Greek
Greek may refer to:
Greece
Anything of, from, or related to Greece, a country in Southern Europe:
*Greeks, an ethnic group.
*Greek language, a branch of the Indo-European language family.
**Proto-Greek language, the assumed last common ancestor ...
extinct
Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
carnivorous
A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, fat and other sof ...
metatheria
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 ...
n
mammal
Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...
s native to South America, related to modern
marsupial
Marsupials are any members of the mammalian infraclass Marsupialia. All extant marsupials are endemic to Australasia, Wallacea and the Americas. A distinctive characteristic common to most of these species is that the young are carried in a po ...
s. They were once considered to be true marsupials, but are now thought to be a separate side branch that split before the last common ancestor of all modern marsupials. A number of these mammalian predators closely resemble
placental
Placental mammals (infraclass Placentalia ) are one of the three extant subdivisions of the class Mammalia, the other two being Monotremata and Marsupial
Marsupials are any members of the mammalian infraclass Marsupialia. All extant marsup ...
predators that evolved separately on other continents, and are cited frequently as examples 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 com ...
. They were first described by
Florentino Ameghino
Florentino Ameghino (born Giovanni Battista Fiorino Giuseppe Ameghino September 19, 1853 – August 6, 1911) was an Argentine naturalist, paleontologist, anthropologist and zoologist, whose fossil discoveries on the Argentine Pampas, especially ...
, from fossils found in the Santa Cruz beds of
Patagonia
Patagonia () refers to a geographical region that encompasses the southern end of South America, governed by Argentina and Chile. The region comprises the southern section of the Andes Mountains with lakes, fjords, temperate rainforests, and gl ...
. Sparassodonts were present throughout South America's long period of "splendid isolation" during the
Cenozoic
The Cenozoic ( ; ) is Earth's current geological era, representing the last 66million years of Earth's history. It is characterised by the dominance of mammals, birds and flowering plants, a cooling and drying climate, and the current configura ...
; during this time, they shared the niches for large warm-blooded predators with the flightless
terror bird
Phorusrhacids, colloquially known as terror birds, are an extinct clade of large carnivorous flightless birds that were one of the largest species of apex predators in South America during the Cenozoic era; their conventionally accepted temporal ...
s. Previously, it was thought that these mammals died out in the face of competition from "more competitive"
placental
Placental mammals (infraclass Placentalia ) are one of the three extant subdivisions of the class Mammalia, the other two being Monotremata and Marsupial
Marsupials are any members of the mammalian infraclass Marsupialia. All extant marsup ...
carnivorans
Carnivora is a monophyletic order of placental mammals consisting of the most recent common ancestor of all cat-like and dog-like animals, and all descendants of that ancestor. Members of this group are formally referred to as carnivorans, ...
during the
Pliocene
The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58Great American Interchange
The Great American Biotic Interchange (commonly abbreviated as GABI), also known as the Great American Interchange and the Great American Faunal Interchange, was an important late Cenozoic paleozoogeographic biotic interchange event in which lan ...
, but more recent research has showed that sparassodonts died out long before eutherian carnivores arrived in South America (aside from
procyonids
Procyonidae is a New World family of the order Carnivora. It comprises the raccoons, ringtails, cacomistles, coatis, kinkajous, olingos, and olinguitos. Procyonids inhabit a wide range of environments and are generally omnivorous.
Characte ...
, which sparassodonts probably did not directly compete with). Sparassodonts have been referred to as borhyaenoids by some authors, but currently the term Borhyaenoidea refers to a restricted subgroup of sparassodonts comprising
borhyaenids
Borhyaenidae is an extinct metatherian family (biology), family of low-slung, heavily built predatory mammals in the order Sparassodonta. Borhyaenids are not true marsupials, but members of a sister taxon, Sparassodonta. Like most metatherians, ...
and their close relatives.
Taxonomy
Classification
Sparassodonts can be divided into six major groups; basal sparassodonts (?earliest
Paleocene
The Paleocene, ( ) or Palaeocene, is a geological epoch (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), E ...
-late
Miocene
The Miocene ( ) is the first 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 means "less recen ...
), species that cannot be easily assigned to any of the other sparassodont groups and whose teeth often exhibit adaptations for
omnivory
An omnivore () is an animal that has the ability to eat and survive on both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize the nut ...
; hathliacynids (late
Oligocene
The Oligocene ( ) is a geologic epoch of the Paleogene Period and extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that define the epoch are well identified but the ...
-early
Pliocene
The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58Pliocene
The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58marten
A marten is a weasel-like mammal in the genus ''Martes'' within the subfamily Guloninae, in the family Mustelidae. They have bushy tails and large paws with partially retractile claws. The fur varies from yellowish to dark brown, depending on t ...
to a
thylacine
The thylacine ( , or , also ) (''Thylacinus cynocephalus'') is an extinct carnivorous marsupial that was native to the Australian mainland and the islands of Tasmania and New Guinea. The last known live animal was captured in 1930 in Tasman ...
in size, and have long, fox-like snouts and teeth strongly adapted for
carnivory
A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, adipose tissue, f ...
; basal borhyaenoids (middle
Eocene
The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
-late
Miocene
The Miocene ( ) is the first 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 means "less recen ...
), borhyaenoids that cannot be easily classified into the families
Borhyaenidae
Borhyaenidae is an extinct metatherian family of low-slung, heavily built predatory mammals in the order Sparassodonta. Borhyaenids are not true marsupials, but members of a sister taxon, Sparassodonta. Like most metatherians, borhyaenids and o ...
,
Thylacosmilidae
Thylacosmilidae is an extinct Family (biology), family of metatherian predators, related to the modern marsupials, which lived in South America between the Miocene and Pliocene epochs. Like other South American mammalian predators that lived prio ...
, or
Proborhyaenidae
Proborhyaenidae is an extinct family of metatherian mammals of the order Sparassodonta, which lived in South America from the Eocene (Mustersan) until the Oligocene (Deseadan). Sometimes it has been included as a subfamily of their relatives, the ...
and vary in size and shape; borhyaenids (early-late
Miocene
The Miocene ( ) is the first 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 means "less recen ...
), the sparassodont group most specialized for running, but not as much as living
carnivorans
Carnivora is a monophyletic order of placental mammals consisting of the most recent common ancestor of all cat-like and dog-like animals, and all descendants of that ancestor. Members of this group are formally referred to as carnivorans, ...
or even
thylacines
The thylacine ( , or , also ) (''Thylacinus cynocephalus'') is an extinct carnivorous marsupial that was native to the Australian mainland and the islands of Tasmania and New Guinea. The last known live animal was captured in 1930 in Tasma ...
; proborhyaenids (middle
Eocene
The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
-late
Oligocene
The Oligocene ( ) is a geologic epoch of the Paleogene Period and extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that define the epoch are well identified but the ...
), robust,
wolverine
The wolverine (), (''Gulo gulo''; ''Gulo'' is Latin for "gluttony, glutton"), also referred to as the glutton, carcajou, or quickhatch (from East Cree, ''kwiihkwahaacheew''), is the largest land-dwelling species of the family Mustelidae. It is ...
-like forms with ever-growing upper and lower
canines
Canine may refer to:
Zoology and anatomy
* a dog-like Canid animal in the subfamily Caninae
** ''Canis'', a genus including dogs, wolves, coyotes, and jackals
** Dog, the domestic dog
* Canine tooth, in mammalian oral anatomy
People with the surn ...
; and thylacosmilids (early
Miocene
The Miocene ( ) is the first 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 means "less recen ...
-late
Pliocene
The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58canines
Canine may refer to:
Zoology and anatomy
* a dog-like Canid animal in the subfamily Caninae
** ''Canis'', a genus including dogs, wolves, coyotes, and jackals
** Dog, the domestic dog
* Canine tooth, in mammalian oral anatomy
People with the surn ...
.
