Systemodon
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''Systemodon'' is a genus of
early Eocene In the geologic timescale the Ypresian is the oldest age (geology), age or lowest stage (stratigraphy), stratigraphic stage of the Eocene. It spans the time between , is preceded by the Thanetian Age (part of the Paleocene) and is followed by th ...
mammal of
Wasatchian The Wasatchian North American Stage on the geologic timescale is the North American faunal stage according to the North American Land Mammal Ages chronology (NALMA), typically set from 55,400,000 to 50,300,000 years Before Present, BP lasting . I ...
age (ca. 55-50 mya). It was one of many mammals originally considered the earliest horses, long classified in the genus ''
Hyracotherium ''Hyracotherium'' ( ; "hyrax-like beast") is an extinction, extinct genus of small (about 60 cm in length) perissodactyl ungulates that was found in the London Clay formation. This small, fox-sized animal is (for some scientists) considered t ...
'' (commonly known as ''Eohippus''). These were dog-sized animals that in life would have looked vaguely like a
paca A paca ia a rodent in South and Central America. Paca or PACA may also refer to: People * William Paca (1740–1799), a Founding Father of the United States * Paca Blanco (Francisca Blanco Díaz, born 1949), Spanish activist * Paca Navas (Franc ...
,
mara Mara or MARA may refer to: Arts and entertainment Fictional characters * Mara (''Doctor Who''), an evil being in two ''Doctor Who'' serials * Mara (She-Ra), fictional characters from the ''She-Ra and the Princesses of Power'' and ''The New Advent ...
, or
chevrotain Chevrotains, or mouse-deer, are small, even-toed ungulates that make up the family Tragulidae, and are the only living members of the infraorder Tragulina. The 10  extant species are placed in three genera, but several species also ar ...
(though they were perissodactyls, not
rodent Rodents (from Latin , 'to gnaw') are mammals of the Order (biology), order Rodentia ( ), which are characterized by a single pair of continuously growing incisors in each of the upper and Mandible, lower jaws. About 40% of all mammal specie ...
s or
artiodactyls Artiodactyls are placental mammals belonging to the order Artiodactyla ( , ). Typically, they are ungulates which bear weight equally on two (an even number) of their five toes (the third and fourth, often in the form of a hoof). The other thre ...
.) The type species, ''S. tapirinus'', is represented by 24 individuals from a locale called the Castillo pocket in the Huerfano Formation of Colorado. This well-preserved deposit allows researchers to reconstruct aspects of the environment and lifestyle of the species.


Taxonomy

The genus was named by E.D. Cope in 1875, who recognized it as different from ''Hyracotherium'' and a basal
perissodactyl Perissodactyla (, ), or odd-toed ungulates, is an order of Ungulate, ungulates. The order includes about 17 living species divided into three Family (biology), families: Equidae (wild horse, horses, Asinus, asses, and zebras), Rhinocerotidae ( ...
. It was later believed to be an
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 ...
and referred to ''Hyracotherium'' (''Hyracotherium tapirinum'', incl. ''H. cristatum'' Wortman 1896, also =''Orohippus tapirinus''). A 1984 analysis revealed this species did not fit well with others in the genus. Such studies revealed that ''Hyracotherium'' had become a
wastebasket taxon Wastebasket taxon (also called a wastebin taxon, dustbin taxon or catch-all taxon) is a term used by some taxonomists to refer to a taxon that has the purpose of classifying organisms that do not fit anywhere else. They are typically defined by e ...
of early perissodactyls; many species have now been reassigned to other genera whose exact relationships are not yet resolved. ''Systemodon'' has been identified as allied to ''Cymbalophus'' and as one of the most basal perissodactyls, making it part of the stem group ancestral to horses, rhinos, tapirs, and the extinct
brontotheres Brontotheriidae is a family of extinct mammals belonging to the order Perissodactyla, the order that includes horses, rhinoceroses, and tapirs. Superficially, they looked rather like rhinos with some developing bony nose horns, and were some of ...
and
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 ...
s. If ''Systemodon'' is a stem perissodactyl, its unusually good preservation gives a rare glimpse into what is basal in the behavior as well as the form of a large group of mammals. Three other species were formerly assigned to this genus: ''S. semihians, S. protapirinus'', and ''S. primaevus''. The first has been reassigned to ''Cardiolophus'', and the second and third synonymized as ''Hyracotherium protapirinus''.


Paleobiology

''S. tapirinus'' lived in patchy woodlands near streams, an environment it shared with a smaller species of perissodactyl, '' Hyracotherium vasacciense''. Other inhabitants of these environments included lemur-like and tarsier-like early primates, and flesh-eating
creodonts Creodonta ("meat teeth") is a former order of extinct carnivorous placental mammals that lived from the early Paleocene to the late Miocene epochs in North America, Europe, Asia and Africa. Originally thought to be a single group of animals ances ...
. Analysis of its teeth and fossil pollen suggests that while ''Systemodon'' did not have the high-crowned molars needed to chew grass, it did not eat forest plants as has often been assumed for early
ungulate Ungulates ( ) are members of the diverse clade Euungulata ("true ungulates"), which primarily consists of large mammals with Hoof, hooves. Once part of the clade "Ungulata" along with the clade Paenungulata, "Ungulata" has since been determined ...
s, but browsed instead on
forb A forb or phorb is a herbaceous flowering plant that is not a graminoid (grass, sedge, or rush). The term is used in botany and in vegetation ecology especially in relation to grasslands and understory. Typically, these are eudicots without woo ...
s and flowers in more open areas. Both females and males had enlarged canine teeth. Fossils show two clear adult size groups that indicate
sexual dimorphism Sexual dimorphism is the condition where sexes of the same species exhibit different Morphology (biology), morphological characteristics, including characteristics not directly involved in reproduction. The condition occurs in most dioecy, di ...
, with male skulls about 15% larger than females, with 40% larger canine teeth. The presence of the genus coincides with a cooling and drying period where the closed tropical forests of the Paleocene-Eocene Thermal Maximum gave way to new, more open environments similar to modern park woodlands and savannas. Based on comparison with the anatomy and habits of living ungulates, it is likely that ''Systemodon'' lived in female groups, herding together as protection from predators in open spaces or to make the best use of localized patches of food in mixed environments. Prime-age males may have competed with each other to live in these groups, and used their canines in ritualized competitive battles or displays.


References

{{Taxonbar, from=Q30894974, from2= Eocene mammals of North America Eocene Perissodactyla Prehistoric Perissodactyla Taxa named by Edward Drinker Cope