Diprotodont
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Diprotodontia (, from
Greek Greek may refer to: 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 of all kno ...
"two forward teeth") is the largest extant
order Order, ORDER or Orders may refer to: * A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica * Categorization, the process in which ideas and objects are recognized, differentiated, and understood ...
of
marsupials Marsupials are a diverse group of mammals belonging to the infraclass Marsupialia. They are natively found in Australasia, Wallacea, and the Americas. One of marsupials' unique features is their reproductive strategy: the young are born in a ...
, with about 155 species, including the
kangaroo Kangaroos are marsupials from the family Macropodidae (macropods, meaning "large foot"). In common use, the term is used to describe the largest species from this family, the red kangaroo, as well as the antilopine kangaroo, eastern gre ...
s,
wallabies A wallaby () is a small or middle-sized macropod native to Australia and New Guinea, with introduced populations in New Zealand, Hawaii, the United Kingdom and other countries. They belong to the same taxonomic family as kangaroos and som ...
,
possums Possum may refer to: Animals * Didelphimorphia, or (o)possums, an order of marsupials native to the Americas ** Didelphis, a genus of marsupials within Didelphimorphia *** Common opossum, native to Central and South America *** Virginia opossum, ...
,
koala The koala (''Phascolarctos cinereus''), sometimes inaccurately called the koala bear, is an arboreal herbivorous marsupial native to Australia. It is the only Extant taxon, extant representative of the Family (biology), family ''Phascolar ...
,
wombat Wombats are short-legged, muscular quadrupedal marsupials of the family Vombatidae that are native to Australia. Living species are about in length with small, stubby tails and weigh between . They are adaptable and habitat tolerant, and are ...
s, and many others. Extinct diprotodonts include the
hippopotamus The hippopotamus (''Hippopotamus amphibius;'' ; : hippopotamuses), often shortened to hippo (: hippos), further qualified as the common hippopotamus, Nile hippopotamus and river hippopotamus, is a large semiaquatic mammal native to sub-Sahar ...
-sized ''
Diprotodon ''Diprotodon'' (Ancient Greek: "two protruding front teeth") is an extinct genus of marsupial from the Pleistocene of Australia containing one species, ''D. optatum''. The earliest finds date to 1.77 million to 780,000 years ago but most speci ...
'', and '' Thylacoleo'', the so-called "marsupial lion".


