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Marsupials; Fossil range: Early Cretaceous–Recent PreЄ Є O S D C P T J K Pg N
	Female Eastern Grey Kangaroo with a joey in her pouch
Scientific classification
Kingdom:	Animalia;
Phylum:	Chordata;
Class:	Mammalia;
Subclass:	Theria;
Infraclass:	Marsupialia; Illiger, 1811
Orders
	Didelphimorphia; Paucituberculata; Microbiotheria; Dasyuromorphia; Peramelemorphia; Notoryctemorphia; Diprotodontia; †Sparassodonta; †Yalkaparidontia;

This article is about mammals. For the marsupial frogs, see Gastrotheca.

Marsupials are an infraclass of mammals, characterized by a distinctive pouch (called the marsupium), in which females carry their young through early infancy.

[edit] History

Isolated petrosals of Djarthia murgonensis, Australia's oldest marsupial fossils^[3]

It was once commonly believed that marsupials were a primitive forerunner of modern placental mammals, but fossil evidence, first presented by researcher M.J. Spechtt in 1982, conflicts with this assumption^{[citation needed]}. Instead, both main branches of the mammal tree appear to have evolved concurrently toward the end of the Mesozoic era. In the absence of soft tissues, such as the pouch and reproductive system, fossil marsupials can be distinguished from placentals by the form of their teeth; primitive marsupials possess four pairs of molar teeth in each jaw, whereas placental mammals never have more than three pairs.^[4]

Using this criterion, the earliest known marsupial is Sinodelphys szalayi, which lived in China around 125 million years ago.^[5]^[6]^[7] This makes it almost contemporary to the earliest placental fossils, which have been found in the same area.^[8]

The discovery of Chinese marsupials appears to support the idea that marsupials reached Australia via Southeast Asia.^[9] There are a few species of marsupials still living in Asia, especially in the Sulawesi region of Indonesia. These marsupials coexist with primates, hooved mammals and other placentals.^{[citation needed]} However, due to the fact that Australia and China were separated by the wide Tethys Sea in the early Cretaceous into the Northern continent of Laurasia and Southern continent of Gondwana, marsupials had to take a much longer route around. From their origin in East Laurasia (modern day China), they spread westwards into modern North America (still attached to Eurasia) and skipped across to South America, which was connected to North America up until around 65MYA. Here they radiated into Borhyaenids and Shrew Opossums, creating a unique fauna found in South America and Antarctica (which were connected until 35MYA). Marsupials reached Australia via Antarctica about 50MYA just after Australia had split off, suggesting a single dispersion event of several of just one species, related to South America's Monito del Monte (Microbiothere), rafted across the widening, but still narrow gap between Australia and Antarctica at that time. In Australia, being the only mammals present (except a few Austrosphenids like echidnas and platypuses) they radiated into the wide varieties we see today, even island hopping some way through the Indonesian archipelagos, almost completing a circumnavigation back to their homeland in China. ^[10]

On most continents, placental mammals were much more successful and no marsupials survived, though in South America the opossums retained a strong presence, and the Tertiary saw the genesis of marsupial predators such as the borhyaenids and the saber-toothed Thylacosmilus. In Australia, however, marsupials displaced placental mammals entirely, and have since dominated the Australian ecosystem. Marsupial success over placental mammals in Australia has been attributed to their comparatively low metabolic rate, a trait which would prove helpful in the hot Australian climate.^{[citation needed]} As a result, native Australian placental mammals (such as hopping mice) are more recent immigrants.

[edit] Description

An early birth removes a developing marsupial from its parent's body much sooner than in placental mammals, and thus marsupials have not developed a complex placenta to protect the embryo from its mother's immune system. Though early birth places the tiny newborn marsupial at a greater environmental risk, it significantly reduces the dangers associated with long pregnancies, as there is no need to carry a large fetus to full-term in bad seasons.

Because newborn marsupials must climb up to their mother's nipples, their front limbs are much more developed than the rest of the body at the time of birth. It is possible that this requirement has resulted in the limited range of locomotor adaptations in marsupials compared to placentals. Marsupials must develop a grasping forepaw during their early youth, making the transition from this limb into a hoof, wing, or flipper, as some groups of placental mammals have done, far more difficult.

There are about 334 species of marsupial, and over 200 are native to Australia and neighboring northern islands. There are also 100 extant American species; these are centered mostly in South America, but the Great American Interchange has provided Central America with 13 species, and North America with one (the Virginia Opossum).

Some structural features can be found in marsupials. Ossified patellae are absent. Marsupials (and also monotremes) also lack a gross communication (corpus callosum) between the right and left brain hemisphere.^[11]

[edit] Reproductive system

Marsupials' reproductive systems differ markedly from those of placental mammals (Placentalia). Females have two lateral vaginas, which lead to separate uteruses, but both open externally through the same orifice. A third canal, the median vagina, is used for birth. This canal can be transitory or permanent.^[11] The males generally have a two-pronged penis, which corresponds to the females' two vaginas.^[12] The penis is used only for discharging semen into females, and there is instead a urogenital sac used to store waste before expulsion.

Pregnant females develop a kind of yolk sac in their wombs, which delivers nutrients to the embryo. Marsupials give birth at a very early stage of development (about 4–5 weeks); after birth, newborn marsupials crawl up the bodies of their mothers and attach themselves to a nipple, which is located inside the marsupium. There they remain for a number of weeks, attached to the nipple. The offspring are eventually able to leave the marsupium for short periods, returning to it for warmth, protection and nourishment.

[edit] Taxonomy

Taxonomically, there are two primary divisions of Marsupialia: American marsupials and the Australian marsupials.^[1]^[2] The Order Microbiotheria (which has only one species, the Monito del Monte) is found in South America but is believed to be more closely related to the Australian marsupials. There are many small arboreal species in each group. The term opossums is properly used to refer to the American species (though possum is a common diminutive), while similar Australian species are properly called possums.

