' Xenosmilus'', ''Homotherium'' itself, and ''Nimravides''. It ...
''. In plan view, the frontal area was broad. The
nasal bones The nasal bones are two small oblong bones, varying in size and form in different individuals; they are placed side by side at the middle and upper part of the face and by their junction, form the bridge of the upper one third of the nose. Eac ...
widened posteriorly. The
infraorbital foramen In human anatomy, the infraorbital foramen is one of two small holes in the skull's upper jawbone (maxillary bone), located below the eye socket and to the left and right of the nose. Both holes are used for blood vessels and nerves. In anatomica ...
was at the level of the fourth premolar, as was the anterior edge of the orbit. Bony ridges attached to the
frontal bone The frontal bone is a bone in the human skull. The bone consists of two portions.'' Gray's Anatomy'' (1918) These are the vertically oriented squamous part, and the horizontally oriented orbital part, making up the bony part of the forehead, pa ...
, which united to form a parietal crest. The zygomatic arch bulged strongly dorsally. On the underside of the skull, the tympanic bulla was markedly distended. As a diagnostic feature, the glenoid fossa was on a raised platform, more prominent than in ''Amphimachairodus'', and was very wide. The anterior palatal window was at the level of the canine, and the largest posterior one was at the level of the third premolar. The upper dentition consisted of three
incisors Incisors (from Latin ''incidere'', "to cut") are the front teeth present in most mammals. They are located in the premaxilla above and on the mandible below. Humans have a total of eight (two on each side, top and bottom). Opossums have 18, wher ...
, one canine, two premolars, and two
molars The molars or molar teeth are large, flat teeth at the back of the mouth. They are more developed in mammals. They are used primarily to grind food during chewing. The name ''molar'' derives from Latin, ''molaris dens'', meaning "millstone to ...
per half of the jaw. The incisors formed a closed arch and increased in size from the inside to the outside. An additional cusp existed on the third incisor. The canine tooth protruded up to 7.4 cm from the jaw and was flat blade-shaped in cross-section. Both the anterior and posterior edges showed fine serration. Adjacent to the canine, a
diastema A diastema (plural diastemata, from Greek διάστημα, space) is a space or gap between two teeth. Many species of mammals have diastemata as a normal feature, most commonly between the incisors and molars. More colloquially, the condition ...
separated the anterior from posterior dentition. Unlike some other saber-toothed cats such as '' Lokotunjailurus'', the second premolar was absent. The third and thus most anterior premolar was not reduced in size, which differs from ''
Homotherium ''Homotherium'', also known as the scimitar-toothed cat or scimitar cat, is an extinct genus of machairodontine saber-toothed predator, often termed scimitar-toothed cats, that inhabited North America, South America, Eurasia, and Africa during ...
'', '' Lokotunjailurus'', or ''
Xenosmilus ''Xenosmilus hodsonae'' (from Greek, , ''xenos'', "strange" + , ''smilē'', " chisel" ) is an extinct species of the Machairodontinae, or saber-toothed cats. Description The species name ''hodsonae'' originates from Debra Hodson, the wife of ...
''. Overall, the tooth appeared elongated at a length of just under 2 cm, as did the much larger posterior premolar, which grew to a good 3.4 cm in length. The typical pointed cusps, the para-, meta-, and protoconus, existed on this, the latter only being small in formation. The anterior shear edge, the parastyle, had an elongate shape, from which the anteriormost section, the preparastyle, stood out only imperceptibly.