Characteristics

Living diprotodonts are almost all
herbivore A herbivore is an animal anatomically and physiologically evolved to feed on plants, especially upon vascular tissues such as foliage, fruits or seeds, as the main component of its diet. These more broadly also encompass animals that eat ...
s, as were most of those that are now
extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
. A few
insectivorous A robber fly eating a hoverfly An insectivore is a carnivorous animal or plant which eats insects. An alternative term is entomophage, which can also refer to the human practice of eating insects. The first vertebrate insectivores we ...
and
omnivorous An omnivore () is an animal that regularly consumes significant quantities of both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize ...
diprotodonts are known, and the
Potoroidae Potoroidae is a family of marsupials, small Australian animals known as bettongs, potoroos, and rat-kangaroos. All are rabbit-sized, brown, jumping marsupials and resemble a large rodent or a very small wallaby. Taxonomy The potoroids are s ...
are almost unique among vertebrates in being largely
fungivorous Fungivory or mycophagy is the process of organisms consuming fungi. Many different organisms have been recorded to gain their energy from consuming fungi, including birds, mammals, insects, plants, amoebas, gastropods, nematodes, bacteria and othe ...
, but these seem to have arisen as relatively recent adaptations from the mainstream herbivorous lifestyle. The extinct thylacoleonids ("marsupial lions") are the only known group to have exhibited
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 nutrition and energy requirements are met by consumption of animal tissues (mainly mu ...
on a large scale. Diprotodonts are restricted to
Australasia Australasia is a subregion of Oceania, comprising Australia, New Zealand (overlapping with Polynesia), and sometimes including New Guinea and surrounding islands (overlapping with Melanesia). The term is used in a number of different context ...
. The earliest known fossils date to the late
Oligocene The Oligocene ( ) is a geologic epoch (geology), epoch of the Paleogene Geologic time scale, Period that extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that defin ...
, but their genesis certainly lies earlier than this, as large gaps occur in Australia's fossil record, with virtually no fossil record at all in geologically active
New Guinea New Guinea (; Hiri Motu: ''Niu Gini''; , fossilized , also known as Papua or historically ) is the List of islands by area, world's second-largest island, with an area of . Located in Melanesia in the southwestern Pacific Ocean, the island is ...
. The great diversity of known Oligocene diprotodonts suggests the order began to diverge well beforehand. Many of the largest and least athletic diprotodonts (along with a wide range of other
Australian megafauna The term Australian megafauna refers to the megafauna in Australia (continent), Australia during the Pleistocene, Pleistocene Epoch. Most of these species became extinct during the latter half of the Pleistocene, as part of the broader global L ...
) became extinct when humans first arrived in Australia about 50,000 years ago. Their extinction possibly occurred as a direct result of hunting, but was more probably a result of widespread habitat changes brought about by human activities—notably the use of fire. Two key anatomical features, in combination, identify Diprotodontia. Members of the order are, first, "diprotodont" (meaning "two front teeth"): they have a pair of large, procumbent incisors on the lower jaw, a common feature of many early groups of mammals and mammaliforms. The diprotodont jaw is short, usually with three pairs of upper incisors (
wombat Wombats are short-legged, muscular quadrupedal marsupials of the family Vombatidae that are native to Australia. Living species are about in length with small, stubby tails and weigh between . They are adaptable and habitat tolerant, and are ...
s, like
rodent Rodents (from Latin , 'to gnaw') are mammals of the Order (biology), order Rodentia ( ), which are characterized by a single pair of continuously growing incisors in each of the upper and Mandible, lower jaws. About 40% of all mammal specie ...
s have only one pair), and no lower canines. The second trait distinguishing diprotodonts is "
syndactyly Syndactyly is a condition wherein two or more digits are fused together. It occurs normally in some mammals, but is an unusual condition in humans. The term is . Classification Syndactyly can be simple or complex. * In simple syndactyly, adja ...
", a fusing of the second and third digits of the foot up to the base of the claws, which leaves the claws themselves separate. Digit five is usually absent, and digit four is often greatly enlarged. Syndactyly is not particularly common (though the Australian omnivorous marsupials share it) and is generally posited as an adaptation to assist in climbing. Many modern diprotodonts, however, are strictly terrestrial, and have evolved further adaptations to their feet to better suit this lifestyle. This makes the history of the
tree-kangaroo Tree-kangaroos are marsupials of the genus ''Dendrolagus'', adapted for arboreal locomotion. They inhabit the tropical rainforests of New Guinea and far northeastern Queensland, Australia along with some of the islands in the region. All tree- ...
s particularly convoluted: it appears that the animals were arboreal at some time in the far distant past, moving afterward to the ground—gaining long kangaroo-like feet in the process — before returning to the trees, where they further developed a shortening and broadening of the hind feet and a novel climbing method.


Fossil record

The earliest known
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserve ...
of Diprotodontia dates back to the
Late Oligocene The Chattian is, in the geologic timescale The geologic time scale or geological time scale (GTS) is a representation of time based on the rock record of Earth. It is a system of chronological dating that uses chronostratigraphy (the pro ...
(23.03 - 28.4 million years ago), and the earliest identifiable species is '' Hypsiprymnodon bartholomaii'' from the
Early Miocene The Early Miocene (also known as Lower Miocene) is a sub-epoch of the Miocene epoch (geology), Epoch made up of two faunal stage, stages: the Aquitanian age, Aquitanian and Burdigalian stages. The sub-epoch lasted from 23.03 ± 0.05 annum, Ma to ...
.The Paleobiology Database
/ref>


