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Marsupial
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This article is about the mammals. For frogs, see Marsupial frog.
Marsupials[1][2]
Temporal range: Paleocene – Holocene, 65–0 Ma
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Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Clade: Metatheria
Infraclass: Marsupialia
Illiger, 1811
Orders
Didelphimorphia
Paucituberculata
Australidelphia
Microbiotheria
Dasyuromorphia
Peramelemorphia
Notoryctemorphia
Diprotodontia
†Yalkaparidontia
Present-day distribution of marsupials.
Marsupials are any members of the mammalian infraclass Marsupialia. All extant marsupials are endemic to Australasia and the Americas. A distinctive characteristic common to these species is that most of the young are carried in a pouch. Well-known marsupials include kangaroos, wallabies, koalas, possums, opossums, wombats, and Tasmanian devils. Some lesser known marsupials are the potoroo and the quokka.
Marsupials represent the clade originating from the last common ancestor of extant metatherians. Like other mammals in the Metatheria, they give birth to relatively undeveloped young that often reside with the mother in a pouch, for a certain amount of time. Close to 70% of the 334 extant species occur on the Australian continent (the mainland, Tasmania, New Guinea and nearby islands). The remaining 100 are found in the Americas — primarily in South America, but thirteen in Central America, and one in North America, north of Mexico.
Contents
[hide]
1 Taxonomy
2 Anatomy
2.1 Description
2.1.1 Skull and teeth
2.1.2 Torso
2.1.3 General and convergences
2.2 Reproductive system
2.2.1 Male reproductive system
2.2.2 Female reproductive system
2.2.2.1 Early development
3 Evolution
4 See also
5 Notes
6 References
7 Further reading
8 External links
Taxonomy[edit]
Taxonomically, the two primary divisions of Marsupialia are American marsupials (Ameridelphia) and Australian marsupials (Australidelphia).[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 Australian marsupials. DNA evidence supports a South American origin for marsupials, with Australian marsupials arising from a single Gondwanan migration of marsupials from South America to Australia.[3][4] There are many small arboreal species in each group. The term "opossum" is used to refer to American species (though "possum" is a common diminutive), while similar Australian species are properly called "possums".
†Extinct
Superorder Ameridelphia
Order Didelphimorphia (93 species)
Family Didelphidae: opossums
Order Paucituberculata (seven species)
Family Caenolestidae: shrew opossums
Superorder Australidelphia
Order Microbiotheria (one species)
Family Microbiotheriidae: monito del monte
?Order †Yalkaparidontia
Order Dasyuromorphia (75 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 (two species)
?Family Notoryctidae: marsupial moles
Order Diprotodontia (137 species)
Family Phascolarctidae: koalas
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 Potoroidae: 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
Marsupialia

