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Machairodontinae

Extinct subfamily of sabertooth cats

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Montage on species of the subfamily Machairodontinae: Homotherium venezuelensis , Machairodus aphanistus , Metailurus sp. and Smilodon fatalis .
George Miquilena PePeEfe Izvora Ryan Somma · CC BY-SA 4.0

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Through Extinction, Homo and Tiger

Machairodonts varied in size from comparable to lynxes to exceeding that of lions. The Machairodontinae contain many of the extinct predators commonly known as "saber-toothed cats", including those with greatly elongated upper maxillary canines, such as the famed genus Smilodon and Megantereon, though the degree of elongation was variable, and in some machairodontines like Dinofelis the length of the upper canines was much more modest. Sometimes, other carnivorous mammals with elongated teeth are also called saber-toothed cats, although they do not belong to the felids. Besides the machairodonts, other saber-toothed predators included gorgonopsians, nimravids, machaeroidines, and thylacosmilids. Unlike living big cats, which generally clamp the muzzle or throat of prey to asphyxiate them, saber-toothed machairodontines are thought to have killed prey using a bite to the neck once immobilised, using their neck muscles to drive the saber teeth into the throat while the lower jaw served as an anchor, causing rapid death via blood loss.

Likely originating in Eurasia during the Middle Miocene, they eventually spread to every continent except Australia and Antarctica. Machairodonts were the dominant group of cats and large mammalian predators across Afro-Eurasia and North America during the late Miocene and Early Pliocene, a time when the ancestors of living cats (Felinae sensu lato) were mostly small sized. Machairodonts began to decline from the end of the Miocene onwards, which accelerated during the Pleistocene, perhaps as a result of environmental change and consequential changes in prey abundance, competition with large living cat lineages such as the pantherins as well as possibly archaic humans. The last species belonging to the genera Smilodon and Homotherium became extinct along with many other large mammals around 12-10,000 years ago as part of the end-Pleistocene extinction event, following human arrival to the Americas at the end of the Late Pleistocene.

Evolution

Family Felidae

Based on mitochondrial DNA sequences extracted from fossils, machairodontines diverged from the ancestors of living cats around 20 million years ago, with the last surviving machairodont genera Homotherium and Smilodon estimated to have diverged from each other about 18 million years ago.

The Machairodontinae originated in the middle Miocene of Europe. The early felid Pseudaelurus quadridentatus showed a trend towards elongated upper canines, and is believed to be at the base of the machairodontine evolution. The earliest known machairodont genus is the middle Miocene Miomachairodus from Africa and Turkey. Until the late Miocene, machairodontines co-existed at several places together with barbourofelids, archaic large carnivores that also bore long sabre-teeth.

Traditionally, three different tribes of machairodontines were recognized, the Smilodontini with typical dirk-toothed forms, such as Megantereon and Smilodon, the Machairodontini or Homotherini with scimitar-toothed cats, such as Machairodus or Homotherium, and the Metailurini, containing genera such as Dinofelis and Metailurus. However, some in the past have regrouped the Metailurini within the other felid subfamily, the Felinae, along with all modern cats. A 2022 phylogenetic study suggests a polyphyletic relationship between Metailurini and Smilodontini, with the genus Paramachairodus being ancestral to both groups.

The name 'saber-toothed tigers' is misleading. Machairodonts were not in the same subfamily as tigers, there is no evidence that they had tiger-like coat patterns, and this broad group of animals did not all live or hunt in the same manner as the modern tiger. DNA analysis published in 2005 confirmed and clarified cladistic analysis in showing that the Machairodontinae diverged early from the ancestors of modern cats and are not closely related to any living feline species.

Classification

Phylogeny

Phylogeny after Werdelin and Flink (2018):

Composite cladogram following Lautenschlager et al. (2020):

Evolutionary history and origin of phenotype

Until the recent discovery of the Late Miocene fossil depository known as Batallones-1 in the 1990s, specimens of Smilodontini and Homotheriini ancestors were rare and fragmentary, so the evolutionary history of the saber-toothed phenotype, a phenotype affecting craniomandibular, cervical forelimb and forelimb anatomy, was largely unknown. Prior to the excavation of Batallones-1, the predominating hypothesis was that the highly derived saber-toothed phenotype arose rapidly through pleiotropic evolution. Batollnes-1 unearthed new specimens of Promegantereon ogygia, a Smilodontini ancestor, and Machairodus aphanistus, a Homotheriini ancestor, shedding light on evolutionary history. (Though the Smilodontini ancestor was originally assigned to the genus Paramachairodus, it was later revised to the genus Promegantereon). The leopard-sized P. ogygia (living 9.0 Ma) inhabited Spain (and perhaps additional territory).

The current hypothesis for the evolution of the saber-toothed phenotype, made possible by Batollnes-1, is that this phenotype arose gradually over time through mosaic evolution. Although the exact cause is uncertain, current findings have supported the hypothesis that a need for the rapid killing of prey was the principal pressure driving the development of the phenotype over evolutionary time. As indicated by high instances of broken teeth, the biotic environment of saber-toothed cats was one marked by intense competition.

Broken teeth indicate the frequency at which teeth contact bone. Increased teeth-bone contact suggests either increased consumption of carcasses, rapid consumption of prey, or increased aggression over kills – all three of which point to decreased prey availability, heightening competition between predators. Such a competitive environment would favor the faster killing of prey, because if prey is taken away before consumption (such as by out-competing) the energetic cost of capturing that prey is not reimbursed, and, if this occurs often enough in the lifetime of a predator, death by exhaustion or starvation would result. The earliest adaptations improving the speed at which prey was killed are present in the skull and mandible of P. ogygia and of M. aphanistus, and in the cervical vertebrae and forelimb of P. ogygia. They provide further morphological evidence for the importance of speed in the evolution of the saber-toothed phenotype.

Machariodont diversity declined from the end of the Miocene onwards, especially during the Pleistocene. By the Late Pleistocene, only two genera of machairodonts remained: Smilodon, and the distantly related Homotherium, both largely confined to the Americas. These two genera became extinct around 13,000-10,000 years ago as part of the wave of extinctions of most large animals across the Americas.

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