Paramachairodus
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''Paramachaerodus'' 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 saber-tooth cat of the
subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end botanical subfamily names with "-oideae", and zo ...
Machairodontinae Machairodontinae (from Ancient Greek μάχαιρα ''Makhaira, 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 ...
, which was
endemic Endemism is the state of a species being found only in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also foun ...
to
Eurasia Eurasia ( , ) is a continental area on Earth, comprising all of Europe and Asia. According to some geographers, Physical geography, physiographically, Eurasia is a single supercontinent. The concept of Europe and Asia as distinct continents d ...
during the Middle and
Late Miocene The Late Miocene (also known as Upper Miocene) is a sub-epoch of the Miocene epoch (geology), Epoch made up of two faunal stage, stages. The Tortonian and Messinian stages comprise the Late Miocene sub-epoch, which lasted from 11.63 Ma (million ye ...
from 15 to 9 Ma. A 2022 phylogenetic analysis suggested that the genus may be
polyphyletic A polyphyletic group is an assemblage that includes organisms with mixed evolutionary origin but does not include their most recent common ancestor. The term is often applied to groups that share similar features known as Homoplasy, homoplasies ...
. ''Paramacherodus'' is one of the earliest known true
saber-toothed 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 ...
s. Many
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserve ...
s were discovered in Cerro de los Batallones, a Late Miocene fossil site near
Madrid Madrid ( ; ) is the capital and List of largest cities in Spain, most populous municipality of Spain. It has almost 3.5 million inhabitants and a Madrid metropolitan area, metropolitan area population of approximately 7 million. It i ...
,
Spain Spain, or the Kingdom of Spain, is a country in Southern Europe, Southern and Western Europe with territories in North Africa. Featuring the Punta de Tarifa, southernmost point of continental Europe, it is the largest country in Southern Eur ...
. One leopard-sized species is known, ''Paramachaerodus orientalis'' from the Turolian. A second species, ''Paramacharodus maximiliani'', has been considered a synonym of ''Paramachaerodus orientalis'' by some authors, but was considered a valid species in the most recent systematic revision. That revision, based on an extensive morphological analysis, also determined that the species ''P. ogygia'' exhibited less derived sabertooth features than the other ''Paramachaerodus'' species and should be assigned to a separate genus, '' Promegantereon''.


Description

The animals were about high at the shoulder, similar to a
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 ...
, but with a more supple body. The shape of its limbs suggests that it may have been an agile climber, and could have hunted relatively large prey.


History and naming

In 1913, Guy Ellcock Pilgrim named the new genus ''Paramachaerodus'' for the species ''Machaerodus orientalis'', ''Machaerodus schlosseri'', and ''Felis ogygia''; at the time, however, he failed to designate a type species. In 1915 he described more material that he assigned to ''Paramachaerodus'' cf. ''schlosseri'' (two hemimandibles, GSI-140 and GSI-141), though Matthew (1929) noted that both of those specimens did not closely resemble others in the genus. He rectified the lack of a type species in 1931 by designating ''Paramachaerodus orientalis'' the type species, now including ''Paramachaerodus schlosseri'' as a junior synonym. However, in the intervening span of time, Miklos Kretzoi had proposed another new genus, ''Pontosmilus'', for the species ''P. orientalis'' (also the type species), ''P. schlosseri'', ''P. ogygia'', ''P. hungaricus'', and the new species ''Pontosmilus indicus'' that he described based on GSI-141. He restricted ''Paramachaerodus'' to another new species, ''P. pilgrimi'', that he described based on GSI-140. He also proposed the genus ''Proamphimachairodus'' for the species ''Machairodus maximiliani''. With Pilgrim's clarification of ''Paramachaerodus'' in 1931, and utter rejection of both ''Pontosmilus'' and ''Proamphimachairodus''-he included ''Machairodus orientalis'', ''Felis ogygia'', and ''Machairodus maximiliani'' as species of ''Paramachaerodus''-''Pontosmilus'' was rendered an invalid genus, for it possessed no valid type species, and both ''Pontosmilus'' and ''Proamphimachairodus'' were designated junior synonyms of ''Paramachaerodus''. A major review of the genus in 2010 designated ''P. matthewi'', ''P. schlosseri'', ''P. hungaricus'' as junior synonyms of ''P. orientalis'', with ''P. maximiliani'' the only other valid species in ''Paramachaerodus'', and assigning ''P. agygia'' back to ''Promegantereon''. It also noted that ''"Pontosmilus" indicus'' (GSI-141) was a feline, and ''"Pontosmilus" pilgrimi'' (GSI-140) a machairodontine not of ''Paramachaerodus'' or ''Promegantereon'', though the authors offered no alternate genus assignments for either species. A third species, ''Paramachaerodus transasiaticus'', was described in 2017 based on analysis of new fossil material from the late Miocene localities of Hezheng, Gansu Province, China, and Hadjidimovo, Bulgaria. These specimens had sabertooth characteristics intermediate between those of ''P. ogygia'' and those of ''P. orientalis'' and ''P. maximiliani''. In 2022, a new species ''Paramachaerodus yingliangi'' was proposed based on fossils from northeastern China; the same paper also proposed separating ''Paramachaerodus schlosseri'' as the new type species and moving ''P. orientalis'' and ''P. maximiliani'' to the resurrected genus ''Pontosmilus''.