Didelphimorphia

Paucituberculata

Australidelphia

Microbiotheria

Diprotodontia

Dasyuromorphia

Notoryctemorphia

Peramelemorphia

Cladogram based on Gallus, S. et al. (2015)[5]
Anatomy[edit]
Koala
(Phascolarctos cinereus)
Marsupials have the typical characteristics of mammals—e.g., mammary glands, three middle ear bones, and true hair. There are, however, striking differences as well as a number of anatomical features that separate them from Eutherians.
In addition to the front pouch, which contains multiple nipples for the protection and sustenance of their young, marsupials have other common structural features. Ossified patellae are absent in most modern marsupials (though a small number of exceptions are reported)[6] and epipubic bones are present. Marsupials (and monotremes) also lack a gross communication (corpus callosum) between the right and left brain hemispheres.[7]
Description[edit]
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Skull and teeth[edit]
The skull has peculiarities in comparison to higher mammals. In general, the skull is relatively small and tight. Holes (foramen lacrimale) are located in the front of the orbit. The cheekbone is enlarged and extends further to the rear. The angular extension (processus angularis) of the lower jaw is bent toward the center. Another feature is the hard palate which, in contrast to the higher mammals' foramina, always have more openings. The teeth differ from that of placental mammals, so that all taxa except wombats have a different number of incisors in the upper and lower jaws. The early marsupials had a dental formula from 5 / 4-1 / 1-3 / 3-4 / 4, that is, per pine half; they have five maxilla or four mandibular incisors, one canine, three premolars and four molars, for a total of 50 teeth. Some taxa, such as the opossum, have the original number of teeth. In other groups the number of teeth is reduced. Marsupials in many cases have 40 to 50 teeth, significantly more than placental mammals. The upper jaw has a high number of incisors, up to ten, and they have more molars than premolars. The second set of teeth grows in only at the 3rd premolar: all remaining teeth are already created as permanent teeth.
Torso[edit]
Few general characteristics describe their skeleton. In addition to details in the construction of the ankle, bones (Ossa epubica) are characteristic, two from the pubic bone of the pelvis, which is a forwardly projecting bone. Since these are present in males and pouchless species, it is believed that they originally had nothing to do with reproduction, but served in the muscular approach to the movement of the hind limbs. The egg-laying platypus have marsupial bones. This could be explained by an original feature of mammals. Marsupial reproductive organs differ from the higher mammals. For them, the reproductive tract is doubled. The females have two uteri and two vaginas, and before birth, a birth canal forms between them, the median vagina.[7] The males have a split or double penis lying in front of the scrotum.[8]
A pouch is present in some but not all species. Some marsupials have a permanent bag, whereas in others the pouch develops during gestation, as with the shrew opossum, where the young are hidden only by skin folds or in the fur of the mother. The arrangement of the pouch is variable to allow the offspring to receive maximum protection. Locomotive kangaroos have a pouch opening at the front, while many others that walk or climb on all fours have the opening in the back. Usually, only females have a pouch, but the male water opossum has a pouch that is used to accommodate his genitalia while swimming or running.
General and convergences[edit]
The sugar glider, a marsupial, (left) and flying squirrel, a rodent, (right) are examples of convergent evolution.
Marsupials have adapted to many habitats, reflected in the wide variety in their build. The largest living marsupial, the red kangaroo, grows up to 1.8 metres (5.9 ft) in height and 90 kilograms (200 lb) in weight, but extinct genera, such as Diprotodon, were significantly larger and heavier. The smallest members of this group are the marsupial mice, which often reach only 5 centimetres (2.0 in) in body length.
Some species resemble higher mammals and are examples of convergent evolution. The extinct Thylacine strongly resembled the placental wolf, hence its nickname "Tasmanian wolf". Flying and the associated ability to glide occurred both with marsupials (as with sugar gliders) and some higher mammals (as with flying squirrels), which developed independently. Other groups such as the kangaroo, however, do not have placental counterparts.
Reproductive system[edit]
See also: Kangaroo § Reproduction and life cycle
Female eastern grey kangaroo with a joey in her pouch
Marsupials' reproductive systems differ markedly from those of placental mammals.[9][10] During embryonic development, a choriovitelline placenta forms in all marsupials. In bandicoots, an additional chorioallantoic placenta forms, although it lacks the chorionic villi found in eutherian placentas.
The evolution of reproduction in marsupials, and speculation about the ancestral state of mammalian reproduction, have engaged discussion since the end of the 19th century. Both sexes possess a cloaca,[10] which is connected to a urogenital sac used to store waste before expulsion. The bladder of marsupials functions as a site to concentrate urine and empties into the common urogenital sinus in both females and males.[10]
Male reproductive system[edit]
Most male marsupials, except for macropods[11] and marsupial moles,[12] have a bifurcated penis, separated into two columns, so that the penis has two ends corresponding to the females' two vaginas.[7][10][13][14][8][15][16] The penis is used only during copulation, and is separate from the urinary tract.[8][10] It curves forward when erect,[17] and when not erect, it is retracted into the body in an S-shaped curve.[8] Neither marsupials nor monotremes possess a baculum.[7] The shape of the glans penis varies among marsupial species.[8][18][19][20]
The male thylacine had a pouch that acted as a protective sheath, covering his external reproductive organs while he ran through thick brush.[21]
The shape of the urethral grooves of the males' genitalia is used to distinguish between Monodelphis brevicaudata, Monodelphis domestica, and Monodelphis americana. The grooves form 2 separate channels that form the ventral and dorsal folds of the erectile tissue.[22] Several species of dasyurid marsupials can also be distinguished by their penis morphology.[23]