Classification

Until recently, only two
suborder Order () is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and recognized ...
s in Diprotodontia were noted:
Vombatiformes The Vombatiformes are one of the three suborders of the large marsupial order Diprotodontia. Seven of the nine known families within this suborder are extinct; only the families Phascolarctidae, with the koala, and Vombatidae, with three ext ...
which encompassed the
wombat Wombats are short-legged, muscular quadrupedal marsupials of the family Vombatidae that are native to Australia. Living species are about in length with small, stubby tails and weigh between . They are adaptable and habitat tolerant, and are ...
s and
koala The koala (''Phascolarctos cinereus''), sometimes inaccurately called the koala bear, is an arboreal herbivorous marsupial native to Australia. It is the only Extant taxon, extant representative of the Family (biology), family ''Phascolar ...
and Phalangerida which contained all other families. Kirsch et al. (1997) split the families into three
suborder Order () is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and recognized ...
s. In addition, the six
Phalangeriformes Phalangeriformes is a paraphyletic suborder of about 70 species of small to medium-sized arboreal locomotion, arboreal marsupials native to Australia, New Guinea, and Sulawesi. The species are commonly known as possums, opossums, gliders, and ...
families are split into two superfamilies. The Macropodiformes are probably nested within the Phalangeriformes, though whether they are sister to Phalangeroidea or Petauroidea is debated. Order Diprotodontia * Suborder
Macropodiformes The Macropodiformes , also known as macropods, are one of the three suborders of the large marsupial order Diprotodontia. They may in fact be nested within one of the suborders, Phalangeriformes. Kangaroos, wallabies and allies, bettongs, p ...
** Family Hypsiprymnodontidae: ( musky rat-kangaroo) ** Family
Macropodidae Macropodidae is a Family (biology), family of marsupials that includes kangaroos, Wallaby, wallabies, tree-kangaroos, wallaroos, pademelons, quokkas, and several other groups. These genera are allied to the suborder Macropodiformes, containing ...
: (
kangaroos Kangaroos are marsupials from the family Macropodidae (macropods, meaning "large foot"). In common use, the term is used to describe the largest species from this family, the red kangaroo, as well as the antilopine kangaroo, eastern grey ...
,
wallabies A wallaby () is a small or middle-sized macropod native to Australia and New Guinea, with introduced populations in New Zealand, Hawaii, the United Kingdom and other countries. They belong to the same taxonomic family as kangaroos and som ...
and allies) ** Family
Potoroidae Potoroidae is a family of marsupials, small Australian animals known as bettongs, potoroos, and rat-kangaroos. All are rabbit-sized, brown, jumping marsupials and resemble a large rodent or a very small wallaby. Taxonomy The potoroids are s ...
: ( bettongs, potoroos, and rat-kangaroos) ** Family † Balbaridae: (basal quadrupedal kangaroos) * Suborder
Phalangeriformes Phalangeriformes is a paraphyletic suborder of about 70 species of small to medium-sized arboreal locomotion, arboreal marsupials native to Australia, New Guinea, and Sulawesi. The species are commonly known as possums, opossums, gliders, and ...
** Superfamily
Petauroidea Petauroidea is a superfamily of marsupials from Australia and New Guinea. It is part of the suborder Phalangeriformes within the order Diprotodontia, which also includes, among others, wombats, kangaroos, cuscuses. The superfamily Phalangeroid ...
*** Family
Acrobatidae The Acrobatidae are a small family of flying and gliding animals, gliding marsupials containing two genera, each with a single species, the feathertail glider (''Acrobates pygmaeus'') from Australia and feather-tailed possum (''Distoechurus penna ...
: (
feathertail glider The feathertail glider (''Acrobates pygmaeus''), also known as the pygmy gliding possum, pygmy glider, pygmy phalanger, flying phalanger and flying mouse, is a species of marsupial native to eastern Australia. It is the world's smallest gliding ...
and
feather-tailed possum The feather-tailed possum (''Distoechurus pennatus'') is a species of marsupial in the family Acrobatidae. It is found in West Papua and Papua New Guinea. It is not to be confused with the feathertail glider, the only other species in the family ...
) *** Family Petauridae: (
striped possum The striped possum or common striped possum (''Dactylopsila trivirgata'') is a member of the marsupial family Petauridae. It is found mainly in New Guinea. The species is black with three white stripes running head to tail, and its head has whi ...
,
Leadbeater's possum Leadbeater's possum (''Gymnobelideus leadbeateri'') is a critically endangered Phalangeriformes, possum largely restricted to small pockets of Eucalyptus delegatensis, alpine ash, Eucalyptus regnans, mountain ash, and Eucalyptus pauciflora, sno ...
,
yellow-bellied glider The yellow-bellied glider (''Petaurus australis''), also known as the fluffy glider, is an arboreal and nocturnal gliding possum that lives in native eucalypt forests in eastern Australia, from northern Queensland south to Victoria. Habitat Th ...
,