Classification

The position of ''Paramachaerodus'' within the Machairodontinae has been subject to much controversy, partially because many names have historically been proposed for it based on only scanty material. While ''Paramachaerodus'' is generally accepted as a close relative and probable forerunner of ''
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 therefore an ancestor of ''
Smilodon ''Smilodon'' is an extinct genus of Felidae, felids. It is one of the best known saber-toothed predators and prehistoric mammals. Although commonly known as the saber-toothed tiger, it was not closely related to the tiger or other modern cats ...
'', the role ''Promegantereon'' played in this is still controversial. While researchers have generally favoured the classification of ''Promegantereon'' as a distinct genus starting in the early 2000's, it was still thought to be a close relative and potential ancestor of ''Paramachaerodus''. Thus, ''Paramachaerodus''' position within the Smilodontini was generally understood as intermediate, with ''Promegantereon'' as the first and ''Smilodon'' as the ultimate representative of that machairodontine tribe. However, a 2022 phylogenetic analysis by Jiangzuo et al. cast doubt on this, and indeed on the
monophyly In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent comm ...
of ''Paramachaerodus'' itself. While it confirmed ''P. orientalis'' as basal to the ''Megantereon-Smilodon'' clade (the Smilodontini
sensu stricto ''Sensu'' is a Latin word meaning "in the sense of". It is used in a number of fields including biology, geology, linguistics, semiotics, and law. Commonly it refers to how strictly or loosely an expression is used in describing any particular c ...
) as proposed earlier, ''P. maxilmiliani'' was instead found to be basal to a clade comprising '' Rhizosmilodon'' and '' Dinofelis''. At the same time, ''Promegantereon'' and ''Paramachaerodus transasiaticus'' were found to group together with '' Metailurus'' and ''
Yoshi Yoshi is a fictional dinosaur who appears in video games published by Nintendo. Yoshi debuted in '' Super Mario World'' (1990) on the SNES as Mario and Luigi's sidekick. Throughout the mainline '' Super Mario'' series, Yoshi typically serves ...
'', which are usually placed in the separate Metailurini tribe. This would make ''Paramachaerodus'', the Smilodontini and the Metailurini as traditionally definded
polyphyletic A polyphyletic group is an assemblage that includes organisms with mixed evolutionary origin but does not include their most recent common ancestor. The term is often applied to groups that share similar features known as Homoplasy, homoplasies ...
, and calls for more work regarding machairodontine systematics, suggesting that a major revision of the subfamily may be needed.


Paleobiology

Based on the morphology of its
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 ...
, ''P. orientalis'' is inferred to have been adapted for wooded environments.


Paleoecology


References


External links


Paramachairodus at the bluelion.org
{{Taxonbar, from1=Q2051668, from2=Q122231603, from3=Q122231608, from4=Q122231610, from5=Q122231612 Smilodontini Miocene felids Prehistoric carnivoran genera Fossil taxa described in 1913