The only accessory sex glands marsupials possess are the prostate and bulbourethral glands.[24] There are no ampullae, seminal vesicles or coagulating glands.[25][13] The prostate is proportionally larger in marsupials than in placental mammals.[8] During the breeding season, the male tammar wallaby's prostate and bulbourethral gland enlarge. However, there does not appear to be any seasonal difference in the weight of the testes.[26]
Female reproductive system[edit]
See also: Birth § Marsupials
Female reproductive anatomy of several marsupial species
Female marsupials have two lateral vaginas, which lead to separate uteri, 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.[7]
Marsupials give birth at a very early stage of development (about four to five weeks); after birth, newborn marsupials crawl up the bodies of their mothers and attach themselves to a nipple, which is located on the underside of the mother either inside a pouch called the marsupium or open to the environment. To crawl to the nipple and attach to it, the marsupial must have well-developed forelimbs and facial structures.[27][28] This is accomplished by accelerating forelimb and facial development in marsupials compared to placental mammals, which results in decelerated development of such structures as the hindlimb and brain. 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.
Early development[edit]
An early birth removes a developing marsupial from its mother's body much sooner than in placental mammals, thus marsupials have not developed a complex placenta to protect the embryo from its mother's immune system. Though early birth puts 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. Marsupials are extremely altricial animals, needing to be intensely cared for immediately following birth (cf. precocial).
Because newborn marsupials must climb up to their mother's nipples, their front limbs are much more developed than the rest of their bodies at the time of birth. This requirement has been argued to have resulted in the limited range of locomotor adaptations in marsupials compared to placentals. Marsupials must develop grasping forepaws during their early youth, making the transition from these limbs into hooves, wings, or flippers, as some groups of placental mammals have done, far more difficult. However, several marsupials do possess atypical forelimb morphologies, such as the hooved forelimbs of the pig-footed bandicoot, suggesting that the range of forelimb speciation is a lot less limited than usually assumed.[29]
An infant marsupial is known as a joey. Marsupials have a very short gestation period (about four to five weeks), and the joey is born in an essentially fetal state. The blind, furless, miniature newborn, the size of a jelly bean,[30] crawls across its mother's fur to make its way into the pouch, where it latches onto a teat for food. It will not re-emerge for several months, during which time it develops fully. After this period, the joey begins to spend increasing lengths of time out of the pouch, feeding and learning survival skills. However, it returns to the pouch to sleep, and if danger threatens, it will seek refuge in its mother's pouch for safety.
Joeys stay in the pouch for up to a year in some species, or until the next joey is born. A marsupial joey is unable to regulate its own body temperature and relies upon an external heat source. Until the joey is well-furred and old enough to leave the pouch, a pouch temperature of 30–32 °C (86–90 °F) must be constantly maintained.
Joeys are born with "oral shields". In species without pouches or with rudimentary pouches these are more developed than in forms with well-developed pouches, implying a role in maintaining the young attached to the mother's nipple.[31]
Evolution[edit]
See also: Evolution of mammals
Isolated petrosals of Djarthia murgonensis, Australia's oldest marsupial fossils[32]
Dentition of the herbivorous eastern grey kangaroo, as illustrated in Knight's Sketches in Natural History
The relationships among the three extant divisions of mammals (monotremes, marsupials, and placentals) were long a matter of debate among taxonomists.[33] Most morphological evidence comparing traits such as number and arrangement of teeth and structure of the reproductive and waste elimination systems as well as most genetic and molecular evidence favors a closer evolutionary relationship between the marsupials and placental mammals than either has with the monotremes.[34]
Phylogenetic tree of marsupials derived from retroposon data[4]
The ancestors of marsupials, part of a larger group called metatherians, probably split from those of placental mammals (eutherians) during the mid-Jurassic period, though no fossil evidence of metatherians themselves are known from this time.[35] Fossil metatherians are distinguished from eutherians by the form of their teeth; metatherians possess four pairs of molar teeth in each jaw, whereas eutherian mammals (including true placentals) never have more than three pairs.[36] Using this criterion, the earliest known metatherian is Sinodelphys szalayi, which lived in China around 125 million years ago (mya).[37] This makes it a contemporary to some early eutherian species which have been found in the same area.[38]
The oldest metatherian fossils are found in present-day China.[39] About 100 mya, the supercontinent Pangaea was in the process of splitting into the northern continent Laurasia and the southern continent Gondwana, with what would become China and Australia already separated by the Tethys Ocean. From there, metatherians spread westward into modern North America (still attached to Eurasia), where the earliest true marsupials are found. Marsupials are difficult to distinguish from other fossils, as they are characterized by aspects of the reproductive system which do not normally fossilize (including pouches) and by subtle changes in the bone and tooth structure that show a metatherian is part of the marsupial crown group (the most exclusive group that contains all living marsupials). The earliest definite marsupial fossil belongs to the species Peradectes minor, from the Paleocene of Montana, dated to about 65 million years ago.[40] From their point of origin in Laurasia, marsupials spread to South America, which was possibly connected to North America at around 65 mya through a ridge that has since moved on to become the Caribbean Archipelago.[41][42] Laurasian marsupials eventually died off, for not entirely clear reasons; convention has it that they disappeared due to competition with placentals, but this is no longer accepted to be the primary reason.[43]
Marsurpials, Peradectes and the related Herpetotheriidae are nested within a clade of metatherians that also included a variety of Cretaceous North American taxa.[44]
Metatheria