The taxonomic classification below follows the latest review of the group, that of Prevosti and Forasiepi (2018), with additions from more recent studies. Although ''Mayulestes'' was originally described as a sparassodont, later phylogenetic analyses have shown that it most likely does not belong to this group; however more recent studies show it to be closely related to sparassodonts. Similarly, while basal borhyaenoids such as ''Lycopsis'' and ''Prothylacynus'' were once thought to belong to a distinct family (Prothylacynidae), phylogenetic analyses have found that these animals do not represent a
monophyletic
In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
group. The exact age of most Eocene species of sparassodonts is uncertain, given the lack of precise stratigraphic information associated with most specimens and the recent division of the
Casamayoran The Casamayoran ( es, Casamayorense) age is a period of geologic time (50.0–48.0 Ma) within the Early Eocene epoch of the Paleogene, used more specifically within the South American land mammal age (SALMA) classification. It follows the Itaboraia ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
s.
* Order Sparassodonta
** Genus '' Allqokirus''
***''Allqokirus australis'' (earliest
Paleocene
The Paleocene, ( ) or Palaeocene, is a geological epoch (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), E ...
,
Tiupampan The Tiupampan ( es, Tiupampense) age is a period of geologic time (64.5–62.5 Ma) within the Paleocene epoch of the Paleogene used more specifically with South American land mammal ages (SALMA). It is the oldest SALMA age and precedes the Peligra ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
** Genus ''
Mayulestes
''Mayulestes'' (Quechua: ''mayu'' river, + Greek: ''lestes'', thief) is a genus of carnivorous metatherian that lived in what is now Tiupampa, Bolivia in the early Paleocene. It shared its habitat with fellow sparrasodont ''Pucadelphys'', and a ...
''
***''Mayulestes ferox'' (earliest
Paleocene
The Paleocene, ( ) or Palaeocene, is a geological epoch (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), E ...
,
Tiupampan The Tiupampan ( es, Tiupampense) age is a period of geologic time (64.5–62.5 Ma) within the Paleocene epoch of the Paleogene used more specifically with South American land mammal ages (SALMA). It is the oldest SALMA age and precedes the Peligra ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
** Genus '' Argyrolestes''
** Genus '' Nemolestes''
***''Nemolestes spalacotherinus'' (late middle
Eocene
The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
** Genus '' Patene''
***''Patene campbelli'' (late ?
Eocene
The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
Eocene
The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
)
***''Patene coluapiensis'' (middle
Eocene
The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
***''Patene simpsoni'' (late
Paleocene
The Paleocene, ( ) or Palaeocene, is a geological epoch (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), E ...
/early
Eocene
The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
,
Itaboraian
The Itaboraian ( pt, Itaboraiense) age is a period within the Early Eocene geologic time (53.0–50.0 Ma) epoch of the Paleogene, used more specifically with South American land mammal ages (SALMA). It follows the Riochican and precedes the Cas ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
** Genus '' Procladosictis''
***''Procladosictis anomala'' (late
Eocene
The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
,
Mustersan The Mustersan age is a period of geologic time (48.0–42.0 Ma) within the Eocene epoch of the Paleogene, used more specifically within the South American land mammal age (SALMA) classification. It follows the Casamayoran and precedes the Divisade ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
** Family
Hondadelphidae
''Hondadelphys'' is an extinct genus of carnivorous sparassodonts, known from the Middle Miocene of Colombia. The type species, ''H. fieldsi'', was described in 1976 from the fossil locality of La Venta, which hosts fossils from the Villavieja Fo ...
*** Genus ''
Hondadelphys
''Hondadelphys'' is an extinct genus of carnivorous sparassodonts, known from the Middle Miocene of Colombia. The type species, ''H. fieldsi'', was described in 1976 from the fossil locality of La Venta, which hosts fossils from the Villavieja For ...
''
****''Hondadelphys fieldsi'' (late middle
Miocene
The Miocene ( ) is the first 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 means "less recen ...
,
Laventan
The Laventan ( es, Laventense) 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 ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
*** Genus '' Stylocynus''
****''Stylocynus paranensis'' (latest
Miocene
The Miocene ( ) is the first 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 means "less recen ...
,
Huayquerian The Huayquerian ( es, Huayqueriense) age is a period of geologic time (9.0–6.8 Ma) within the Late Miocene epoch of the Neogene, used more specifically within the SALMA classification. It follows the Mayoan and precedes the Montehermosan age.
...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
** Family Hathliacynidae
*** Genus '' Acyon''
****''Acyon ?herrerae'' (early
Miocene
The Miocene ( ) is the first 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 means "less recen ...
,
Colhuehuapian The Colhuehuapian age is a period of geologic time (21.0–17.5 Ma) within the Early Miocene epoch of the Neogene, used more specifically within the SALMA classification in South America. It follows the Deseadan and precedes the Santacrucian ag ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
****''Acyon myctoderos'' (late middle
Miocene
The Miocene ( ) is the first 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 means "less recen ...
,
Laventan
The Laventan ( es, Laventense) 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 ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
****''Acyon tricuspidatus'' (late early
Miocene
The Miocene ( ) is the first 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 means "less recen ...
,
Santacrucian
The Santacrucian age is a period of geologic time (17.5 – 16.3 Ma) within the Early Miocene epoch of the Neogene, used more specifically with SALMA classification in South America. It follows the Colhuehuapian and precedes the Friasian age.
...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
*** Genus '' Australogale''
****''Australogale leptognathus'' (late middle
Miocene
The Miocene ( ) is the first 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 means "less recen ...
,
Laventan
The Laventan ( es, Laventense) 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 ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
*** Genus '' Borhyaenidium''
****''Borhyaenidium altiplanicus'' (latest
Miocene
The Miocene ( ) is the first 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 means "less recen ...
,
Huayquerian The Huayquerian ( es, Huayqueriense) age is a period of geologic time (9.0–6.8 Ma) within the Late Miocene epoch of the Neogene, used more specifically within the SALMA classification. It follows the Mayoan and precedes the Montehermosan age.
...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
****''Borhyaenidium riggsi'' (unknown, either early or late
Pliocene
The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58Montehermosan
The Montehermosan age is a period of geologic time (6.8–4.0 Ma) within the Miocene and Pliocene epochs of the Neogene used more specifically with South American Land Mammal Ages. It follows the Huayquerian and precedes the Chapadmalalan
The C ...
or
Chapadmalalan
The Chapadmalalan age is a period of geologic time (4.0–3.0 Ma) within the Pliocene epoch of the Neogene used more specifically with South American Land Mammal Ages. It follows the Montehermosan and precedes the Uquian age.
Fossil content
F ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
****''Borhyaenidium musteloides'' (latest
Miocene
The Miocene ( ) is the first 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 means "less recen ...
,
Huayquerian The Huayquerian ( es, Huayqueriense) age is a period of geologic time (9.0–6.8 Ma) within the Late Miocene epoch of the Neogene, used more specifically within the SALMA classification. It follows the Mayoan and precedes the Montehermosan age.
...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
*** Genus '' Chasicostylus''
****''Chasicostylus castroi'' (early late
Miocene
The Miocene ( ) is the first 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 means "less recen ...
,
Chasicoan The Chasicoan ( es, Chasiquense or es, Chasicoense) age is a period of geologic time from 10–9 Ma within the Late Miocene epoch of the Neogene, used more specifically within the SALMA classification in South America. It follows the Mayoan and ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
*** Genus ''
Cladosictis
''Cladosictis'' is an extinct genus of South American metatherian from Patagonia, Argentina ( Chichinales, Cerro Bandera, Sarmiento and Santa Cruz Formations) and Chile ( Río Frias Formation).Miocene
The Miocene ( ) is the first 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 means "less recen ...
,
Colhuehuapian The Colhuehuapian age is a period of geologic time (21.0–17.5 Ma) within the Early Miocene epoch of the Neogene, used more specifically within the SALMA classification in South America. It follows the Deseadan and precedes the Santacrucian ag ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
****''Cladosictis patagonica'' (late early to earliest middle
Miocene
The Miocene ( ) is the first 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 means "less recen ...
,
Santacrucian
The Santacrucian age is a period of geologic time (17.5 – 16.3 Ma) within the Early Miocene epoch of the Neogene, used more specifically with SALMA classification in South America. It follows the Colhuehuapian and precedes the Friasian age.
...
-
Friasian The Friasian age is a period of geologic time (16.3–15.5 Ma) within the Early Miocene epoch of the Neogene, used more specifically within the SALMA classification of South America. It follows the Santacrucian and precedes the Colloncuran age. ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
*** Genus '' Notictis''
****''Notictis ortizi'' (latest
Miocene
The Miocene ( ) is the first 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 means "less recen ...