Fossil finds

The only fossil find of ''Taowu'' to date comes from Fancun, a locality about 36 km east of the township of Taigu, which in turn is part of the city of
Jinzhong Jinzhong, formerly Yuci, is a prefecture-level city in east central Shanxi province of the People's Republic of China, bordering Hebei province to the east. As of the 2020 census, its total population was 3,379,498 inhabitants whom 1,226,617 ...
in the
Chinese Chinese can refer to: * Something related to China * Chinese people, people of Chinese nationality, citizenship, and/or ethnicity **''Zhonghua minzu'', the supra-ethnic concept of the Chinese nation ** List of ethnic groups in China, people of ...
Shanxi province. The site and other areas in the region were excavated as early as the 1930s by American paleontologist Childs Frick. The various localities yielded numerous fossil materials, including several
carnivores A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, fat and other so ...
such as ''
Homotherium ''Homotherium'', also known as the scimitar-toothed cat or scimitar cat, is an extinct genus of machairodontine saber-toothed predator, often termed scimitar-toothed cats, that inhabited North America, South America, Eurasia, and Africa during ...
'' as another representative of the saber-toothed cats, an extinct form of the lynx as well as the
badger Badgers are short-legged omnivores in the family Mustelidae (which also includes the otters, wolverines, martens, minks, polecats, weasels, and ferrets). Badgers are a polyphyletic rather than a natural taxonomic grouping, being united by ...
, ''
Xenocyon ''Xenocyon'' ("strange dog") is an extinct subgenus of ''Canis''. The group includes ''Canis'' (''Xenocyon'') ''africanus'', ''Canis'' (''Xenocyon'') ''antonii'' and ''Canis'' (''Xenocyon'') ''falconeri'' that gave rise to ''Canis'' (''Xenocyon'' ...
'' as a member of the dogs, and the hyena ''
Pachycrocuta ''Pachycrocuta'' is an extinct genus of prehistoric hyenas. The largest and most well-researched species is ''Pachycrocuta brevirostris'', colloquially known as the giant short-faced hyena as it stood about at the shoulder and it is estimated t ...
''. In addition, early relatives of modern horses also appeared. Especially the latter mentioned imply a predominant position in the
Pleistocene The Pleistocene ( , often referred to as the ''Ice age'') is the geological epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was finally confirmed in ...
, with the lower section being more likely.Lorenzo Rook: ''The Plio-Pleistocene Old World Canis (Xenocyon) ex gr. falconeri.'' Bollettino della Societa Paleontologica Italiana 33 (1), 1994, S. 71–82Véra Eisenmann und Tao Deng: ''Equus qingyangensis (Equidae, Perissodactyla) of the Upper Pliocene of Bajiazui, China: Evidence for the North American origin of an Old World lineage distinct from E. stenonis.'' Quaternaire 2 (2), 2005, S. 113–122