Holoclemensia

Pappotherium

Sulestes

Oklatheridium

Tsagandelta

Lotheridium

Deltatheroides

Deltatheridium

Nanocuris

Atokatheridium

Marsupialiformes

Gurlin Tsav skull

Borhyaenidae

Mayulestes

Jaskhadelphys

Andinodelphys

Pucadelphys

Asiatherium

Iugomortiferum

Kokopellia

Aenigmadelphys

Anchistodelphys

Glasbiidae
Glasbius

Pediomyidae
Pediomys

Stagodontidae

Pariadens

Eodelphis

Didelphodon

Alphadontidae

Turgidodon

Alphadon

Albertatherium

Marsupialia

In South America, the opossums evolved and developed a strong presence, and the Paleogene also saw the evolution of shrew opossums (Paucituberculata) alongside non-marsupial metatherian predators such as the borhyaenids and the saber-toothed Thylacosmilus. South American niches for mammalian carnivores were dominated by these marsupial and sparassodont metatherians. While placental predators were absent, the metatherians did have to contend with avian (terror bird) and terrestrial crocodylomorph competition. South America and Antarctica remained connected until 35 mya, as shown by the unique fossils found there. North and South America were disconnected until about three million years ago, when the Isthmus of Panama formed. This led to the Great American Interchange. Sparassodonts disappeared for unclear reasons – again, this has classically assumed as competition from carnivoran placentals, but the last sparassodonts co-existed with a few small carnivorans like procyonids and canines, and disappeared long before the arrival of macropredatory forms like felines,[45] while didelphimorphs (opossums) invaded Central America, with the Virginia opossum reaching as far north as Canada.
Marsupials reached Australia via Antarctica about 50 mya, shortly after Australia had split off. This suggests a single dispersion event of just one species, most likely a relative to South America's monito del monte (a microbiothere, the only New World australidelphian). This progenitor may have rafted across the widening, but still narrow, gap between Australia and Antarctica. In Australia, they radiated into the wide variety seen today. Modern marsupials appear to have reached the islands of New Guinea and Sulawesi relatively recently via Australia.[46][47][48] A 2010 analysis of retroposon insertion sites in the nuclear DNA of a variety of marsupials has confirmed all living marsupials have South American ancestors. The branching sequence of marsupial orders indicated by the study puts Didelphimorphia in the most basal position, followed by Paucituberculata, then Microbiotheria, and ending with the radiation of Australian marsupials. This indicates that Australidelphia arose in South America, and reached Australia after Microbiotheria split off.[3][4]
In Australia, terrestrial placental mammals disappeared early in the Cenozoic (their most recent known fossils being 55 million-year-old teeth resembling those of condylarths) for reasons that are not clear, allowing marsupials to dominate the Australian ecosystem.[46] Extant native Australian terrestrial placental mammals (such as hopping mice) are relatively recent immigrants, arriving via island hopping from Southeast Asia.[47]
Genetic analysis suggests a divergence date between the marsupials and the placentals at 160 million years ago.[49] The ancestral number of chromosomes has been estimated to be 2n = 14.
A new hypothesis suggests that South American microbiotheres resulted from a back-dispersal from eastern Gondwana due to new cranial and post-cranial marsupial fossils from the Djarthia murgonensis from the early Eocene Tingamarra Local Fauna in Australia that indicate the Djarthia murgonensis is the most plesiomorphic, the oldest unequivocal australidelphian, and may be the ancestral morphotype of the Australian marsupial radiation.[50]
[show]
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t
e
Geological time scale of marsupial evolution
Phanerozoic
Cenozoic
Quaternary
Holocene
10,000 ya – present
First Europeans visit Australia in 1606, settlements begin in 1788.
Dingo introduced 3500-4000 ya. Thylacine and Tasmanian devil subsequently disappear from Australian mainland.
Pleistocene
1.75 Mya – 10,000
Last glacial maximum 18,000–20,000 ya
Extinction of megafauna 45,000–55,000 ya.
First humans arrive in Australia at least 70,000~65,000 ya.
Neogene
Pliocene
5.3–1.7 Mya
Growing diversity in grazing marsupials as a result of grasslands and arid habitats development.
First appearance of large marsupials.
Miocene
Late Miocene 11–5.3 Mya
'Dim age' of marsupial fossils in Australia. Forest-dwellers diminish.
Middle Miocene 16.4–11 Mya
Icehouse conditions result in the number of forest and forest-dwelling marsupials to decrease.
Early Miocene 23.5–16.4 Mya
Greenhouse conditions in Australia result in great diversity of Australian marsupials.
Paleogene
Oligocene 33–23 Mya
Appearance of marsupials in Australian fossil record
Eocene 53–33.7 Mya
Australia separates from Antarctica.
Paleocene
65–53 Mya
High marsupial diversity in South America. Appearance of the oldest Australian marsupial in late Paleocene.
Dinosaurs are wiped off the Earth after an asteroid collision.
Mesozoic
Cretaceous
Late Cretaceous 97–65 Mya
The northern landmass, Laurasia, is inhabited by marsupials. Some of them start dispersing to South America.
Early Cretaceous 135–97 Mya
First appearance of marsupial and placental fossils.
Jurassic 203–135 Mya
Break apart of the great southern landmass, Gondwana. Marsupials and placentals diverge.
Triassic 250–203 Mya
First mammals appear in late Triassic in the supercontinent, Pangaea.
See also[edit]
Wikimedia Commons has media related to Marsupial reproductive system.
Marsupial lawn
Metatheria
Notes[edit]
References[edit]
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Jump up ^ Schneider, Nanette Yvette (August 2011). "The development of the olfactory organs in newly hatched monotremes and neonate marsupials". J Anat. 219 (2): 229–242. doi:10.1111/j.1469-7580.2011.01393.x.
Jump up ^ Beck, Robin M. D.; Godthelp, Henk; Weisbecker, Vera; Archer, Michael; Hand, Suzanne J. (2008). Hawks, John, ed. "Australia's oldest marsupial fossils and their biogeographical implications". PLoS ONE. 3 (3): e1858. PMC 2267999  . PMID 18365013. doi:10.1371/journal.pone.0001858.
