''Yoshi'' is an
extinct
Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
genus
Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
machairodontine
Machairodontinae (from Ancient Greek μάχαιρα ''machaira,'' a type of Ancient Greek sword and ὀδόντος ''odontos'' meaning tooth) is an extinct subfamily of carnivoran mammals of the cat family Felidae, representing the earliest d ...
sabertooth cat
Machairodontinae (from Ancient Greek μάχαιρα '' machaira,'' a type of Ancient Greek sword and ὀδόντος ''odontos'' meaning tooth) is an extinct subfamily of carnivoran mammals of the cat family Felidae, representing the earliest ...
in the tribe
Metailurini
Metailurini is an extinct taxonomic tribe of large saber-toothed cats that lived in Africa, Asia, Europe, and North America from the Miocene to the Pleistocene.
The best known Metalurini genera are '' Dinofelis'' and '' Metailurus''. Metailurini ...
. Its remains were first discovered in
Pikermi, Greece, and were assigned by Reinhold Friedrich Hensel to ''Machairodus'' (under the new species ''M. parvulus''. In 1938, Miklós Kretzoi assigned it to a new genus, ''Pikermia''. Subsequently, it was realised that the Greek taxon was closely related to another from China, then called ''Parapseudailurus minor'', and both were synonymised. In 2014, a new genus, ''Yoshi'', was erected to encompass the two, resurrecting ''Y. minor'' as a valid species. As ''M. parvulus'' was a ''
nomen dubium
In binomial nomenclature, a ''nomen dubium'' (Latin for "doubtful name", plural ''nomina dubia'') is a scientific name that is of unknown or doubtful application.
Zoology
In case of a ''nomen dubium,'' it may be impossible to determine whether a ...
'', a new species, ''Yoshi garevskii'', was designated as the
type
Type may refer to:
Science and technology Computing
* Typing, producing text via a keyboard, typewriter, etc.
* Data type, collection of values used for computations.
* File type
* TYPE (DOS command), a command to display contents of a file.
* ...
. In the intervening years, three more species have been assigned to the genus: ''Y. faie'', ''Y. obscura'', and ''Y. yongdengenesis''.
In some regards, such as the shape of its skull, ''Yoshi'' was
convergent with
cheetahs
The cheetah (''Acinonyx jubatus'') is a large cat and the fastest land animal. It has a tawny to creamy white or pale buff fur that is marked with evenly spaced, solid black spots. The head is small and rounded, with a short snout and blac ...
, and it was originally suggested that it may have been more
cursorial
A cursorial organism is one that is adapted specifically to run. An animal can be considered cursorial if it has the ability to run fast (e.g. cheetah) or if it can keep a constant speed for a long distance (high endurance). "Cursorial" is often ...
(adapted for fast running) than other metailurines. However, the anatomy of its
postcranial skeleton, such as the fact that its hind limbs were longer than its forelimbs, suggest that, while probably somewhat cursorial, it was not to the same extent as cheetahs. ''Yoshi'' instead had several adaptations for leaping, like many modern cats. It appears to have frequented open habitats more often than contemporary machairodonts.
Taxonomy
Early history
In 1862,
Reinhold Friedrich Hensel
Reinhold is a German language, German, male given name, originally composed of two elements. The first is from ''regin'', meaning "the (German)Gods" or as an emphatic prefix (very) and ''wald'' meaning "powerful". The second element having been re ...
described the species ''Machairodus parvulus'' based poorly preserved skulls from
Pikermi, Greece, which he believed came from an animal the size of a
lynx
A lynx ( ; : lynx or lynxes) is any of the four wikt:extant, extant species (the Canada lynx, Iberian lynx, Eurasian lynx and the bobcat) within the medium-sized wild Felidae, cat genus ''Lynx''. The name originated in Middle Engl ...
. In 1938,
Miklós Kretzoi
Miklós Kretzoi (9 February 1907 – 15 March 2005) was a Hungarian people, Hungarian geologist, paleontologist and paleoanthropologist and Széchenyi Prize winner. Exhibition in the Hungarian National Museum, 9 February - 24 May 2004
Kr ...
reassigned ''M. parvulus'' to a new genus, ''Pikermia'', creating the new combination ''Pikermia parvula''. In the same paper, he suggested that "''
Metailurus''" ''minor'', from China, could be reassigned to a genus of its own, ''Parapseudailurus''.