Classifications

''Taowu'' is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
from the extinct subfamily of saber-toothed cats (Machairodontinae) within the family of cats (Felidae). The saber-toothed cats represent an ancient branch of cats, which according to
molecular genetics Molecular genetics is a sub-field of biology that addresses how differences in the structures or expression of DNA molecules manifests as variation among organisms. Molecular genetics often applies an "investigative approach" to determine the ...
studies, forms the sister group of today's
big cats The term "big cat" is typically used to refer to any of the five living members of the genus ''Panthera'', namely the tiger, lion, jaguar, leopard, and snow leopard. Despite enormous differences in size, various cat species are quite similar ...
and small cats. Their separation from the common branch occurred about 20 million years ago in the Lower
Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...
.Johanna L. A. Paijmans, Ross Barnett, M. Thomas P. Gilbert, M. Lisandra Zepeta-Mendoza, Jelle W. F. Reumer, John de Voss, Grant Zazula, Doris Nagel, Gennady F. Baryshnikov, Jennifer A. Leonard, Nadine Rohland, Michael V. Westbury, Axel Barlow, Michael Hofreiter: ''Evolutionary history of sabre-toothed cats based on ancient mitogenomics.'' Current Biology 27, 2017, S. 3330–3336, doi:10.1016/j.cub.2017.09.033Michael V. Westbury, Ross Barnett, Marcela Sandoval-Velasco, Graham Gower, Filipe Garrett Vieira, Marc de Manuel, Anders J. Hansen, Nobuyuki Yamaguchi, Lars Werdelin, Tomas Marques-Bonet, M. Thomas P Gilbert, Eline D Lorenzen: ''A genomic exploration of the early evolution of extant cats and their sabre-toothed relatives.'' Open Research Europe 1, 2021, S. 25, doi:10.12688/openreseurope.13104.2Alexandre Hassanin, Géraldine Veron, Anne Ropiquet, Bettine Jansen van Vuuren, Alexis Lécu, Steven M. Goodman, Jibran Haider, Trung Thanh Nguyen: ''Evolutionary history of Carnivora (Mammalia, Laurasiatheria) inferred from mitochondrial genomes.'' PLOS One 16 (3), 2021, S. e0249387, doi:10.1371/journal.pone.0240770 The feature that gave saber-toothed cats their name is their significantly enlarged
canine teeth In mammalian oral anatomy, the canine teeth, also called cuspids, dog teeth, or (in the context of the upper jaw) fangs, eye teeth, vampire teeth, or vampire fangs, are the relatively long, pointed teeth. They can appear more flattened howeve ...
. However, this shows certain variations within the kinship community, which led to the establishment of different subgroups. Here, the Machairodontini have saber-like canines, the
Smilodontini Smilodontini is an extinct tribe within the Machairodontinae or "saber-toothed cat" subfamily of the Felidae. The tribe is also known as the "dirk-toothed cats". They were endemic to South America, North America, Europe, Asia, and Africa during t ...
have dagger-like canines, and the Metailurini have less conspicuously large canines.Paolo Piras, Daniele Silvestro, Francesco Carotenuto, Silvia Castiglione, Anastassios Kotsakis, Leonardo Maiorino, Marina Melchionna, Alessandro Mondanaro, Gabriele Sansalone, Carmela Serio, Veronica Anna Vero, Pasquale Raia: ''Evolution of the sabertooth mandible: A deadly ecomorphological specialization.'' Palaeogeography, Palaeoclimatology, Palaeoecology 496, 2018, S. 166–174, doi:10.1016/j.palaeo.2018.01.034 With its leopard-like body size, ''Taowu'' represents one of the smallest representatives of the saber-toothed cats of the
Pliocene The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58Pleistocene The Pleistocene ( , often referred to as the ''Ice age'') is the geological epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was finally confirmed in ...
. In overall dentition, the form can be assumed to be somewhat intermediate between the more primitive ''
Amphimachairodus ''Amphimachairodus'' is an extinct genus of large machairodonts. It is also a member of the tribe Homotherini within Machairodontinae and is most closely related to such species as '' Xenosmilus'', ''Homotherium'' itself, and ''Nimravides''. It ...
'' and the terminal form ''
Homotherium ''Homotherium'', also known as the scimitar-toothed cat or scimitar cat, is an extinct genus of machairodontine saber-toothed predator, often termed scimitar-toothed cats, that inhabited North America, South America, Eurasia, and Africa during ...
'' within the Machairodontini. This is recognizable, for example, from the third premolars, which are only slightly reduced in size. On the other hand, the anteriorly shifted
incisors Incisors (from Latin ''incidere'', "to cut") are the front teeth present in most mammals. They are located in the premaxilla above and on the mandible below. Humans have a total of eight (two on each side, top and bottom). Opossums have 18, wher ...
also suggest certain more modern characteristics.Paolo Piras, Daniele Silvestro, Francesco Carotenuto, Silvia Castiglione, Anastassios Kotsakis, Leonardo Maiorino, Marina Melchionna, Alessandro Mondanaro, Gabriele Sansalone, Carmela Serio, Veronica Anna Vero, Pasquale Raia: ''Evolution of the sabertooth mandible: A deadly ecomorphological specialization.'' Palaeogeography, Palaeoclimatology, Palaeoecology 496, 2018, S. 166–174, doi:10.1016/j.palaeo.2018.01.034 The skull underlying the genus ''Taowu'' was found as early as the 1930s in Fancun, part of the city of
Jinzhong Jinzhong, formerly Yuci, is a prefecture-level city in east central Shanxi province of the People's Republic of China, bordering Hebei province to the east. As of the 2020 census, its total population was 3,379,498 inhabitants whom 1,226,617 ...
, in the
Chinese Chinese can refer to: * Something related to China * Chinese people, people of Chinese nationality, citizenship, and/or ethnicity **''Zhonghua minzu'', the supra-ethnic concept of the Chinese nation ** List of ethnic groups in China, people of ...
Shanxi province. It was mentioned in individual publications in which various researchers classified it as belonging to ''
Machairodus ''Machairodus'' (from el, μαχαίρα , 'knife' and el, ὀδούς 'tooth') is a genus of large machairodontine saber-toothed cats that lived in Africa, Eurasia and North America during the late Miocene. It is the animal from which the su ...
'', but a precise characteristic description was not developed.Germaine Petter und F. Clark Howell: ''Machairodus africanus Arambourg, 1970 (Carnivora Mammalia) du Villafranchien d'Aïn Brimba, Tunisie.'' Bulletin du Muséum National d'Histoire Naturelle 4 (9), 1987, S. 97–119

Denis Geraads, Tanju Kaya und Vahdet Tuna: ''A skull of Machairodus giganteus (Felidae, Mammalia) from the Late Miocene of Turkey.'' Neues Jahrbuch für Geologie und Paläontologie Monatshefte 2004 (2), 2004, S. 95–110 Studies in 2022 necessitated a reevaluation. Jiangzuo Qigao and colleagues then established ''Taowu'' as a distinct form of saber-toothed cat in their scientific first description and presented the skull in detail. The generic name ''Taowu'' refers to the creature of the same name in
Chinese mythology Chinese mythology () is mythology that has been passed down in oral form or recorded in literature in the geographic area now known as Greater China. Chinese mythology includes many varied myths from regional and cultural traditions. Much of t ...
. Its striking characteristics include powerful tusks reminiscent of saber teeth. The species ''T. liui'', introduced together with the genus, uses its epithet to honor the Chinese scientist Liu Jinyi, who had made significant contributions to the study of
Pleistocene The Pleistocene ( , often referred to as the ''Ice age'') is the geological epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was finally confirmed in ...
East Asian predators. Below is a phylogenetic tree of the Machairodontinae, based on the work of Jiangzuo and colleagues (2022).


References

{{Machairodontinae Machairodontinae Fossil taxa described in 2022 Pleistocene mammals of Asia Fossils of China Pleistocene carnivorans