Jump up ^ Moyal, Ann Mozley (2004). Platypus: The Extraordinary Story of How a Curious Creature Baffled the World. Baltimore: The Johns Hopkins University Press. ISBN 0-8018-8052-1.
Jump up ^ van Rheede, T.; Bastiaans, T.; Boone, D.; Hedges, S.; De Jong, W.; Madsen, O. (2006). "The platypus is in its place: nuclear genes and indels confirm the sister group relation of monotremes and therians". Molecular Biology and Evolution. 23 (3): 587–597. PMID 16291999. doi:10.1093/molbev/msj064.
Jump up ^ Zhe-Xi Luo; Chong-Xi Yuan; Qing-Jin Meng; Qiang Ji (2011). "A Jurassic eutherian mammal and divergence of marsupials and placentals". Nature. 476 (7361): 442–445. PMID 21866158. doi:10.1038/nature10291.
Jump up ^ Benton, Michael J. (1997). Vertebrate Palaeontology. London: Chapman & Hall. p. 306. ISBN 0-412-73810-4.
Jump up ^ Rincon, Paul (2003-12-12). "Oldest Marsupial Ancestor Found, BBC, Dec 2003". BBC News. Retrieved 2010-03-16.
Jump up ^ Hu, Y; Meng, J; Li, C; Wang, Y (2010). "New basal eutherian mammal from the Early Cretaceous Jehol biota, Liaoning, China". Proceedings of the Royal Society B. 277 (1679): 229–236. PMC 2842663  . PMID 19419990. doi:10.1098/rspb.2009.0203.
Jump up ^ Luo, Zhe-Xi; Ji, Qiang; Wible, John R.; Yuan, Chong-Xi (2003-12-12). "An early Cretaceous tribosphenic mammal and metatherian evolution". Science. 302 (5652): 1934–1940. PMID 14671295. doi:10.1126/science.1090718.
Jump up ^ O'Leary, Maureen A.; Bloch, Jonathan I.; Flynn, John J.; Gaudin, Timothy J.; Giallombardo, Andres; Giannini, Norberto P.; Goldberg, Suzann L.; Kraatz, Brian P.; Luo, Zhe-Xi; Meng, Jin; Ni, Michael J.; Novacek, Fernando A.; Perini, Zachary S.; Randall, Guillermo; Rougier, Eric J.; Sargis, Mary T.; Silcox, Nancy b.; Simmons, Micelle; Spaulding, Paul M.; Velazco, Marcelo; Weksler, John r.; Wible, Andrea L.; Cirranello, A. L. (8 February 2013). "The Placental Mammal Ancestor and the Post–K-Pg Radiation of Placentals". Science. 339 (6120): 662–667. PMID 23393258. doi:10.1126/science.1229237.
Jump up ^ Kemp, Thomas Stainforth (2005). The origin and evolution of mammals (PDF). Oxford: Oxford University Press. p. 217. ISBN 0 19 850760 7.
Jump up ^ Boschman, Lydian M.; van Hinsbergen, Douwe J.J.; Torsvik, Trond H.; Spakman, Wim; Pindell, James L. (23 August 2014). "Kinematic reconstruction of the Caribbean region since the Early Jurassic". Earth-Science Reviews. 138: 102–136. doi:10.1016/j.earscirev.2014.08.007. Retrieved 11 October 2017.
Jump up ^ Sánchez-Villagra, Marcelo (2012). "Why are There Fewer Marsupials than Placentals? On the Relevance of Geography and Physiology to Evolutionary Patterns of Mammalian Diversity and Disparity". Journal of Mammalian Evolution. 20 (4): 279–290. doi:10.1007/s10914-012-9220-3.
Jump up ^ Wilson, G.P.; Ekdale, E.G.; Hoganson, J.W.; Calede, J.J.; Linden, A.V. (2016). "A large carnivorous mammal from the Late Cretaceous and the North American origin of marsupials". Nature Communications. 7. doi:10.1038/ncomms13734.
Jump up ^ Prevosti, Francisco J.; Forasiepi, Analía; Zimicz, Natalia (2011). "The Evolution of the Cenozoic Terrestrial Mammalian Predator Guild in South America: Competition or Replacement?". Journal of Mammalian Evolution. 20: 3–21. doi:10.1007/s10914-011-9175-9.
^ Jump up to: a b Dawkins, Richard (2005). The Ancestor's Tale : A Pilgrimage to the Dawn of Evolution. Boston: Mariner Books. p. 223. ISBN 0-618-61916-X.
^ Jump up to: a b Hand, Suzanne J.; Long, John; Archer, Michael; Flannery, Timothy Fridtjof (2002). Prehistoric mammals of Australia and New Guinea: one hundred million years of evolution. Baltimore: Johns Hopkins University Press. ISBN 0-8018-7223-5.
Jump up ^ Kemp, T.S. (2005). The origin and evolution of mammals. Oxford [Oxfordshire]: Oxford University Press. ISBN 0-19-850761-5.
Jump up ^ Graves JA, Renfree MB (2013) Marsupials in the age of genomics. Annu Rev Genomics Hum Genet
Jump up ^ Beck, Robin M. D.; Godthelp, Henk; Weisbecker, Vera; Archer, Michael; Hand, Suzanne J. (2008). "Australia's Oldest Marsupial Fossils and their Biogeographical Implications". PLoS ONE. 3 (3): e1858. PMC 2267999  . PMID 18365013. doi:10.1371/journal.pone.0001858.
Further reading[edit]
Austin, C. R.; Russell Austin, Colin; Valentine Short, Roger, eds. (21 March 1985). Reproduction in Mammals: Volume 4, Reproductive Fitness. Cambridge University Press. pp. 4–. ISBN 978-0-521-31984-3.
Bronson, F. H. (1989). Mammalian Reproductive Biology. University of Chicago Press. ISBN 978-0-226-07559-4.
Dawson, Terence J. (1995). Kangaroos: Biology of the Largest Marsupials. Cornell University Press. ISBN 0-8014-8262-3.
Flannery, Tim (2002). The Future Eaters: An Ecological History of the Australasian Lands and People. Grove Press. pp. 67–75. ISBN 978-0-8021-3943-6.
Flannery, Tim (2008). Chasing kangaroos : a continent, a scientist, and a search for the world's most extraordinary creature. (1st American ed.). New York: Grove. ISBN 9780802143716.
Flannery, Tim (2005). Country : a continent, a scientist & a kangaroo (2nd ed.). Melbourne: Text Pub. ISBN 1-920885-76-5.
Frith, H. J. and J. H. Calaby. Kangaroos. New York: Humanities Press, 1969.
McKay, George (2006). The Encyclopedia of MAMMALS. Weldon Owen. ISBN 978-1-74089-352-7.
Hunsaker, Don. The Biology of Marsupials. New York: Academic Press, 1977.
Johnson, M. H.; Everitt, Barry J. (1988). Essential Reproduction. Blackwell Scientific. ISBN 978-0-632-02183-3.
Jones, Menna; Dickman, Chris; Archer, Mike (2003). Predators with pouches : the biology of carnivorous marsupials. Collingwood, Victoria: Australia). ISBN 9780643066342.
Knobill, Ernst; Neill, Jimmy D., eds. (1998). Encyclopedia of Reproduction. 3. New York: Academic Press.
McCullough, Dale R. (2000). Kangaroos in Outback Australia: Comparative Ecology and Behavior of Three Coexisting Species. Columbia University Press. ISBN 978-0-231-11916-0. |first2= missing |last2= in Authors list (help)
Nowak, Ronald M. (7 April 1999). Walker's Mammals of the World. JHU Press. ISBN 978-0-8018-5789-8.
Taylor, Andrea C.; Taylor, Paul (1997). "Sex of Pouch Young Related to Maternal Weight in Macropus eugeni and M. parma". Australian Journal of Zoology. 45 (6): 573–578. doi:10.1071/ZO97038.