,
Huayquerian The Huayquerian ( es, Huayqueriense) age is a period of geologic time (9.0–6.8 Ma) within the Late Miocene epoch of the Neogene, used more specifically within the SALMA classification. It follows the Mayoan and precedes the Montehermosan age.
...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
*** Genus '' Notocynus''
****''Notocynus hermosicus'' (early
Pliocene
The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58Montehermosan
The Montehermosan age is a period of geologic time (6.8–4.0 Ma) within the Miocene and Pliocene epochs of the Neogene used more specifically with South American Land Mammal Ages. It follows the Huayquerian and precedes the Chapadmalalan
The C ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
*** Genus '' Notogale''
****''Notogale mitis'' (late
Oligocene
The Oligocene ( ) is a geologic epoch of the Paleogene Period and extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that define the epoch are well identified but the ...
,
Deseadan The Deseadan ( es, Deseadense) age is a period of geologic time (29.0–21.0 Ma) within the Oligocene epoch of the Paleogene to the Early Miocene epoch of the Neogene, used more specifically within the SALMA classification of South America. It fo ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
*** Genus '' Pseudonotictis''
****''Pseudonotictis chubutensis'' (early middle
Miocene
The Miocene ( ) is the first 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 means "less recen ...
,
Colloncuran
The Colloncuran ( es, Colloncurense) age is a period of geologic time (15.5–13.8 Ma) within the Middle Miocene epoch of the Neogene, used more specifically within the SALMA classification in South America. It follows the Friasian and precedes ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
****''Pseudonotictis pusillus'' (late early
Miocene
The Miocene ( ) is the first 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 means "less recen ...
,
Santacrucian
The Santacrucian age is a period of geologic time (17.5 – 16.3 Ma) within the Early Miocene epoch of the Neogene, used more specifically with SALMA classification in South America. It follows the Colhuehuapian and precedes the Friasian age.
...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
*** Genus '' Perathereutes''
****''Perathereutes pungens'' (late early
Miocene
The Miocene ( ) is the first 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 means "less recen ...
,
Santacrucian
The Santacrucian age is a period of geologic time (17.5 – 16.3 Ma) within the Early Miocene epoch of the Neogene, used more specifically with SALMA classification in South America. It follows the Colhuehuapian and precedes the Friasian age.
...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
*** Genus '' Sallacyon''
****''Sallacyon hoffstetteri'' (late
Oligocene
The Oligocene ( ) is a geologic epoch of the Paleogene Period and extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that define the epoch are well identified but the ...
,
Deseadan The Deseadan ( es, Deseadense) age is a period of geologic time (29.0–21.0 Ma) within the Oligocene epoch of the Paleogene to the Early Miocene epoch of the Neogene, used more specifically within the SALMA classification of South America. It fo ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
*** Genus '' Sipalocyon''
****''Sipalocyon externus'' (early
Miocene
The Miocene ( ) is the first 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 means "less recen ...
,
Colhuehuapian The Colhuehuapian age is a period of geologic time (21.0–17.5 Ma) within the Early Miocene epoch of the Neogene, used more specifically within the SALMA classification in South America. It follows the Deseadan and precedes the Santacrucian ag ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
****''Sipalocyon gracilis'' (late early
Miocene
The Miocene ( ) is the first 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 means "less recen ...
,
Santacrucian
The Santacrucian age is a period of geologic time (17.5 – 16.3 Ma) within the Early Miocene epoch of the Neogene, used more specifically with SALMA classification in South America. It follows the Colhuehuapian and precedes the Friasian age.
...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
****''Sipalocyon "obusta"'' (late early
Miocene
The Miocene ( ) is the first 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 means "less recen ...
,
Santacrucian
The Santacrucian age is a period of geologic time (17.5 – 16.3 Ma) within the Early Miocene epoch of the Neogene, used more specifically with SALMA classification in South America. It follows the Colhuehuapian and precedes the Friasian age.
...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
** Superfamily
Borhyaenoidea
Sparassodonta (from Greek to tear, rend; and , gen. , ' tooth) is an extinct order of carnivorous metatherian mammals native to South America, related to modern marsupials. They were once considered to be true marsupials, but are now th ...
*** Genus '' Angelocabrerus''
****''Angelocabrerus daptes'' (middle
Eocene
The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
)
*** Genus '' Chlorocyon''
****''Chlorocyon phantasma'' (late
Eocene
The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
,
Mustersan The Mustersan age is a period of geologic time (48.0–42.0 Ma) within the Eocene epoch of the Paleogene, used more specifically within the South American land mammal age (SALMA) classification. It follows the Casamayoran and precedes the Divisade ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
*** Genus '' Dukecynus''
****''Dukecynus magnus'' (late middle
Miocene
The Miocene ( ) is the first 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 means "less recen ...
,
Laventan
The Laventan ( es, Laventense) 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 ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
*** Genus '' Fredszalaya''
****''Fredszalaya hunteri'' (late
Oligocene
The Oligocene ( ) is a geologic epoch of the Paleogene Period and extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that define the epoch are well identified but the ...
,
Deseadan The Deseadan ( es, Deseadense) age is a period of geologic time (29.0–21.0 Ma) within the Oligocene epoch of the Paleogene to the Early Miocene epoch of the Neogene, used more specifically within the SALMA classification of South America. It fo ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
*** Genus ''
Lycopsis
''Lycopsis'' is an extinct genus of South American metatherian, that lived during the Miocene in Argentina and Colombia.
History
Although not named until 1927, Florentino Ameghino described a species now seen as synonymous with ''Lycopsis tor ...
''
****''Lycopsis longirostrus'' (late middle
Miocene
The Miocene ( ) is the first 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 means "less recen ...
,
Laventan
The Laventan ( es, Laventense) 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 ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
****''Lycopsis padillai'' (middle
Miocene
The Miocene ( ) is the first 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 means "less recen ...
,
Colloncuran
The Colloncuran ( es, Colloncurense) age is a period of geologic time (15.5–13.8 Ma) within the Middle Miocene epoch of the Neogene, used more specifically within the SALMA classification in South America. It follows the Friasian and precedes ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
****''Lycopsis torresi'' (early
Miocene
The Miocene ( ) is the first 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 means "less recen ...
,
Colhuehuapian The Colhuehuapian age is a period of geologic time (21.0–17.5 Ma) within the Early Miocene epoch of the Neogene, used more specifically within the SALMA classification in South America. It follows the Deseadan and precedes the Santacrucian ag ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
****''Lycopsis viverensis'' (early late
Miocene
The Miocene ( ) is the first 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 means "less recen ...
,
Chasicoan The Chasicoan ( es, Chasiquense or es, Chasicoense) age is a period of geologic time from 10–9 Ma within the Late Miocene epoch of the Neogene, used more specifically within the SALMA classification in South America. It follows the Mayoan and ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
))
*** Genus ''
Pharsophorus
''Pharsophorus'' is an extinct genus of borhyaenoid sparassodont that inhabited South America during the Middle to Late Oligocene epoch.
''
****''Pharsophorus lacerans'' (middle to late
Oligocene
The Oligocene ( ) is a geologic epoch of the Paleogene Period and extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that define the epoch are well identified but the ...
, La Cantera Local Fauna to
Deseadan The Deseadan ( es, Deseadense) age is a period of geologic time (29.0–21.0 Ma) within the Oligocene epoch of the Paleogene to the Early Miocene epoch of the Neogene, used more specifically within the SALMA classification of South America. It fo ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
****''Pharsophorus tenax'' (late
Oligocene
The Oligocene ( ) is a geologic epoch of the Paleogene Period and extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that define the epoch are well identified but the ...