However, in 1951, Erich Thenius suggested that ''P. minor'' represented the same taxon as Hensel's ''M. parvulus'', a conclusion also reached by G. D. Beaumont in 1961, who went as far as assigning a mandible from Iran to the latter.
Consequently, ''P. minor'' was treated as a synonym of ''M. parvulus'' by subsequent authors. Further material from southern/southeastern Europe and Iran was assigned to ''M. parvulus'' over the years, though much of it is incomplete.
Reclassification as a new genus
In 2015, Nikolai Spassov and Denis Geraads published a paper discussing the taxonomy of the aforementioned taxa. They concluded that Hensel's ''Machairodus parvulus'' was a
nomen dubium
In binomial nomenclature, a ''nomen dubium'' (Latin for "doubtful name", plural ''nomina dubia'') is a scientific name that is of unknown or doubtful application.
Zoology
In case of a ''nomen dubium,'' it may be impossible to determine whether a ...
, due to the generally fragmented nature of all material assigned to the species. All of the remains were assigned to a new genus, ''Yoshi'', named after Spassov's pet cat. Two species were assigned: the
type species
In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...
, ''Y. garevskii'', and ''Y. minor'', the taxon discussed by Kretzoi.
In 2022, two new species ''Yoshi faie'' and ''Yoshi yongdengensis'' were proposed based on fossils found in northeastern China.
The next year, a paper discussing the predatory mammals of the Langebaanweg locality in South Africa assigned a fifth species, ''Y. obscura'', to the genus. ''Y. obscura'' had previously been assigned to ''
Adelphailurus'', ''
Felis
''Felis'' is a genus of small and medium-sized cat species native to most of Africa and south of 60° latitude in Europe and Asia to Indochina. The genus includes the domestic cat. The smallest of the seven ''Felis'' species is the black-footed ...
'', ''
Megantereon
''Megantereon'' is an extinct genus of prehistoric machairodontine saber-toothed cat that lived in Eurasia, Africa and possibly North America from the late Pliocene to the Middle Pleistocene. It is a member of the tribe Smilodontini, and close ...
'', and ''
Metailurus''. The authors noted that the very small taxon ''
Tchadailurus adei'' seemed very close to the genus ''Yoshi'', though a synonymy was not proposed in the text.
Below is a cladogram depicting the phylogenetic analysis from that paper:
Description
''Yoshi'' was a fairly large metailurine. In the paper describing it, ''Y. garevskii'' was described as being intermediate in size between a
lynx
A lynx ( ; : lynx or lynxes) is any of the four wikt:extant, extant species (the Canada lynx, Iberian lynx, Eurasian lynx and the bobcat) within the medium-sized wild Felidae, cat genus ''Lynx''. The name originated in Middle Engl ...
and a
cheetah
The cheetah (''Acinonyx jubatus'') is a large Felidae, cat and the Fastest animals, fastest land animal. It has a tawny to creamy white or pale buff fur that is marked with evenly spaced, solid black spots. The head is small and rounded, wit ...
, though it was closer to the latter, with an estimated body mass of .
The largest, ''Y. yongdengensis'', fell within a similar size range.
Though too fragmentary for precise estimates, ''Y. obscura'' is believed to have been the smallest, roughly the size of a female
leopard
The leopard (''Panthera pardus'') is one of the five extant cat species in the genus ''Panthera''. It has a pale yellowish to dark golden fur with dark spots grouped in rosettes. Its body is slender and muscular reaching a length of with a ...
.
Skull and dentition
The skull length of the ''Y. garevskii''
type specimen
In biology, a type is a particular wikt:en:specimen, specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally associated. In other words, a type is an example that serves to ancho ...
, KAR-69, was in length. The skull of ''Y. garevskii'' is short, broad and high, and is morphologically quite similar to that of cheetahs, with a facial region that was similarly long and sloping. However, that of ''Y. minor'' was longer, and appears to have been less sloping. The frontal area in ''Y. garevskii'' bulged outwards, similar to cheetahs and
snow leopards
The snow leopard (''Panthera uncia'') is a species of large cat in the genus ''Panthera'' of the family Felidae. The species is native to the mountain ranges of Central and South Asia. It is listed as Vulnerable on the IUCN Red List because ...