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Researchers Publish First Marsupial Genome Sequence The National Institutes of Health May 2007
First marsupial genome released. Most differences between the opossom and placental mammals stem from non-coding DNA
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One Tree Hill (season 3)
From Wikipedia, the free encyclopedia
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One Tree Hill (season 3)
One Tree Hill Season 3 DVD cover
Country of origin
United States
No. of episodes
22
Release
Original network
The WB
Original release
October 5, 2005 (2005-10-05) – May 3, 2006 (2006-05-03)
Season chronology
← Previous
Season 2
Next →
Season 4
List of One Tree Hill episodes
The third season of One Tree Hill, an American teen drama television series, began airing on October 5, 2005. The season concluded on May 3, 2006, after 22 episodes. It is the final season that aired on The WB television network. Season three dipped in ratings, averaging 2.8 million viewers weekly.[1]
Contents
[hide]
1 Overview
2 Cast and characters
2.1 Regular
2.2 Recurring
2.3 Guest Cast
3 Episodes
4 Ratings
5 DVD release
6 References
Overview[edit]
This season focuses on the first half of senior year at Tree Hill High School. Starting three months after the second season finale, Haley returns to Tree Hill and tries to save her marriage with Nathan. Peyton gets to know her birth mother. Lucas and Brooke begin a relationship. Keith and Karen get engaged. Nathan patches things up with Lucas, while Mouth begins a complicated relationship with newcomer Rachel, who also spreads a lot of drama through the core characters. After a shooting at the school things between Lucas and Peyton heat up despite Peyton's proposal to Jake.
Cast and characters[edit]
See also: List of One Tree Hill characters
Regular[edit]
Chad Michael Murray as Lucas Scott (22 episodes)
James Lafferty as Nathan Scott (22 episodes)
Hilarie Burton as Peyton Sawyer (22 episodes)
Bethany Joy Galeotti as Haley James Scott (22 episodes)
Paul Johansson as Dan Scott (20 episodes)
Sophia Bush as Brooke Davis (22 episodes)
Barbara Alyn Woods as Deb Scott (13 episodes)
Lee Norris as Mouth McFadden (18 episodes)
Barry Corbin as Whitey Durham (12 episodes)
Craig Sheffer as Keith Scott (13 episodes)
Moira Kelly as Karen Roe (20 episodes)
Recurring[edit]
Danneel Harris as Rachel Gatina (19 episodes)
Bevin Prince as Bevin Mirskey (16 episodes)
Tyler Hilton as Chris Keller (12 episodes)
Sheryl Lee as Ellie Harp (8 episodes)
Vaughn Wilson as Fergie Thompson (7 episodes)
Antwon Tanner as Antwon "Skills" Taylor (7 episodes)
Cullen Moss as Junk Moretti (6 episodes)
Kevin Kilner as Larry Sawyer (6 episodes)
Brett Claywell as Tim Smith (5 episodes)
Kelsey Chow as Gigi Silveri (5 episodes)
Shawn Shepard as Principal Turner (4 episodes)
Guest Cast[edit]
Michael Trucco as Cooper Lee (3 episodes)
Bryan Greenberg as Jake Jagielski (3 episodes
Colin Fickes as Jimmy Edwards (2 episodes)
Amber Wallace as Glenda Farrell (1 episode)
Allison Scagliotti as Abigail "Abby" Brown (1 episode)
Episodes[edit]
See also: List of One Tree Hill episodes
No.
overall
No. in
season
Title
Directed by
Written by
Original air date
Prod.
code
U.S. viewers
(millions)
46
1 "Like You Like an Arsonist" Greg Prange Mark Schwahn October 5, 2005 (2005-10-05) 2T6151 3.46[3]
In the third season premiere, we finally discover Dan's fate after the fire. In addition to the situation with his father, Nathan also has to deal with Haley's reappearance in Tree Hill and the future of their relationship. Peyton tries to find the truth about the mysterious woman claiming to be her mother. Lucas looks forward to Brooke's return after a summer apart.[2]
This episode is named after a song by Paris, Texas.
47
2 "From the Edge of the Deep Green Sea" Kevin Dowling Mark Schwahn October 12, 2005 (2005-10-12) 2T6152 3.12[4]
Brooke has an end of the summer party on the beach. Nathan is home from basketball camp. Lucas deals with Brooke's rules for non-exclusive dating. Brooke ends up having a ball with many guys whereas Lucas spends the night alone and then tells Brooke that he is the one for her, which marks the beginning of his mission to become exclusive. Peyton deals with family issues, by confronting her mom about her drug problem, after Lucas witnesses her buying. Haley tries to tell Nathan how she feels, however he is not ready for a relationship yet and is still angry with her.
This episode is named after a song by The Cure.
48
3 "First Day on a Brand New Planet" Billy Dickson Terrence Coli October 19, 2005 (2005-10-19) 2T6153 3.12[5]
It's the first day of senior year for the Tree Hill gang. Lucas tells Dan he saved his life. Peyton discovers Ellie's secret. Nathan tells Haley he's considering a divorce. Dan accuses several people of starting the dealership fire and tries to get Whitey replaced as Ravens coach.
This episode is named after a song by the band Jesse James.
49
4 "An Attempt to Tip the Scales" Janice Cooke Stacy Rukeyser October 26, 2005 (2005-10-26) 2T6154 3.36[6]
It's Halloween in Tree Hill and everybody's celebrating at TRIC, with special guest band Fall Out Boy. The masquerade ball gives Haley the chance to win Nathan back, while Lucas keeps trying to convince Brooke to be exclusive. Peyton decides to face her new mother and get to know her. Dan tries to use blackmail to become mayor but things don't turn out the way he thought they would when Deb gets in his way.
This episode is named after a song by Bright Eyes.
50
5 "A Multitude of Casualties" Thomas J. Wright R. Lee Fleming, Jr November 2, 2005 (2005-11-02) 2T6155 3.03[7]
The Ravens begin basketball practice. New team captain Nathan clashes with Lucas over Haley, and a new cheerleader named Rachel Gatina sets her sights on Lucas. Dan's mayoral bid takes a surprising turn when an unlikely opponent emerges in Karen.
This episode is named after a song by The Hold Steady.
51
6 "Locked Hearts & Hand Grenades" Marita Grabiak James Stoteraux and Chad Fiveash November 9, 2005 (2005-11-09) 2T6156 3.42[8]