,
Deseadan The Deseadan ( es, Deseadense) age is a period of geologic time (29.0–21.0 Ma) within the Oligocene epoch of the Paleogene to the Early Miocene epoch of the Neogene, used more specifically within the SALMA classification of South America. It fo ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
*** Genus '' Plesiofelis''
****''Plesiofelis schlosseri'' (late
Eocene
The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
,
Mustersan The Mustersan age is a period of geologic time (48.0–42.0 Ma) within the Eocene epoch of the Paleogene, used more specifically within the South American land mammal age (SALMA) classification. It follows the Casamayoran and precedes the Divisade ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
*** Genus '' Prothylacynus''
****''Prothylacynus patagonicus'' (late early to earliest middle
Miocene
The Miocene ( ) is the first 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 means "less recen ...
,
Santacrucian
The Santacrucian age is a period of geologic time (17.5 – 16.3 Ma) within the Early Miocene epoch of the Neogene, used more specifically with SALMA classification in South America. It follows the Colhuehuapian and precedes the Friasian age.
...
-
Friasian The Friasian age is a period of geologic time (16.3–15.5 Ma) within the Early Miocene epoch of the Neogene, used more specifically within the SALMA classification of South America. It follows the Santacrucian and precedes the Colloncuran age. ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
*** Genus '' Pseudolycopsis''
****''Pseudolycopsis cabrerai'' (early late
Miocene
The Miocene ( ) is the first 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 means "less recen ...
,
Chasicoan The Chasicoan ( es, Chasiquense or es, Chasicoense) age is a period of geologic time from 10–9 Ma within the Late Miocene epoch of the Neogene, used more specifically within the SALMA classification in South America. It follows the Mayoan and ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
*** Genus '' Pseudothylacynus''
****''Pseudothylacynus rectus'' (early
Miocene
The Miocene ( ) is the first 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 means "less recen ...
,
Colhuehuapian The Colhuehuapian age is a period of geologic time (21.0–17.5 Ma) within the Early Miocene epoch of the Neogene, used more specifically within the SALMA classification in South America. It follows the Deseadan and precedes the Santacrucian ag ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
*** Family
Borhyaenidae
Borhyaenidae is an extinct metatherian family of low-slung, heavily built predatory mammals in the order Sparassodonta. Borhyaenids are not true marsupials, but members of a sister taxon, Sparassodonta. Like most metatherians, borhyaenids and o ...
**** Genus '' Acrocyon''
*****''Acrocyon riggsi'' (early
Miocene
The Miocene ( ) is the first 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 means "less recen ...
,
Colhuehuapian The Colhuehuapian age is a period of geologic time (21.0–17.5 Ma) within the Early Miocene epoch of the Neogene, used more specifically within the SALMA classification in South America. It follows the Deseadan and precedes the Santacrucian ag ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
*****''Acrocyon sectorius'' (late early
Miocene
The Miocene ( ) is the first 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 means "less recen ...
,
Santacrucian
The Santacrucian age is a period of geologic time (17.5 – 16.3 Ma) within the Early Miocene epoch of the Neogene, used more specifically with SALMA classification in South America. It follows the Colhuehuapian and precedes the Friasian age.
...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
**** Genus '' Arctodictis''
*****''Arctodictis munizi'' (late early
Miocene
The Miocene ( ) is the first 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 means "less recen ...
,
Santacrucian
The Santacrucian age is a period of geologic time (17.5 – 16.3 Ma) within the Early Miocene epoch of the Neogene, used more specifically with SALMA classification in South America. It follows the Colhuehuapian and precedes the Friasian age.
...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
*****''Arctodictis sinclairi'' (early
Miocene
The Miocene ( ) is the first 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 means "less recen ...
,
Colhuehuapian The Colhuehuapian age is a period of geologic time (21.0–17.5 Ma) within the Early Miocene epoch of the Neogene, used more specifically within the SALMA classification in South America. It follows the Deseadan and precedes the Santacrucian ag ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
**** Genus ''
Australohyaena
''Australohyaena'' is an extinct genus of carnivorous mammal, belonging to the order Sparassodonta. It lived during the Late Oligocene, and its fossilized remains were discovered in Argentina.
Description
This genus is mainly known from cran ...
''
*****''Australohyaena antiquua'' (late
Oligocene
The Oligocene ( ) is a geologic epoch of the Paleogene Period and extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that define the epoch are well identified but the ...
,
Deseadan The Deseadan ( es, Deseadense) age is a period of geologic time (29.0–21.0 Ma) within the Oligocene epoch of the Paleogene to the Early Miocene epoch of the Neogene, used more specifically within the SALMA classification of South America. It fo ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
**** Genus ''
Borhyaena
''Borhyaena'' is an extinct genus of South American metatherian, living between 17.5 and 15.5 million years ago in Patagonia, Argentina ( Santa Cruz and Sarmiento Formations) and Chile ( Río Frias Formation).Miocene
The Miocene ( ) is the first 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 means "less recen ...
,
Colhuehuapian The Colhuehuapian age is a period of geologic time (21.0–17.5 Ma) within the Early Miocene epoch of the Neogene, used more specifically within the SALMA classification in South America. It follows the Deseadan and precedes the Santacrucian ag ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
*****''Borhyaena tuberata'' (late early
Miocene
The Miocene ( ) is the first 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 means "less recen ...
,
Santacrucian
The Santacrucian age is a period of geologic time (17.5 – 16.3 Ma) within the Early Miocene epoch of the Neogene, used more specifically with SALMA classification in South America. It follows the Colhuehuapian and precedes the Friasian age.
...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
Proborhyaenidae
Proborhyaenidae is an extinct family of metatherian mammals of the order Sparassodonta, which lived in South America from the Eocene (Mustersan) until the Oligocene (Deseadan). Sometimes it has been included as a subfamily of their relatives, the ...
Eocene
The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
**** Genus '' Paraborhyaena''
*****''Paraborhyaena boliviana'' (late
Oligocene
The Oligocene ( ) is a geologic epoch of the Paleogene Period and extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that define the epoch are well identified but the ...
,
Deseadan The Deseadan ( es, Deseadense) age is a period of geologic time (29.0–21.0 Ma) within the Oligocene epoch of the Paleogene to the Early Miocene epoch of the Neogene, used more specifically within the SALMA classification of South America. It fo ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
**** Genus '' Proborhyaena''
*****''Proborhyaena gigantea'' (late
Oligocene
The Oligocene ( ) is a geologic epoch of the Paleogene Period and extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that define the epoch are well identified but the ...
,
Deseadan The Deseadan ( es, Deseadense) age is a period of geologic time (29.0–21.0 Ma) within the Oligocene epoch of the Paleogene to the Early Miocene epoch of the Neogene, used more specifically within the SALMA classification of South America. It fo ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
Eomakhaira
''Eomakhaira'' is an extinct genus of thylacosmilid sparassodont known from the Oligocene Abanico Formation of Chile
Chile, officially the Republic of Chile, is a country in the western part of South America. It is the southernmost count ...
''
******''Eomakhaira molossus'' (early
Oligocene
The Oligocene ( ) is a geologic epoch of the Paleogene Period and extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that define the epoch are well identified but the ...
, ?
Tinguirirican The Tinguirirican ( es, Tinguiririquense) age is a period of geologic time (36.0–29.0 Ma) within the Late Eocene and Early Oligocene epochs of the Paleogene, used more specifically within the SALMA classification in South America. It follows th ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
***** Genus ''
Anachlysictis
''Anachlysictis gracilis'' is an extinct carnivorous mammal belonging to the group Sparassodonta, which were metatherians (a group including marsupials and their close relatives) that inhabited South America during the Cenozoic. ''Anachlysictis' ...
''
******''Anachlysictis gracilis'' (late middle
Miocene
The Miocene ( ) is the first 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 means "less recen ...
,
Laventan
The Laventan ( es, Laventense) 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 ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
***** Genus ''
Patagosmilus
''Patagosmilus'' ("Patagonian knife" in Greek) is an extinct genus of meat-eating metatherian mammal of the family Thylacosmilidae, that lived in the Middle Miocene in South America.Miocene
The Miocene ( ) is the first 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 means "less recen ...
,
Colloncuran
The Colloncuran ( es, Colloncurense) age is a period of geologic time (15.5–13.8 Ma) within the Middle Miocene epoch of the Neogene, used more specifically within the SALMA classification in South America. It follows the Friasian and precedes ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
***** Genus ''
Thylacosmilus
''Thylacosmilus'' is an extinct genus of Saber-toothed predator, saber-toothed metatherian mammals that inhabited South America from the Miocene, Late Miocene to Pliocene Epoch (geology), epochs. Though ''Thylacosmilus'' looks similar to the "Mac ...