, while that of ''Y. minor'' was somewhat flatter. The two species' skulls also differed in that the muzzle of ''Y. garevskii'' was steeper, and the
orbits
In celestial mechanics, an orbit (also known as orbital revolution) is the curved trajectory of an physical body, object such as the trajectory of a planet around a star, or of a natural satellite around a planet, or of an satellite, artificia ...
(eye sockets) were elevated on the skull. On the
frontal bone
In the human skull, the frontal bone or sincipital bone is an unpaired bone which consists of two portions.'' Gray's Anatomy'' (1918) These are the vertically oriented squamous part, and the horizontally oriented orbital part, making up the bo ...
, the
zygomatic processes are short and rounded, like those of cheetahs. The
zygomatic arches are broad and thick, though differ from modern large felids in that their bases are narrower.
''Yoshi''
's dentition was dissimilar to that of other machairodonts. The upper
canines were short and uncrenulated. The lower canines, too, were small, and hardly surpassed the level of the upper
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 ...
'
alveoli
Alveolus (; pl. alveoli, adj. alveolar) is a general anatomical term for a concave cavity or pit.
Uses in anatomy and zoology
* Pulmonary alveolus, an air sac in the lungs
** Alveolar cell or pneumocyte
** Alveolar duct
** Alveolar macrophage
* M ...
while the jaw was closed. The
diastema
A diastema (: diastemata, from Greek , '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 may be referred to ...
(gap) between the canines and the
cheek teeth
Cheek teeth or postcanines comprise the molar and premolar teeth in mammals. Cheek teeth are multicuspidate (having many folds or tubercles). Mammals have multicuspidate molars (three in placentals, four in marsupials, in each jaw quadrant) and ...
(
premolars
The premolars, also called premolar teeth, or bicuspids, are transitional teeth located between the canine and molar teeth. In humans, there are two premolars per quadrant in the permanent set of teeth, making eight premolars total in the mout ...
and
molars
The molars or molar teeth are large, flat tooth, teeth at the back of the mouth. They are more developed in mammal, mammals. They are used primarily to comminution, grind food during mastication, chewing. The name ''molar'' derives from Latin, '' ...
) was fairly small. The third upper and lower premolars lacked a
mesial
This is a list of definitions of commonly used terms of location and direction in dentistry. This set of terms provides orientation within the oral cavity, much as anatomical terms of location provide orientation throughout the body.
Terms
...
accessory
cuspid. The first upper molar was small and button-shaped,
while the equivalent lower molar bore a distinct
talonid
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 tooth ...
.
Postcranial skeleton
The
postcranial skeleton of ''Yoshi'' is best known from a specimen from Greece, initially assigned to "''Metailurus parvulus''" (''Y. garevskii''). The specimen consists of a mandible, most limb elements, some
cervical (neck) and
thoracic
The thorax (: thoraces or thoraxes) or chest is a part of the anatomy of mammals and other tetrapod animals located between the neck and the abdomen.
In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main ...
(upper back)
vertebrae
Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spinal ...
, and ribs. The
fossa for the
teres major
The teres major muscle is a muscle of the upper limb. It attaches to the scapula and the humerus and is one of the seven scapulohumeral muscles. It is a thick but somewhat flattened muscle.
The teres major muscle (from Latin ''teres'', meanin ...
covers half of the
scapula's posterior (rearward) border, and is well defined by a prominent crest. The head of the
humerus
The humerus (; : humeri) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius (bone), radius and ulna, and consists of three sections. The humeral upper extrem ...
is almost circular in outline. The lateral compression of the humeral
shaft
Shaft may refer to:
Rotating machine elements
* Shaft (mechanical engineering), a rotating machine element used to transmit power
* Line shaft, a power transmission system
* Drive shaft, a shaft for transferring torque
* Axle, a shaft around whi ...
is comparable to that of leopards, being thicker than that of
lions
The lion (''Panthera leo'') is a large cat of the genus ''Panthera'', native to Sub-Saharan Africa and India. It has a muscular, broad-chested body; a short, rounded head; round ears; and a dark, hairy tuft at the tip of its tail. It is se ...
and cheetahs. The crest at the posteromedial (rear middle) margin of the
ulna
The ulna or ulnar bone (: ulnae or ulnas) is a long bone in the forearm stretching from the elbow to the wrist. It is on the same side of the forearm as the little finger, running parallel to the Radius (bone), radius, the forearm's other long ...
was well-developed, providing a strong insertion point for the large head of the
triceps brachii
The triceps, or triceps brachii (Latin for "three-headed muscle of the arm"), is a large muscle on the back of the upper limb of many vertebrates. It consists of three parts: the medial, lateral, and long head. All three heads cross the elbow jo ...