Brooke starts a "Fantasy Boy Draft", after the cheerleaders start fighting over the same guys. Haley and Peyton continue to argue about Nathan. Dan butts heads with Karen as the election draws nearer. Lucas tries to keep up with the fast pace of basketball.
This episode is named after a song by Plan A Project.
52
7 "Champagne for My Real Friends, Real Pain for My Sham Friends" Paul Johansson Mark Schwahn November 16, 2005 (2005-11-16) 2T6157 3.52[9]
The fantasy draft dates begin, Lucas finds himself with Rachel and a very familiar setting. Brooke is on a double date with a surprising person and a surprising couple. Skills goes to Bevin's house for the night, but isn't quite interested in what she has in mind. Mouth and Peyton hang out. Karen and Deb take the campaigning for mayor to a new level, to bring down Dan. Nathan gets mad when Chris gets close to Haley.
This episode is named after a song by Fall Out Boy.
53
8 "The Worst Day Since Yesterday" John Asher Mike Herro & David Strauss November 30, 2005 (2005-11-30) 2T6158 3.41[8]
As the town readies for the Ravens’ first basketball game, Lucas worries that his head and heart aren't in it. Nathan accuses Lucas of being the Ravens' weak link. Peyton and Haley mend fences to support Brooke in her time of need. When the mayoral race tightens, Dan resorts to a smear campaign.
This episode is named after a song by Flogging Molly.
54
9 "How a Resurrection Really Feels" Greg Prange Mark Schwahn December 7, 2005 (2005-12-07) 2T6159 3.33[10]
It is revealed that Lucas knows who tried to kill Dan and he confronts the guilty party. Brooke, Peyton and Haley find themselves behind bars. Peyton is annoyed by the presence of her guilty conscience. Chris Keller convinces Nathan to be his wingman at a cash poker game. The residents of Tree Hill finally choose a mayor.
This episode is named after a song by The Hold Steady.
55
10 "Brave New World" John Asher John A. Norris January 11, 2006 (2006-01-11) 2T6160 3.23[11]
Lucas and Peyton take a road trip together to find Ellie. While out of town, Lucas reconnects with a friend from the past. Brooke, Haley and Mouth successfully launch the Clothes Over Bro's website, only to find that demand far exceeds supply. Deb reveals to Nathan that she tried to kill Dan. Nathan and Haley finally get back together.
This episode is named after a song by Iron Maiden from the album of the same name.
56
11 "Return of the Future" Bethany Rooney Terrence Coli January 18, 2006 (2006-01-18) 2T6161 2.67[12]
The Ravens lose their second game in a row, Whitey decides to teach them a tough lesson. Peyton and Ellie look for support for their album and hope to attract Nada Surf to the project. Both Dan and Haley upset Nathan, while Brooke plans a romantic thank-you for Lucas.
This episode is named after a song by Scooter.
57
12 "I've Got Dreams to Remember" Stuart Gillard Mike Herro & David Strauss January 25, 2006 (2006-01-25) 2T6162 2.70[13]
As the students meet with the school counselor to plan for the future, Nathan and Haley realize they want to attend colleges across the country from one another. Brooke discovers Rachel's reasons for submitting her designs to Rogue Vogue. Meanwhile, Ellie and Peyton bond as they finish the benefit album.
This episode is named after a song by Otis Redding.
58
13 "The Wind That Blew My Heart Away" David Jackson Stacy Rukeyser February 1, 2006 (2006-02-01) 2T6163 3.01[14]
A violent rainstorm knocks out the power in Tree Hill, leading Haley to Nathan's doorstep and Brooke out into the downpour for a dramatic confrontation with Lucas. Peyton finally confronts the truth about Ellie's illness. Mouth is tempted by his new friend, Rachel. Before morning, secrets are revealed, relationships are tested and romance heats up.
This episode is named after a song by Fruit Bats.
59
14 "All Tomorrow's Parties" David Paymer Anna Lotto February 8, 2006 (2006-02-08) 2T6164 2.89[15]
Brooke is forced to grow up in a hurry when her trip to a New York City fashion show prevents her from attending an annual cheerleading tournament. Nathan and Lucas are influenced by the father of a friend. Skills tells Bevin he can't swim to trick her into giving him a swimming lesson. Struggling with her birth mother's cancer, Peyton joins Rachel for a night of blowing off steam. Haley and Nathan use their hotel room as the honeymoon suite they never had.
This episode is named after a song by The Velvet Underground and Nico.
60
15 "Just Watch the Fireworks" Billy Dickson James Stoteraux & Chad Fiveash February 15, 2006 (2006-02-15) 2T6165 2.85[16]
When someone unleashes the students' time capsule on Tree Hill High, Lucas and Mouth must confront a person from their past. Brooke deals with the revelation that secrets aren't the only things she exposed in the time capsule. Dan tries to forgive Keith. Haley is forced to reconcile the man Nathan was before with the man he has become. Peyton's Breast Cancer Benefit Concert at TRIC features performances by Haley and Jack's Mannequin and the return of Fall Out Boy.
This episode is named after a song by Jimmy Eat World.
61
16 "With Tired Eyes, Tired Minds, Tired Souls, We Slept" Greg Prange Mark Schwahn March 1, 2006 (2006-03-01) 2T6166 3.36[17]
In the aftermath of the release of the time capsule, chaos breaks out, Jimmy Edwards holds students of Tree Hill High School hostage with a gun. While lives hang in the balance, Nathan and Lucas put themselves at risk to protect their friends and loved ones. Young lives are shattered and two will end. Dan takes revenge on a close relative.
This episode is named after a song by Explosions in the Sky.
62
17 "Who Will Survive, and What Will Be Left of Them" John Asher Mark Schwahn March 29, 2006 (2006-03-29) 2T6167 2.82[18]
The aftermath of the school shooting and the death of Jimmy Edwards force everyone in Tree Hill to examine their lives. Nathan reassesses his relationship with Haley, while Brooke and Rachel join forces to heal the student body. Dan must deal with the ramifications of killing Keith. Lucas and Peyton consider the implications of their kiss in the library.
This episode is named after an album by Murder by Death. The title is also the tagline of the 1974 horror movie The Texas Chain Saw Massacre.
63
18 "When It Isn't Like It Should Be" Paul Johansson R. Lee Fleming, Jr. April 5, 2006 (2006-04-05) 2T6168 2.93[19]