''
******''Thylacosmilus atrox'' (latest
Miocene
The Miocene ( ) is the first 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 means "less recen ...
to late
Pliocene
The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58Huayquerian The Huayquerian ( es, Huayqueriense) age is a period of geologic time (9.0–6.8 Ma) within the Late Miocene epoch of the Neogene, used more specifically within the SALMA classification. It follows the Mayoan and precedes the Montehermosan age.
...
-
Chapadmalalan
The Chapadmalalan age is a period of geologic time (4.0–3.0 Ma) within the Pliocene epoch of the Neogene used more specifically with South American Land Mammal Ages. It follows the Montehermosan and precedes the Uquian age.
Fossil content
F ...
SALMA
The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as a ...
)
Several other metatherian taxa have been suggested to be sparassodonts or closely related to sparassodonts. The australian
Murgon
Murgon is a rural town and locality in the South Burnett Region, Queensland, Australia. In the , the locality of Murgon had a population of 2,378 people.
Geography
Murgon is in the region of Queensland known as the South Burnett, the southern ...
taxa '' Archaeonothos'' has been noted as being similar to sparassodonts, but currently its relationships are not fully concluded. Carneiro (2018) recovered the genus '' Varalphadon'' from the
Late Cretaceous
The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', the ...
of North America as a basal member of Sparassodonta. However, this interpretation of ''Varalphadon'' as a sparassodont has not been supported by later phylogenetic analyses, and most of the purported synapomorphies between ''Varalphadon'' and sparassodonts are not actually present in ''Varalphadon'' or have been suggested to be due to convergent evolution. Sparassodonts have sometimes been considered closely related to the "
Gurlin Tsav skull The "Gurlin Tsav" skull is a currently unnamed carnivorous metatherian fossil from the Nemegt Formation of Mongolia. Composed of a single semi-complete skull, this specimen is notable in regards to the evolution and systematics of Metatheria as a wh ...
" an unnamed metatherian known from a partial skull found in the Late Cretaceous
Nemegt Formation
The Nemegt Formation (also known as Nemegtskaya Svita) is a geological formation in the Gobi Desert of Mongolia, dating to the Late Cretaceous. The formation consists of river channel sediments and contains fossils of fish, turtles, crocodilian ...
of Mongolia.
The following
cladogram
A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...
of sparassodont interrelationships is after Engelman ''et al.'', 2020. Not all studies agree on the
sister group
In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree.
Definition
The expression is most easily illustrated by a cladogram:
Taxon A and t ...
relationship between
Thylacosmilidae
Thylacosmilidae is an extinct Family (biology), family of metatherian predators, related to the modern marsupials, which lived in South America between the Miocene and Pliocene epochs. Like other South American mammalian predators that lived prio ...
and
Borhyaenidae
Borhyaenidae is an extinct metatherian family of low-slung, heavily built predatory mammals in the order Sparassodonta. Borhyaenids are not true marsupials, but members of a sister taxon, Sparassodonta. Like most metatherians, borhyaenids and o ...
recovered here, with other studies finding thylacosmilids to be within
Proborhyaenidae
Proborhyaenidae is an extinct family of metatherian mammals of the order Sparassodonta, which lived in South America from the Eocene (Mustersan) until the Oligocene (Deseadan). Sometimes it has been included as a subfamily of their relatives, the ...
. The relationships among hathliacynids are also relatively unstable.
Within
Metatheria
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 ...
, a 2016 phylogenetic analysis group found that borhyaenids form a clade with the Asian "Gurlin Tsav skull" as well as other South American taxa. The same phylogeny found that marsupials group among various North American Cretaceous species. The phylogenetic tree is reproduced below.
Evolution
The early history of the Sparassodonta is poorly known, as most
Paleocene
The Paleocene, ( ) or Palaeocene, is a geological epoch (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), E ...
and
Eocene
The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
members of this group are only known from isolated teeth and fragmentary jaws. However, one species, the middle
Eocene
The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
''Callistoe vincei'', is known from a nearly complete, articulated skeleton. As ''Callistoe'' belongs to one of the most specialized groups of sparassodonts, this indicates that the other major groups (e.g. borhyaenids, hathliacynids, etc.) must have also arisen by this time. Originally, the early
Paleocene
The Paleocene, ( ) or Palaeocene, is a geological epoch (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), E ...
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 ...
''
Mayulestes
''Mayulestes'' (Quechua: ''mayu'' river, + Greek: ''lestes'', thief) is a genus of carnivorous metatherian that lived in what is now Tiupampa, Bolivia in the early Paleocene. It shared its habitat with fellow sparrasodont ''Pucadelphys'', and a ...
'' was considered to be the earliest known member of the Sparassodonta, but phylogenetic analyses suggest that this species represents an independent radiation of carnivorous
metatheria
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 ...
ns more closely related to ''
Pucadelphys
''Pucadelphys'' is an extinct genus of non-marsupial metatherian. The genus contains a single species, ''P. andinus''. Fossils of ''Pucadelphys'' have been found in the Santa Lucía Formation in Tiupampa in Bolivia.
''; however, recent studies show that these taxa were closely related to borhyaenids. As of this writing, the earliest known true sparassodonts are either ''Allqokirus australis'', a species from the same site as ''Mayulestes'' that may turn out to not be a sparassodont, and an isolated
astragalus
''Astragalus'' is a large genus of over 3,000 species of herbs and small shrubs, belonging to the legume family Fabaceae and the subfamily Faboideae. It is the largest genus of plants in terms of described species. The genus is native to tempe ...
from the earliest
Paleocene
The Paleocene, ( ) or Palaeocene, is a geological epoch (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), E ...
site of Punta Peligro,
Argentina
Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the second-largest country in South America after Brazil, th ...
.
Sparassodonts have been suggested to be related to a variety of other groups of metatherians.
Florentino Ameghino
Florentino Ameghino (born Giovanni Battista Fiorino Giuseppe Ameghino September 19, 1853 – August 6, 1911) was an Argentine naturalist, paleontologist, anthropologist and zoologist, whose fossil discoveries on the Argentine Pampas, especially ...
, who first described fossils of the group, thought that sparassodonts were closely related to
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 ance ...
and were a transitional group between metatherians and carnivorous placentals (including modern
carnivorans
Carnivora is a monophyletic order of placental mammals consisting of the most recent common ancestor of all cat-like and dog-like animals, and all descendants of that ancestor. Members of this group are formally referred to as carnivorans, ...
). Contemporary authors in the late 19th and early 20th century rejected this hypothesis and considered sparassodonts to be closely related to Australian
thylacines
The thylacine ( , or , also ) (''Thylacinus cynocephalus'') is an extinct carnivorous marsupial that was native to the Australian mainland and the islands of Tasmania and New Guinea. The last known live animal was captured in 1930 in Tasma ...
and dasyurids. The most popular hypothesis for much of the 20th century was that sparassodonts were closely related to
opossums
Opossums () are members of the marsupial order Didelphimorphia () endemic to the Americas. The largest order of marsupials in the Western Hemisphere, it comprises 93 species in 18 genera. Opossums originated in South America and entered North A ...
. In 1990, Marshall et al. (1990) considered the
Cretaceous
The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of th ...
stagodontids to be members of Sparassodonta, but this was criticized by later authors. Marshall and Kielan-Jaworowska (1992) considered sparassodonts to be closely related to
deltatheroida
Deltatheroida is an extinct group of basal metatherians that were distantly related to modern marsupials. The majority of known members of the group lived in the Cretaceous; one species, '' Gurbanodelta kara'', is known from the late Paleocene (G ...
ns, but this was also criticized. Most of these hypotheses were based on similar adaptations for
carnivorous
A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, fat and other sof ...
diets in sparassodonts, opossums, dasyuromorphians, stagodonts, and deltatheroidans, which are highly prone to convergent evolution within mammals. Szalay (1994) considered sparassodonts to be closely related to
paucituberculata
Paucituberculata is an order of South American marsupials. Although currently represented only by the seven living species of shrew opossums, this order was formerly much more diverse, with more than 60 extinct species named from the fossil rec ...
ns based on features of the ankle. In recent years there has been a growing consensus that sparassodonts are positioned just outside of
crown-group
In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor. ...