sugar glider The sugar glider (''Petaurus breviceps'') is a small, omnivorous, arboreal, and nocturnal gliding possum. The common name refers to its predilection for sugary foods such as sap and nectar and its ability to glide through the air, much lik ...
,
mahogany glider The mahogany glider (''Petaurus gracilis'') is an endangered gliding possumsquirrel glider The squirrel glider (''Petaurus norfolcensis'') is a nocturnal gliding possum. The squirrel glider is one of the wrist-winged gliders of the genus '' Petaurus''. Habitat This species' home range extends from Bordertown near the South Austral ...
) *** Family
Pseudocheiridae Pseudocheiridae is a family of arboreal marsupials containing 17 extant species of ringtailed possums and close relatives. They are found in forested areas and shrublands throughout Australia and New Guinea. Characteristics Physically, they app ...
: ( ring-tailed possums and allies) *** Family Tarsipedidae: (
honey possum The honey possum or noolbenger (''Tarsipes rostratus''), is a tiny species of marsupial that feeds on the nectar and pollen of a diverse range of flowering plants. Found only in southwest Australia, it is an important pollinator for such plants ...
) ** Superfamily Phalangeroidea *** Family
Burramyidae The pygmy possums are a family of small possums that together form the marsupial family Burramyidae. The five extant species of pygmy possum are grouped into two genera. Four of the species are endemic to Australia, with one species also co-occu ...
: ( pygmy possums) *** Family
Phalangeridae The Phalangeridae are a family of mostly nocturnal marsupials native to Australia, New Guinea, and Eastern Indonesia, including the cuscuses, brushtail possums, and their close relatives. Considered a type of possum, most species are arboreal, a ...
: ( brushtail possums and
cuscus Cuscus ( or ) is the common name generally given to the species within the four genera of Australasian possum of the family Phalangeridae with the most tropical distribution: * '' Ailurops'' * '' Phalanger'' * '' Spilocuscus'' * '' Strigocu ...
es) *** Family † Ektopodontidae: (
sprite possum ''Ektopodon'' is an extinct genus of marsupial, and is the type genus of the family Ektopodontidae which occurred in forested environments in South Australia, Queensland and Victoria. The last species of this group went extinct in the early Ple ...
s) * Suborder
Vombatiformes The Vombatiformes are one of the three suborders of the large marsupial order Diprotodontia. Seven of the nine known families within this suborder are extinct; only the families Phascolarctidae, with the koala, and Vombatidae, with three ext ...
** Family
Phascolarctidae The Phascolarctidae (''φάσκωλος (phaskolos)'' - pouch or bag, ''ἄρκτος (arktos)'' - bear, from the Greek ''phascolos'' + ''arctos'' meaning pouched bear) is a family of marsupials of the order Diprotodontia, consisting of only one ...
:
koala The koala (''Phascolarctos cinereus''), sometimes inaccurately called the koala bear, is an arboreal herbivorous marsupial native to Australia. It is the only Extant taxon, extant representative of the Family (biology), family ''Phascolar ...
(one species) ** Family Vombatidae:
wombat Wombats are short-legged, muscular quadrupedal marsupials of the family Vombatidae that are native to Australia. Living species are about in length with small, stubby tails and weigh between . They are adaptable and habitat tolerant, and are ...
s (three species) ** Family †
Diprotodontidae Diprotodontidae is an extinct family of large herbivorous marsupials, endemic to Australia and New Guinea during the Oligocene through Pleistocene periods from 28.4 million to 40,000 years ago. Description The family primarily consisted of lar ...
: ( giant wombats) ** Family †
Ilariidae Ilariidae is a family of extinct vombatiform marsupials. Most ilariids are found in the middle Tertiary faunal assemblages of South Australia South Australia (commonly abbreviated as SA) is a States and territories of Australia, state in t ...
** Family †
Maradidae ''Marada arcanum'' is a species of Vombatiformes discovered in 2001 at the Oligocene Hiatus Site at Australian Fossil Mammal Sites (Riversleigh), Riversleigh. It is the only member of the genus ''Marada''. It exhibits plesiomorphic and apomorphic ...
** Family † Palorchestidae: ( marsupial tapirs) ** Family † Thylacoleonidae: ( marsupial lions) ** Family †
Wynyardiidae Wynyardiidae is an extinct family of possum-like marsupials from the early Miocene of Wynyard in Tasmania, long been considered to display skeletal features that are intermediate between the primitive polyprotodont and the advanced diprotodont ...
† means extinct family, genus or species


See also

* List of mammal genera *
List of recently extinct mammals Recently extinct mammals are defined by the International Union for Conservation of Nature (IUCN) as any mammals that have become extinct since the year 1500 CE. Since then, roughly 80 mammal species have become extinct. Extinction of taxa ...
*
List of prehistoric mammals This is an incomplete list of prehistoric mammals. It does not include List of mammals, extant mammals or List of extinct mammals, recently extinct mammals. For extinct primate species, see: list of fossil primates.Mikko's Phylogeny Archiv Ma ...


References

{{Authority control Mammal orders Extant Chattian first appearances Taxa named by Richard Owen