Rachel invites the gang for a weekend getaway at her family's cabin. Craziness will ensue, including a game of "I Never" which will cause people to do things they normally wouldn't. A nasty little issue Mouth has been harboring toward Rachel will surface. Meanwhile, Karen confronts Dan about Keith's death.
This episode is named after a song by Saves the Day.
64
19 "I Slept with Someone in Fall Out Boy and All I Got Was This Stupid Song Written About Me" Moira Kelly William H. Brown April 12, 2006 (2006-04-12) 2T6169 2.76[20]
Lucas attempts to play in the first game after Keith's death, but he cannot bring himself to do it. After seeing Lucas's pain, Nathan asks Whitey to forfeit the game. Lucas finally tells Whitey, Nathan, and Karen about his heart condition (HCM) after Deb convinces him to be there for Karen after Keith's death. Nathan and Haley prepare to renew their vows. Mouth tries to get closer to Rachel. Peyton gets an invitation to go to Chicago, from Pete. Brooke gets revenge on Rachael, by making posters of Rachael who got multiple surgeries to become attractive, because she was obese when she was 14.
This episode is named after a song by Fall Out Boy.
65
20 "Everyday Is a Sunday Evening" Billy Dickson Mark Schwahn April 19, 2006 (2006-04-19) 2T6170 2.67[21]
Nathan deals with a player who is a rival both on and off the court. Dan is given a restraining order from Deb and Nathan. Brooke and Mouth look to get even with Rachel, who has tricked Cooper into thinking she is 26 and a model. Dan has a run-in with Cooper. Peyton makes a major decision and goes to see Jake and Jenny who are in Savannah. At the end of the episode Peyton asks Jake to marry her.
This episode is named after an album by The Blackouts.
66
21 "Over the Hills and Far Away" Thomas J. Wright Mark Schwahn April 26, 2006 (2006-04-26) 2T6171 2.87[22]
Haley argues with Brooke about her wedding dress. Brooke plans an engagement party for Nathan and Haley. Karen and Lucas get back in town. Peyton is surprised by her own feelings when she mumble she loves Lucas in her sleep and Jake overhears.
This episode is named after a song by Led Zeppelin and another by Gary Moore which was covered by Nightwish.
67
22 "The Show Must Go On" Mark Schwahn Mark Schwahn May 3, 2006 (2006-05-03) 2T6172 3.06[23]
Nathan worries about Haley as they prepare to renew their vows. Deb confesses to Dan about the fire. Peyton and Brooke's friendship becomes strained when Peyton confesses she still loves Lucas. Nathan, Cooper and Rachel's lives hang in the balance after a car accident. Someone might be pregnant.
This episode is named after a song by Queen.
Ratings[edit]
Season
Timeslot
Season premiere
Season finale
TV season
Rank
Viewers
(in millions)
3rd
Wednesday 8/7C
October 5, 2005 May 3, 2006 2005–2006
#137
2.8[1]
DVD release[edit]
The DVD release of season three was released after the season has completed broadcast on television. It has been released in Regions 1, 2 and 4. As well as every episode from the season, the DVD release features bonus material such as audio commentaries on some episodes from the creator and cast, deleted scenes, gag reels and behind-the-scenes featurettes.
The Complete Third Season
Set details[24] Special features[24]
22 episodes
937 minutes (Region 1);[25] 968 minutes (Region 2);[26] 900 minutes (Region 4) [27]
6-disc set
1.78:1 aspect ratio
Languages:
English (Dolby Digital 2.0 Surround)
Subtitles:
Spanish and French (Region 1)
English, Danish, Finnish, Portuguese, Norwegian, Swedish, English for the Hearing Impaired (Regions 2 and 4) [27]
Audio commentaries:
"With Tired Eyes, Tired Minds, Tired Souls, We Slept" - with creator/executive producer/writer Mark Schwahn, executive producer Joe Davola, actors Colin Fickes, Sophia Bush, Lee Norris and music supervisor Lindsey Wolfington
"The Show Must Go On" - with creator/executive producer/writer/director Mark Schwahn, executive producer Joe Davola, actors Sophia Bush and Lee Norris
Deleted scenes:
Episodes: 2, 3, 4, 5, 9, 10, 14, 15, 16, 17, 20 & 22
"Anatomy of an Episode: A Behind-the-Scenes Look at the Creation of "With Tired Eyes, Tired Minds, Tired Souls, We Slept"
Gag reel
Release dates
United States United Kingdom Australia
September 26, 2006[24] October 23, 2006[26] July 4, 2007[27]
References[edit]
^ Jump up to: a b "Hollywood Reporter: 2005-06 primetime wrap". Hollywood Reporter. May 26, 2006. Archived from the original on January 14, 2007.
Jump up ^ Mark Schwahn (writer) & Gregory Prange (director) (2004-10-05). "Like You Like an Arsonist". One Tree Hill. Season 3. Episode 1. 42 minutes in. The WB.
Jump up ^ [1]
Jump up ^ [2]
Jump up ^ [3]
Jump up ^ [4]
Jump up ^ [5]
^ Jump up to: a b [6]
Jump up ^ [7]
Jump up ^ [8]
Jump up ^ [9]
Jump up ^ [10]
Jump up ^ "Archived copy". Archived from the original on 2014-07-28. Retrieved 2008-07-29.
Jump up ^ [11]
Jump up ^ [12]
Jump up ^ [13]
Jump up ^ Kissell, Rick (March 2, 2006). "Super sweeps sesh". Variety.com. Archived from the original on August 5, 2011. Retrieved August 6, 2011.
Jump up ^ [14]
Jump up ^ [15]
Jump up ^ [16]
Jump up ^ [17]
Jump up ^ [18]
Jump up ^ "Archived copy". Archived from the original on 2009-06-01. Retrieved 2009-09-23.
^ Jump up to: a b c "One Tree Hill - The Complete 3rd Season". TVShowsOnDVD.com. Retrieved 2008-02-09.
Jump up ^ "One Tree Hill - The Complete Third Season (2006)". Amazon.co.uk. Retrieved 2008-02-11.
^ Jump up to: a b "One Tree Hill - Series 3". Amazon.co.uk. Retrieved 2008-02-11.
^ Jump up to: a b c "One Tree Hill; Complete Season 3: M15 (DVD)". Sanity. Archived from the original on 2007-05-02. Retrieved 2008-02-11.
[hide]
v
t
e
One Tree Hill
Season 1
Season 2
Season 3
Season 4
Season 5
Season 6
Season 7
Season 8
Season 9
Characters
Lucas Scott
Nathan Scott
Peyton Sawyer
Haley James Scott
Brooke Davis
Mouth McFadden
Jamie Scott
Millicent Huxtable
Julian Baker
Clay Evans
Quinn James
Episodes
"Pilot"
"With Tired Eyes, Tired Minds, Tired Souls, We Slept"
"4 Years, 6 Months, 2 Days"
Related
Official soundtracks
"I Don't Want to Be"
Retrieved from "https://en.wikipedia.org/w/index.php?title=One_Tree_Hill_(season_3)&oldid=828346803"
Categories:
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