Marsupialia
Marsupials are any members of the mammalian infraclass Marsupialia. All extant marsupials are endemic to Australasia, Wallacea and the Americas. A distinctive characteristic common to most of these species is that the young are carried in a pou ...
, in a broader clade ( Pucadelphyida) including pucadelphyids as well as sparassodonts.
Sparassodonts are currently considered to be
endemic
Endemism is the state of a species being found 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 found elsew ...
to South America, and have not even been found in nearby continents like
Antarctica
Antarctica () is Earth's southernmost and least-populated continent. Situated almost entirely south of the Antarctic Circle and surrounded by the Southern Ocean, it contains the geographic South Pole. Antarctica is the fifth-largest contine ...
(which otherwise shares many groups of mammals with South America such as
litopterns
Litopterna (from grc, λῑτή πτέρνα "smooth heel") is an extinction, extinct order of fossil hoofed mammals from the Cenozoic era. The order is one of the five great orders of Meridiungulata, South American ungulates that were endemic to ...
,
astrapotheres
Astrapotheria is an extinct order of South American and Antarctic hoofed mammals that existed from the late Paleocene to the Middle Miocene, ."The uruguaytheriine Astrapotheriidae from the rich middle Miocene Honda Group of the upper Magdalen ...
Almost all sparassodonts have an exceptionally shortened snout—most especially thylacosmylids. Hathliacynids usually have a longer snout than the other groups. The
nasal bone
The nasal bones are two small oblong bones, varying in size and form in different individuals; they are placed side by side at the middle and upper part of the face and by their junction, form the bridge of the upper one third of the nose.
Eac ...
s extend past the
eye sockets
In anatomy, the orbit is the cavity or socket of the skull in which the eye and its appendages are situated. "Orbit" can refer to the bony socket, or it can also be used to imply the contents. In the adult human, the volume of the orbit is , o ...
, often reaching the
lacrimal bone
The lacrimal bone is a small and fragile bone of the facial skeleton; it is roughly the size of the little fingernail. It is situated at the front part of the medial wall of the orbit. It has two surfaces and four borders. Several bony landmarks of ...
. Except for thylacosmylids beyond ''
Patagosmilus
''Patagosmilus'' ("Patagonian knife" in Greek) is an extinct genus of meat-eating metatherian mammal of the family Thylacosmilidae, that lived in the Middle Miocene in South America.postorbital process
The postorbital process is a projection on the frontal bone near the rear upper edge of the eye socket. In many mammals, it reaches down to the zygomatic arch, forming the postorbital bar.
References
See also
* Orbital process
In the human ...
es which would otherwise form the
postorbital bar The postorbital bar (or postorbital bone) is a bony arched structure that connects the frontal bone of the skull to the zygomatic arch, which runs laterally around the eye socket. It is a trait that only occurs in mammalian taxa, such as most strep ...
connecting the forehead to the cheek, thus framing the eye. They exhibit marked
postorbital constriction
In physical anthropology, post-orbital constriction is the narrowing of the cranium (skull) just behind the eye sockets (the orbits, hence the name) found in most non-human primates and early hominins. This constriction is very noticeable in non-h ...
. The
orbital process
In the human skull, the zygomatic bone (from grc, ζῠγόν, zugón, yoke), also called cheekbone or malar bone, is a paired irregular bone which articulates with the maxilla, the temporal bone, the sphenoid bone and the frontal bone. It is si ...
(between the cheek and the eye socket) is usually diminished, though the
zygomatic arch
In anatomy, the zygomatic arch, or cheek bone, is a part of the skull formed by the zygomatic process of the temporal bone (a bone extending forward from the side of the skull, over the opening of the ear) and the temporal process of the zygomati ...
(the cheekbone) is strong. They feature a prominent
sagittal crest
A sagittal crest is a ridge of bone running lengthwise along the midline of the top of the skull (at the sagittal suture) of many mammalian and reptilian skulls, among others. The presence of this ridge of bone indicates that there are exceptiona ...
along the midline of the flattened skull, the crest strength is quite variable among borhyaenids. They have an expanded
occipital bone
The occipital bone () is a neurocranium, cranial dermal bone and the main bone of the occiput (back and lower part of the skull). It is trapezoidal in shape and curved on itself like a shallow dish. The occipital bone overlies the occipital lobe ...
with a well defined
nuchal crest
The nuchal lines are four curved lines on the external surface of the occipital bone:
* The upper, often faintly marked, is named the highest nuchal line, but is sometimes referred to as the Mempin line or linea suprema, and it attaches to the epi ...
.
Sparassodonts spanned a wide range of body sizes, from 2.2-pound (1 kg)
weasel
Weasels are mammals of the genus ''Mustela'' of the family Mustelidae. The genus ''Mustela'' includes the least weasels, polecats, stoats, ferrets and European mink. Members of this genus are small, active predators, with long and slender bo ...
or
civet
A civet () is a small, lean, mostly nocturnal mammal native to tropical Asia and Africa, especially the tropical forests. The term civet applies to over a dozen different species, mostly from the family Viverridae. Most of the species diversity ...
-like forms to ''
Thylacosmilus
''Thylacosmilus'' is an extinct genus of Saber-toothed predator, saber-toothed metatherian mammals that inhabited South America from the Miocene, Late Miocene to Pliocene Epoch (geology), epochs. Though ''Thylacosmilus'' looks similar to the "Mac ...
'', which was the size of a
leopard
The leopard (''Panthera pardus'') is one of the five extant species in the genus '' Panthera'', a member of the cat family, Felidae. It occurs in a wide range in sub-Saharan Africa, in some parts of Western and Central Asia, Southern Russia, a ...
. Along with the Australian thylacoleonids, sparassodonts include some of the largest metatherian carnivores.
Sparassodonts have highly reduced
epipubic
Epipubic bones are a pair of bones projecting forward from the pelvic bones of modern marsupials, monotremes and fossil mammals like multituberculates, and even basal eutherians (the ancestors of placental mammals, who lack them).
They first occ ...
bones (pelvic bones which support the pouch), to the point that early analysis could not even find evidence for them. This is a characteristic shared with the Australian
thylacine
The thylacine ( , or , also ) (''Thylacinus cynocephalus'') is an extinct carnivorous marsupial that was native to the Australian mainland and the islands of Tasmania and New Guinea. The last known live animal was captured in 1930 in Tasman ...
, and historically argued as a
synapomorphy
In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to have ...
, though nowadays it is considered to have developed independently for poorly understood reasons. Like with thylacines, it is very likely that they possessed long cartilaginous elements instead.
Teeth
The
dental formula
Dentition pertains to the development of teeth and their arrangement in the mouth. In particular, it is the characteristic arrangement, kind, and number of teeth in a given species at a given age. That is, the number, type, and morpho-physiolo ...
of sparassodonts varies considerably. In borhyaenids, it is , with three upper and lower
incisor
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, whe ...
s, one upper and lower canine, three upper and lower
premolar
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 mouth ...
s, and four upper and lower
molars
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 to ...
in each half of either jaw. Proborhyaenids usually only have two lower incisors instead of three, except for '' Callistoe''. Thylacosmylids have at least two upper and only two lower incisors (the uppers grew into elongated
sabers
A sabre (French: sabʁ or saber in American English) is a type of backsword with a curved blade associated with the light cavalry of the early modern and Napoleonic periods. Originally associated with Central European cavalry such as the ...
), and two upper and lower premolars. Some specimens of ''
Borhyaena
''Borhyaena'' is an extinct genus of South American metatherian, living between 17.5 and 15.5 million years ago in Patagonia, Argentina ( Santa Cruz and Sarmiento Formations) and Chile ( Río Frias Formation).Arctodictis'' are also missing the last upper molar, showing that the presence of this tooth was variable in these species.
Sparassodonta is characterized by dental
synapomorphies
In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to have ...
that distinguish the group from other closely related mammals. Unequivocal traits uniting the earliest Sparassodonts include:
* a snout that forms a pronounced bulge around the canine teeth when viewed from above
* a ridge on the upper molar (preparacrista) oriented anterobuccally (towards the cheek) with respect to the long axis of the tooth.