. Unlike other
machairodontines (with the exception of ''
Homotherium
''Homotherium'' is an extinct genus of Homotherini, scimitar-toothed cat belonging to the extinct subfamily Machairodontinae that inhabited North America, Eurasia, and Africa, as well as possibly South America during the Pliocene and Pleistocene ...
''), the trochlear notch lacked a groove on its superomedial (above and at the midline) edge.
Palaeobiology
Initially, based on similarities between ''Yoshi'' and modern cheetahs, it was suggested that the former may have been
cursorial
A cursorial organism is one that is adapted specifically to run. An animal can be considered cursorial if it has the ability to run fast (e.g. cheetah) or if it can keep a constant speed for a long distance (high endurance). "Cursorial" is often ...
to a similar degree to the latter.
However, analysis of the postcranial skeleton indicates that this cursoriality was only moderate: the ovoid shaft cross-section of the
femoral shaft, greater in diameter transversely (across) than
anteroposteriorly (from front to back), which is a trait associated with reduced cursoriality. Furthermore, the shaft bows anteriorly (forwards), and the hind limbs were somewhat longer than the forelimbs, typical of felids with leaping abilities.
Overall, ''Yoshi'' appears to have been intermediate between cursorial and non-cursorial machairodonts,
and was more well-adapted to open environments than larger contemporaries, such as ''
Paramachaerodus
''Paramachaerodus'' is an extinct genus of saber-tooth cat of the subfamily Machairodontinae, which was endemic to Eurasia during the Middle and Late Miocene from 15 to 9 Ma. A 2022 phylogenetic analysis suggested that the genus may be polyphyl ...
''.
Distribution and palaeoenvironment
Specimens assigned to ''Yoshi'' have been recovered from Greece, Spain, Ukraine, China, and Iran, though with the exception of the Greek and Chinese fossils, most are too fragmentary to confidently assign to the genus.
The most complete postcranial skeleton of ''Yoshi'' is known from the K1 locality of northern Greece, which also preserves fossils from the
suid
The Unix and Linux access rights flags setuid and setgid (short for ''set user identity'' and ''set group identity'') allow users to run an executable with the file system permissions of the executable's owner or group respectively and to change ...
''
Microstonyx
''Microstonyx'' was an extinct genus of suid that existed during the Miocene in Asia and Europe.
Geographic range
Fossils of the species ''M. major'' have been found in Spain, North Macedonia, Turkiye, and China
China, officially the ...
'', the
giraffids ''
Helladotherium
''Helladotherium'' is an extinct genus of sivatheriine giraffid from Europe, Africa, and Asia during the Miocene. The most complete skeleton is that of a female, based on a comparison with an intact female ''Sivatherium giganteum'' skull.
Onl ...
'', ''
Palaeotragus
''Palaeotragus'' ("ancient goat") is a genus of very large, primitive, okapi-like giraffids from the Miocene to Early Pleistocene of Africa and Eurasia.
''Palaeotragus primaevus'' is the older species, being found in early to mid-Miocene strata ...
'', and ''
Samotherium
''Samotherium'' ("beast of Samos") is an extinct genus of Giraffidae from the Miocene and Pliocene of Eurasia and Africa. ''Samotherium'' had two ossicones on its head, and long legs. The ossicones usually pointed upward, and were curved back ...
'', the antelopes ''
Gazella
A gazelle is one of many antelope species in the genus ''Gazella'' . There are also seven species included in two further genera; '' Eudorcas'' and '' Nanger'', which were formerly considered subgenera of ''Gazella''. A third former subgenus, ' ...
