Thylacosmilid
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Thylacosmilidae is an extinct
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
of
metatherian Metatheria is a mammalian clade that includes all mammals more closely related to marsupials than to placentals. First proposed by Thomas Henry Huxley in 1880, it is a more inclusive group than the marsupials; it contains all marsupials as well ...
predators, related to the modern
marsupials Marsupials are a diverse group of mammals belonging to the infraclass Marsupialia. They are natively found in Australasia, Wallacea, and the Americas. One of marsupials' unique features is their reproductive strategy: the young are born in a ...
, which lived in
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...
between the
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 ...
and
Pliocene The Pliocene ( ; also Pleiocene) is the epoch (geology), epoch in the geologic time scale that extends from 5.33 to 2.58
, these animals belonged to the order
Sparassodonta Sparassodonta (from Ancient Greek, Greek to tear, rend; and , gen.
, ' The comma is a punctuation mark that appears in several variants in different languages. Some typefaces render it as a small line, slightly curved or straight, but inclined from the vertical; others give it the appearance of a miniature fille ...
tooth) is an extinct order (biology), order of carnivore, carnivorous metatherian mammals native to South America, related to modern marsupials. They were once cons ...
, which occupied the ecological niche of many
eutherian Eutheria (from Greek , 'good, right' and , 'beast'; ), also called Pan-Placentalia, is the clade consisting of placental mammals and all therian mammals that are more closely related to placentals than to marsupials. Eutherians are distingu ...
mammals of the order
Carnivora Carnivora ( ) is an order of placental mammals specialized primarily in eating flesh, whose members are formally referred to as carnivorans. The order Carnivora is the sixth largest order of mammals, comprising at least 279 species. Carnivor ...
from other continents. The family's most notable feature are the elongated, laterally flattened fangs, which is a remarkable evolutionary convergence with other saber-toothed mammals like '' Barbourofelis'' and ''
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 ...
''.


Taxonomic history

The family Thylacosmilidae was originally erected by Riggs in 1933, to accommodate '' Thylacosmilus'', found in the Pliocene Brochero Formation of Argentina. Later, the family was demoted to a subfamily, as Thylacosmilinae, within Borhyaenidae, a group of superficially canid-like sparassodonts, under the assumption that ''Thylacosmilus'' was merely a late and specialized borhyaenid. Later, with the discovery of fragmentary specimens of new sparassodonts related to ''Thylacosmilus'' from Miocene and Pliocene strata, Thylacosmilidae was promoted back to familial status.Analía M. Forasiepi & Alfredo A. Carlini (2010). A new thylacosmilid (Mammalia, Metatheria, Sparassodonta) from the Miocene of Patagonia, Argentina. ''Zootaxa'' 2552: 55–68 In 1997, a second genus and species of thylacosmilid was described from the
Laventan The Laventan () age is a period of geologic time (13.8 to 11.8 Ma) within the Middle Miocene epoch of the Neogene, used more specifically within the SALMA classification in South America. It follows the Colloncuran and precedes the Mayoan age. ...
Honda Group at the
Lagerstätte A Fossil-Lagerstätte (, from ''Lager'' 'storage, lair' '' Stätte'' 'place'; plural ''Lagerstätten'') is a sedimentary deposit that preserves an exceptionally high amount of palaeontological information. ''Konzentrat-Lagerstätten'' preserv ...
La Venta La Venta is a pre-Columbian archaeological site of the Olmec civilization located in the present-day Mexican state of Tabasco. Some of the artifacts have been moved to the museum "Parque - Museo de La Venta", which is in nearby Villaherm ...
in
Colombia Colombia, officially the Republic of Colombia, is a country primarily located in South America with Insular region of Colombia, insular regions in North America. The Colombian mainland is bordered by the Caribbean Sea to the north, Venezuel ...
, '' Anachlysictis gracilis''. This animal, less specialized than ''Thylacosmilus'', was the first indication that the family's origin dates back to before the end of the Miocene. In fact, the anatomy of ''Anachlysictis molar teeth suggests a closer relationship with basal sparassodonts like '' Hondadelphys'' than with advanced sparassodonts like '' Borhyaena''.Goin, F.J. 2003. Early marsupial radiations in South America. En: M. Jones, C. Dickman y M. Archer (eds.), ''Predators with Pouches, The Biology of Carnivorous Marsupials'', CSIRO Publishing, Australia, pp. 30-42. Also, additional materials of a small predatory sparassodont of Colombia have been found, which has certain features diagnostic of thylacosmilids, but much less specialized, as well as indeterminate remains in
Uruguay Uruguay, officially the Oriental Republic of Uruguay, is a country in South America. It shares borders with Argentina to its west and southwest and Brazil to its north and northeast, while bordering the Río de la Plata to the south and the A ...
and the Argentinean
Patagonia Patagonia () is a geographical region that includes parts of Argentina and Chile at the southern end of South America. The region includes the southern section of the Andes mountain chain with lakes, fjords, temperate rainforests, and glaciers ...
, from the early Pliocene, has been tentatively assigned to family. Forasiepi and Carlini in 2010 unveiled a third genus and species, '' Patagosmilus goini'', from the Collón Cura Formation of Argentina from the mid-Miocene, with characteristics intermediate between ''Anachlysictis'' and ''Thylacosmilus''.


Description

The family is perhaps the best known among sparassodonts because of their dental and cranial specializations, which are superficially similar to that of the
saber-toothed cats 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 ...
, often cited as an example of
convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last comm ...
between placental and metatherian mammals. However, there were several differences between thylacosmilids and the other saber-toothed mammals, and these unique traits diagnose the family: unique traits include canine teeth that grew continuously, less specialized
carnassial Carnassials are paired upper and lower teeth modified in such a way as to allow enlarged and often self-sharpening edges to pass by each other in a shearing manner. This adaptation is found in carnivorans, where the carnassials are the modified f ...
molars and tremendous,
flange A flange is a protruded ridge, lip or rim (wheel), rim, either external or internal, that serves to increase shear strength, strength (as the flange of a steel beam (structure), beam such as an I-beam or a T-beam); for easy attachment/transfer o ...
-like outgrowths of the lower jaw that protected the saber-teeth. In ''Thylacosmilus'', the last and most specialized known member, the incisors are very small and the lower teeth are poorly developed, and tack-shaped; in the other genera these elements are unknown. In ''Thylacosmilus'', the upper incisors are unknown other than by wear indentations on the lower incisors, as no premaxilla has ever been found complete in this genus. Another evolutionary trend in families is the progressive reduction of
masseter In anatomy, the masseter is one of the muscles of mastication. Found only in mammals, it is particularly powerful in herbivores to facilitate chewing of plant matter. The most obvious muscle of mastication is the masseter muscle, since it is the ...
and temporal muscles, resulting in relatively weak bites, but compensated by the increase in the size of the neck muscles to lower the head and the fangs into the necks of their prey. Fossils of ''Thylacosmilus'' forelimbs, the only reported for this group, indicate that animals were not fast runners, and were in turn adapted to exert force in order to subdue their prey, helping with their semiopposable thumb. A recent study has since proposed a lack of analogy between thylacosmilids and saber toothed eutherians, speculating that rather than using the fangs to pierce prey they were instead used to open up corpses. Their lack of incisors and crushing molars suggest an entrail-based diet, and irrigated bone on the maxillae suggests some sort of extensive, soft tissue-based structure.


References

{{Taxonbar, from=Q5493808 Sparassodonta Prehistoric mammal families Neogene mammals of South America Miocene first appearances Pliocene extinctions