* a pronounced keel near the base of the front of the paraconid
* ridges on lower molars (postprotocristid-metacristid) parallel or oblique with respect to lower jaw axis.
* a very tall protoconid (>90% tooth length, secondarily lost in ''Hondadelphys'' and ''Stylocynus'') that bulges to the side and is wider at its midpoint than its base
* talonid (crushing end) of lower molar narrow in relation to trigonid (shearing end).
In borhyaenids, only the third premolar was ever replaced in the animal's lifetime, similar to other metatherians. In thylacosmilids, only the lower third premolar was replaced.
The cusps of the sparassodont molar correlate to a cutting function rather than a crushing one. In the upper molars, the paracone (on the lingual, or tongueward, side) is reduced and fused to the metacone (distal, towards the back of the mouth), inflating the postmetacrista (the lingual border of the metacone); and they almost always lack the stylar shelf (on the buccal, or cheekward, side) and associated stylar cusps. In the lower molars, the trigonids (the buccal shearing side) have an inflated paracristid and marginalized or absent metaconid; and the talonid (the distal, or backendwards, crushing side) is either reduced or gone.
Paleobiology
Diet
Sparassodonts were carnivorous, and with the exception of some basal members of all members of this group were
hypercarnivorous
A hypercarnivore is an animal which has a diet that is more than 70% meat, either via active predation or by scavenging. The remaining non-meat diet may consist of non-animal foods such as fungi, fruits or other plant material. Some extant exampl ...
(having diets composed of more than 70% meat). Only ''Hondadelphys'' and ''Stylocynus'' appear to have exhibited adaptations for
omnivory
An omnivore () is an animal that has the ability to eat and survive on both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize the nut ...
, and even then ''Stylocynus'' may have had a more mesocarnivorous diet similar to
canids
Canidae (; from Latin, ''canis'', "dog") is a biological family of dog-like carnivorans, colloquially referred to as dogs, and constitutes a clade. A member of this family is also called a canid (). There are three subfamilies found within th ...
than an omnivorous one. Medium-to-large
caviomorph
Caviomorpha is the rodent infraorder or parvorder that unites all New World hystricognaths. It is supported by both fossil and molecular evidence. The Caviomorpha was for a time considered to be a separate order outside the Rodentia, but is now a ...
rodents
Rodents (from Latin , 'to gnaw') are mammals of the order Rodentia (), which are characterized by a single pair of continuously growing incisors in each of the upper and lower jaws. About 40% of all mammal species are rodents. They are nat ...
and rodent-like mammals (e.g., small
notoungulates
Notoungulata is an extinct order of mammalian ungulates that inhabited South America from the early Paleocene to the Holocene, living from approximately 61 million to 11,000 years ago. Notoungulates were morphologically diverse, with forms resemb ...
) appear to have been common prey items of sparassodonts. The subadult holotype of '' Lycopsis longirostrus'' preserves remains of the dinomyid ''Scleromys colombianus'' as fossilized gut contents. Sparassodont
coprolites
A coprolite (also known as a coprolith) is fossilized feces. Coprolites are classified as trace fossils as opposed to body fossils, as they give evidence for the animal's behaviour (in this case, diet) rather than morphology. The name is de ...
from the
Santa Cruz Formation
The Santa Cruz Formation is a geological formation in the Magallanes/Austral Basin in southern Patagonia in Argentina and in adjacent areas of Chile. It dates to the late Early Miocene epoch, and is contemporaneous with eponymous Santacrucian ...
preserve the bones of chinchillid and octodontoid rodents inside them. Bite marks from medium-sized sparassodonts have been found on the small
notoungulate
Notoungulata is an extinct order of mammalian ungulates that inhabited South America from the early Paleocene to the Holocene, living from approximately 61 million to 11,000 years ago. Notoungulates were morphologically diverse, with forms resemb ...
'' Paedotherium''. Stable isotope data from the early late Miocene ''
Lycopsis
''Lycopsis'' is an extinct genus of South American metatherian, that lived during the Miocene in Argentina and Colombia.
History
Although not named until 1927, Florentino Ameghino described a species now seen as synonymous with ''Lycopsis tor ...
viverensis'' and ''
Thylacosmilus
''Thylacosmilus'' is an extinct genus of Saber-toothed predator, saber-toothed metatherian mammals that inhabited South America from the Miocene, Late Miocene to Pliocene Epoch (geology), epochs. Though ''Thylacosmilus'' looks similar to the "Mac ...
atrox'' suggests that these species fed on C3grazers in open habitats, likely notoungulates.
Bite marks likely pertaining to hathliacynid sparassodonts have been found on the remains of penguins and flightless marine ducks in ancient seabird nesting colonies, suggesting that sparassodonts raided seabird colonies for eggs, carrion, and other prey like many predatory mammals do today.
Borhyaenid and proborhyaenid sparassodonts have been interpreted as being capable of crushing bones similar to modern
hyenas
Hyenas, or hyaenas (from Ancient Greek , ), are feliform carnivoran mammals of the family Hyaenidae . With only four extant species (each in its own genus), it is the fifth-smallest family in the Carnivora and one of the smallest in the cla ...
,
wolverines
The wolverine (), (''Gulo gulo''; ''Gulo'' is Latin for " glutton"), also referred to as the glutton, carcajou, or quickhatch (from East Cree, ''kwiihkwahaacheew''), is the largest land-dwelling species of the family Mustelidae. It is a muscul ...
, or the
Tasmanian devil
The Tasmanian devil (''Sarcophilus harrisii'') (palawa kani: purinina) is a carnivorous marsupial of the family Dasyuridae. Until recently, it was only found on the island state of Tasmania, but it has been reintroduced to New South Wales in ...
(''Sarcophilus harrisii'') based on their deep jaws, bulbous premolars with deep roots and pronounced wear at their tips, extensive fused or interlocking mandibular symphyses, large
masseteric fossa
Masseteric is an adjective meaning "of or pertaining to the Masseter muscle", such as:
* Masseteric artery
The masseteric artery is small and passes laterally through the mandibular notch to the deep surface of the masseter muscle, which it suppl ...
e, microfractures in their tooth enamel, and high estimated bite forces. ''Australohyaena antiquua'' shows particularly pronounced adaptations for bone-cracking, with a very deep jaw and strongly arched
nasals
In phonetics, a nasal, also called a nasal occlusive or nasal stop in contrast with an oral stop or nasalized consonant, is an occlusive consonant produced with a lowered velum, allowing air to escape freely through the nose. The vast majorit ...
similar to what is seen in modern
hyaenids
Hyenas, or hyaenas (from Ancient Greek , ), are feliform carnivoran mammals of the family Hyaenidae . With only four extant species (each in its own genus), it is the fifth-smallest family in the Carnivora and one of the smallest in the cla ...
.
Based on studies of the
postcranial skeleton Postcrania (postcranium, adjective: postcranial) in zoology and vertebrate paleontology is all or part of the skeleton apart from the skull. Frequently, fossil remains, e.g. of dinosaurs or other extinct tetrapods, consist of partial or isolated sk ...
, it appears as though most sparassodonts were scansorial (adapted for climbing), although terrestrial adaptations evolved in ''
Lycopsis
''Lycopsis'' is an extinct genus of South American metatherian, that lived during the Miocene in Argentina and Colombia.
History
Although not named until 1927, Florentino Ameghino described a species now seen as synonymous with ''Lycopsis tor ...
longirostrus'', borhyaenids, proborhyaenids, and thylacosmilids. Most sparassodonts were
plantigrade
151px, Portion of a human skeleton, showing plantigrade habit
In terrestrial animals, plantigrade locomotion means walking with the toes and metatarsals flat on the ground. It is one of three forms of locomotion adopted by terrestrial mammals. T ...
, ''Borhyaena'' has been suggested to have been
digitigrade
In terrestrial vertebrates, digitigrade () locomotion is walking or running on the toes (from the Latin ''digitus'', 'finger', and ''gradior'', 'walk'). A digitigrade animal is one that stands or walks with its toes (metatarsals) touching the groun ...
but this has been questioned. The one exception was ''
Thylacosmilus
''Thylacosmilus'' is an extinct genus of Saber-toothed predator, saber-toothed metatherian mammals that inhabited South America from the Miocene, Late Miocene to Pliocene Epoch (geology), epochs. Though ''Thylacosmilus'' looks similar to the "Mac ...