The Koala
(Phascolarctos cinereus)

The Common Brushtail Possum (Trichosurus vulpecula)

The Sugar Glider (Petaurus breviceps)

The Squirrel Glider
(Petaurus norfolcensis)

The Virginia Opossum (Didelphis virginiana), the only North American marsupial north of Mexico.

Superorder Ameridelphia
- Order Didelphimorphia (93 species)
  - Family Didelphidae: opossums
- Order Paucituberculata (6 species)
  - Family Caenolestidae: shrew opossums
Superorder Australidelphia
- Order †Yalkaparidontia
- Order Microbiotheria (1 species)
  - Family Microbiotheriidae: Monito del Monte
- Order Dasyuromorphia (71 species)
  - Family †Thylacinidae: Thylacine
  - Family Dasyuridae: antechinuses, quolls, dunnarts, Tasmanian Devil, and relatives
  - Family Myrmecobiidae: Numbat
- Order Peramelemorphia (24 species)
  - Family Thylacomyidae: bilbies
  - Family †Chaeropodidae: Pig-footed Bandicoot
  - Family Peramelidae: bandicoots and allies
- Order Notoryctemorphia (2 species)
  - Family Notoryctidae: marsupial moles
- Order Diprotodontia (137 species)
  - Family Phascolarctidae: Koala
  - Family Vombatidae: wombats
  - Family †Diprotodontidae: diprotodon
  - Family Phalangeridae: brushtail possums and cuscuses
  - Family Burramyidae: pygmy possums
  - Family Tarsipedidae: Honey Possum
  - Family Petauridae: Striped Possum, Leadbeater's Possum, Yellow-bellied Glider, Sugar Glider, Mahogany Glider, Squirrel Glider
  - Family Pseudocheiridae: ringtailed possums and relatives
  - Family Potoridae: potoroos, rat kangaroos, bettongs
  - Family Acrobatidae: Feathertail Glider and Feather-tailed Possum
  - Family Hypsiprymnodontidae: Musky Rat-kangaroo
  - Family Macropodidae: kangaroos, wallabies, and relatives
  - Family †Thylacoleonidae: marsupial lions
- Order †Sparassodonta

† indicates extinction

[edit] See also

Metatheria

[edit] References

^ ^a ^b Gardner, Alfred (2005-11-16). Wilson, D. E., and Reeder, D. M.. ed. Mammal Species of the World (3rd ed.). Johns Hopkins University Press. pp. 3–21. ISBN 0-801-88221-4. http://www.bucknell.edu/msw3.
^ ^a ^b Groves, C. (2005). Wilson, D. E., & Reeder, D. M.. ed. Mammal Species of the World (3rd ed.). Johns Hopkins University Press. pp. 22–70. ISBN 0-801-88221-4. http://www.bucknell.edu/msw3.
^ http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0001858;jsessionid=A57F0FDB595AC49992E2B5A390FA104C
^ Benton, Michael J. (1997). Vertebrate Palaeontology. London: Chapman & Hall. p. 306. ISBN 0-412-73810-4.
^ Rincon, P., Oldest Marsupial Ancestor Found, BBC, Dec 2003
^ Pickrell, J., Oldest Marsupial Fossil Found in China, National Geographic, December 2003
^ Klinger, M.A., Sinodelphys szalayi, Carnegie Mellon Natural History, 2003
^ Ji, Q., et al., The Earliest Known Eutherian Mammal, Nature, 416, Pages 816-822, Apr 2002
^ Harrison, L., The Migration Route of the Australian Marsupial Fauna, Australian Zoologist, Volume 3, Pages 247-263, 1924
^ [2005. T.S. Kemp The origin and evolution of mammals.]
^ ^a ^b Nowak, Ronald M. (1999). Walker's Book Of Mammals, Sixth Edition. Baltimore and London: Johns Hopkins University Press. p. 5. ISBN 0-8018-5789-9.
^ [1] Iowa State University Biology Dept. Discoveries about Marsupial Reproduction Anna King 2001. webpage] (note shows code, html extension omitted)

Tim Flannery (1994),The Future Eaters: An Ecological History of the Australasian Lands and People, pages 67–75. ISBN 0-8021-3943-4 ISBN 0-7301-0422-2
Tim Flannery, Country: a continent, a scientist & a kangaroo, pages 196–200. ISBN 1-920885-76-5
Austin, C.R. ed. Reproduction in Mammals. Melbourne: Cambridge University Press,1982.
Bronson, F. H. Mammalian Reproductive Biology. Chicago: University of Chicago Press, 1989.
Dawson, Terrence J. Kangaroos: Biology of Largest Marsupials. New York: Cornell University Press, 1995.
Frith, H. J. and J. H. Calaby. Kangaroos. New York: Humanities Press, 1969.
Gould, Edwin and George McKay. Encyclopedia of Mammals. San Diego: Academic Press, 1998.
Hunsaker, Don. The Biology of Marsupials. New York: Academic Press, 1977.
Johnson, Martin H. and Barry J. Everitt. Essential Reproduction. Boston: Blackwell Scientific Publications, 1984.
Knobill, Ernst and Jimmy D. Neill ed. Encyclopedia of Reproduction. V. 3 New York: Academic Press, 1998
McCullough, Dale R. and Yvette McCullough. Kangaroos in Outback Australia: Comparative Ecology and Behavior of Three Coexisting Species. New York: Columbia University Press, 2000.
Taylor, Andrea C. and Paul Sunnucks. Sex of Pouch Young Related to Maternal Weight in Macropus eugeni and M. parma. Australian Journal of Zoology 1997 V. 45 pp. 573–578