'' and ''
Tragoportax
''Tragoportax'' is an extinct genus of Bovidae, bovid ungulate. It lived from the upper Miocene to the earliest Pliocene, and its fossils have been found in southeastern Europe, parts of Africa, and the Indian subcontinent. ''Tragoportax'' was fo ...
'', the
equid
Equidae (commonly known as the horse family) is the Taxonomy (biology), taxonomic Family (biology), family of Wild horse, horses and related animals, including Asinus, asses, zebra, zebras, and many extinct species known only from fossils. The fa ...
''
Hipparion
''Hipparion'' is an extinct genus of three-toed, medium-sized equine belonging to the extinct tribe Hipparionini, which lived about 10-5 million years ago. While the genus formerly included most hipparionines, the genus is now more narrowly defi ...
'', the
chalicothere
Chalicotheriidae (from Ancient Greek ''khálix'', "gravel", and ''theríon'', "beast") is an extinct family of herbivorous, odd-toed ungulate (perissodactyl) mammals that lived in North America, Eurasia, and Africa from the Middle Eocene to the ...
''
Ancylotherium
''Ancylotherium'' (from Greek, meaning "hooked beast") is an extinct genus of the family Chalicotheriidae, subfamily Schizotheriinae, endemic to Europe, Asia, and Africa during the Late Miocene-Early Pleistocene (11.6—1.8 mya), existing for ...
'', the
aardvark
Aardvarks ( ; ''Orycteropus afer'') are medium-sized, burrowing, nocturnal mammals native to Africa. Aardvarks are the only living species of the family Orycteropodidae and the order Tubulidentata. They have a long proboscis, similar to a pi ...
''
Orycteropus
''Orycteropus'' is a genus of mammals in the family Orycteropodidae within Tubulidentata. The genus is known from Late Miocene to recent of Africa
Africa is the world's second-largest and second-most populous continent after Asia. At ab ...
'', and possible remains from the hyena ''
Adcrocuta
''Adcrocuta'' is an extinct genus of large hyena that lived in Africa and Eurasia during the late Miocene epoch.
Distribution and chronology
Fossils of ''A. eximia'' are known from across Eurasia during the Vallesian-Turolian age of the Late ...
'' and the
elephant
Elephants are the largest living land animals. Three living species are currently recognised: the African bush elephant ('' Loxodonta africana''), the African forest elephant (''L. cyclotis''), and the Asian elephant ('' Elephas maximus ...
''Choerolophodon''.
The Chinese ''Yoshi'' species come from the end-Miocene Xingjiawan locality, a humid environment consisting open and forested biomes. The Xingjiawan preserves the elephant ''
Stegodon
''Stegodon'' (from the Ancient Greek στέγω (''stégō''), meaning "to cover", and ὀδούς (''odoús''), meaning "tooth", named for the distinctive ridges on the animal's molars) is an extinct genus of proboscidean, related to elephants ...
'', the hyaenids ''Adcrocuta'', ''
Hyaenictitherium
''Hyaenictitherium'' is an extinct genus of hyaenids that lived throughout Africa, Asia, and possibly Europe during the Late Miocene and Pliocene. The type species, ''H. hyaenoides'', was similar in size to a striped hyena, whereas the smaller ''H. ...
'' and ''
Ictitherium
''Ictitherium'' (meaning "weasel beast") is an extinct genus belonging to the family Hyaenidae and the subfamily Ictitheriinae erected by Trouessart in 1897. ''Ictitherium'' species were endemic to Africa and Eurasia during the Middle Miocene t ...
'', and the felids ''
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 genera as '' Xenosmilus'', ''Homotherium'' itself, and ''Machairodus''. It inhab ...
'' and ''
Pristifelis
''Pristifelis'' is an extinct genus of feline from the late Miocene. It contains a single species, ''Pristifelis attica''. The first fossil skull of ''P. attica'' was excavated near Pikermi in Attica, Greece.
Fossils were also excavated near th ...
''.
References
{{Taxonbar, from1=Q28432831, from2=Q120433497, from3=Q107410997, from4=Q107411031, from5=Q120433536, from6=Q122047315
Metailurini
Miocene carnivorans
Miocene genus extinctions
Prehistoric carnivoran genera
Miocene mammals of Europe
Fossil taxa described in 2014