'', which has been interpreted as having a digitigrade forefoot and a semiplantigrade hindfoot, this has been supported by fossil tracks.
One unusual aspect of sparassodont paleoecology is that at most fossil localities their remains are nearly ten times rarer than would be expected based on comparisons with carnivorous mammals at fossil sites in other parts of the world. The exact reasons for this are not clear, though this appears to be a broader pattern applicable to other groups of Cenozoic South American terrestrial carnivores (i.e.,
terror birds
Phorusrhacids, colloquially known as terror birds, are an extinct clade of large carnivorous flightless birds that were one of the largest species of apex predators in South America during the Cenozoic era; their conventionally accepted temporal ...
).
Sociality
Little is known of the behavior and biology of sparassodonts outside of general locomotor and dietary habits. Argot (2004) proposed that ''Thylacosmilus atrox'' may have exhibited protracted parental care after weaning of the offspring, given that saber teeth in general have been suggested to require long juvenile periods for the young to gain the skill necessary to use them effectively. However, this has not been tested further. Sparassodonts have relatively large and complex brains for metatherians, comparable to those of some Australian marsupials like Australian possums, though the body masses used to produce these estimates of relative brain size are low compared later studies suggesting these values could be overestimated.
Wounds have been documented on the face of specimens of ''Borhyaena tuberata'' and ''Sipalocyon gracilis'', potentially suggesting aggressive habits similar to the modern
Tasmanian devil
The Tasmanian devil (''Sarcophilus harrisii'') (palawa kani: purinina) is a carnivorous marsupial of the family Dasyuridae. Until recently, it was only found on the island state of Tasmania, but it has been reintroduced to New South Wales in ...
(''Sarcophilus harrisii'').
Senses
Sparassodonts appear to have had very little binocular vision, with
borhyaenids
Borhyaenidae is an extinct metatherian family (biology), family of low-slung, heavily built predatory mammals in the order Sparassodonta. Borhyaenids are not true marsupials, but members of a sister taxon, Sparassodonta. Like most metatherians, ...
having the greatest degree of depth perception (but still lower than modern carnivorans) and the eyes of ''
Thylacosmilus
''Thylacosmilus'' is an extinct genus of Saber-toothed predator, saber-toothed metatherian mammals that inhabited South America from the Miocene, Late Miocene to Pliocene Epoch (geology), epochs. Though ''Thylacosmilus'' looks similar to the "Mac ...
'' facing almost completely to the sides. However, later studies have found that ''Thylacosmilus'' likely held its head in a downward-facing position, which would have allowed for more binocular vision than previously thought.
Pathology
Several specimens of hathliacynids (''Sipalocyon'' and ''Cladosictis'') show a pathological disorder characterized by the presence of growths on the surface of the mandible, which in the most extreme cases can result in the loss of several teeth due to bony pathological growths. The exact cause of this condition (i.e., infection, virus, parasite) and why it seems to only occur in small sparassodonts is unknown, though this condition has also been documented in
microbiotheria
Microbiotheria is an australidelphian marsupial order that encompasses two families, Microbiotheriidae and Woodburnodontidae, and is represented by only one extant species, the monito del monte, and a number of extinct species known from fossi ...
ns.
Extinction
After the middle Miocene, sparassodonts began to slowly decline in diversity. Basal borhyaenoids are last known from the early late Miocene (''Pseudolycopsis cabrerai'' and ''Lycopsis viverensis''), and after this time were at least partially replaced by large-bodied basal sparassodonts such as ''Stylocynus''. It has been suggested that this shift in dominance was due to the more omnivorous habits of basal sparassodonts, which may have been better able to exploit the more seasonal climates of South America during the late
Neogene
The Neogene ( ), informally Upper Tertiary or Late Tertiary, is a geologic period and system that spans 20.45 million years from the end of the Paleogene Period million years ago ( Mya) to the beginning of the present Quaternary Period Mya. ...
. Borhyaenids are last known from the latest
Miocene
The Miocene ( ) is the first 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 means "less recen ...
, though only fragmentary remains of this group are known from this period. Later remains assigned to this group have since been reidentified as thylacosmilids or
procyonids
Procyonidae is a New World family of the order Carnivora. It comprises the raccoons, ringtails, cacomistles, coatis, kinkajous, olingos, and olinguitos. Procyonids inhabit a wide range of environments and are generally omnivorous.
Characte ...
. By the
Pliocene
The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58Hathliacynidae and the
Thylacosmilidae
Thylacosmilidae is an extinct Family (biology), family of metatherian predators, related to the modern marsupials, which lived in South America between the Miocene and Pliocene epochs. Like other South American mammalian predators that lived prio ...
.
Pliocene
The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58didelphids such as ''
Lutreolina
''Lutreolina'' is a genus of opossum found in South America. Both extant species in this genus are known as lutrine opossums. They have an otter-like body plan and occasionally semiaquatic tendencies, hence the genus name ''Lutreolina'', which is ...
'' that appeared around this time. Hathliacynids are last definitively known from the early
Pliocene
The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58 It is still not certain why Sparassodonta declined in diversity and became extinct during the late Cenozoic, but it appears as though competition from
eutherian
Eutheria (; from Greek , 'good, right' and , 'beast'; ) is the clade consisting of all therian mammals that are more closely related to placentals than to marsupials.
Eutherians are distinguished from noneutherians by various phenotypic tra ...
carnivorans
Carnivora is a monophyletic order of placental mammals consisting of the most recent common ancestor of all cat-like and dog-like animals, and all descendants of that ancestor. Members of this group are formally referred to as carnivorans, ...
was not a factor, as the placental analogues of sparassodonts (
dogs
The dog (''Canis familiaris'' or ''Canis lupus familiaris'') is a domesticated descendant of the wolf. Also called the domestic dog, it is derived from the extinct Pleistocene wolf, and the modern wolf is the dog's nearest living relative. Do ...
,
weasels
Weasels are mammals of the genus ''Mustela'' of the family Mustelidae. The genus ''Mustela'' includes the least weasels, polecats, stoats, ferrets and European mink. Members of this genus are small, active predators, with long and slender bodi ...
, and
saber-toothed cats Sabretooth or sabertooth may refer to:
Animals
* Saber-toothed cat, several prehistoric felines
** ''Smilodon'', a prehistoric genus of felidae
* Sabertooth fish, a deep-sea fish found in the tropics
** Sabre-toothed blenny, ''Aspidontus taenia ...
) did not enter South America until the middle Pleistocene, several million years after their sparassodont counterparts became extinct. Sparassodonts did coexist with ''
Cyonasua
''Cyonasua'' (meaning "dog-coati" in Greek) is an extinct genus of procyonid from the Late Miocene to Middle Pleistocene of South America. Fossils of ''Cyonasua'' have been found in Argentina ( Ituzaingó, Epecuén, Huayquerías, Monte Hermoso, ...
''-group procyonids during the late Miocene and Pliocene, but ''
Cyonasua
''Cyonasua'' (meaning "dog-coati" in Greek) is an extinct genus of procyonid from the Late Miocene to Middle Pleistocene of South America. Fossils of ''Cyonasua'' have been found in Argentina ( Ituzaingó, Epecuén, Huayquerías, Monte Hermoso, ...
''-group procyonids appear to have been primarily omnivorous and filled ecological niches that sparassodonts never occupied, which may be one reason that these animals were able to colonize South America despite the diverse predator guild in the late Miocene. The overall decline in sparassodont diversity from the late Miocene to the end of the Pliocene may be linked to the cooling climates following the middle Miocene climatic optimum and the onset of the
Pleistocene
The Pleistocene ( , often referred to as the ''Ice age'') is the geological Epoch (geology), epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fina ...
ice ages
An ice age is a long period of reduction in the temperature of Earth's surface and atmosphere, resulting in the presence or expansion of continental and polar ice sheets and alpine glaciers. Earth's climate alternates between ice ages and gree ...