Marsupial

Taxonomy and naming

The scientific name Marsupialia derives from the Latin marsupium, meaning 'pouch', which is in turn borrowed from the ancient Greek word marsippos.^[7] The term was introduced in 1811 by the German zoologist Johann Karl Wilhelm Illiger, who initially erected Marsupialia as a family within his mammalian order Pollicata alongside several primate groups.^{[8][9][10][11]} In 1816, French anatomist Henri Marie Ducrotay de Blainville proposed the term Didelphia (meaning 'double womb') for marsupials, contrasting them with Monodelphia (placentals) and Ornithodelphia (monotremes), though his terminology was largely superseded.^[11][12] Later in 1816, Georges Cuvier recognized all marsupials within a single taxonomic order.^[8][13] In 1880, English biologist Thomas Henry Huxley introduced the term Metatheria ('beyond beasts') to designate a transitional evolutionary rank between Prototheria and Eutheria.^[11] Modern systematists generally treat Marsupialia as the crown group containing the last common ancestor of all living marsupials and its descendants, whereas Metatheria is used as a broader total clade that also incorporates stem taxa diverging before this common ancestor.^[14][15] In 1997, J. A. W. Kirsch and collaborators formalized infraclass rank for Marsupialia.^[13]

Historically, marsupial classification relied heavily on cranial and dental morphology, particularly the distinction between polyprotodont dentition (multiple unspecialized lower incisors) and diprotodont dentition (a single pair of enlarged, forward-pointing lower incisors), as well as syndactyly, the fusing of the second and third digits of the hindfoot inside a common skin sheath.^[16][17] During the 20th century, these traits caused taxonomic confusion because South American shrew opossums (Paucituberculata) share diprotodonty with Australian diprotodonts, while Australian bandicoots share syndactyly with diprotodonts.^[16] A serological investigation in 1977 ruled out a close phylogenetic affinity between Paucituberculata and Diprotodontia, demonstrating that both diprotodonty and syndactyly represent convergent evolutionary traits.^[18][19][20]

In 1982, Frederick S. Szalay divided marsupials into two grand clades: Ameridelphia (New World marsupials) and Australidelphia (Australian marsupials and their close relatives) based on the tarsal morphology of the ankle joints, specifically the articulation between the astragalus and calcaneus.^{[21][22][23][24]} Later morphological and molecular phylogenetics demonstrated that Ameridelphia is paraphyletic.^{[24][25][26][27]} The South American monito del monte (Dromiciops gliroides), placed in the order Microbiotheria, shares the derived astragalar-calcaneal joint structure of Australian marsupials, placing it inside Australidelphia rather than with the other South American lineages.^{[22][24][25][28][29]}

Living marsupials are categorized into seven orders containing approximately 21 extant families.^[16][30][31] The two American orders are Didelphimorphia, comprising American opossums with over 90 living species in the family Didelphidae, and Paucituberculata, comprising shrew opossums in the family Caenolestidae.^[16] Australidelphia includes the South American Microbiotheria (Microbiotheriidae, containing Dromiciops), alongside four exclusively Australasian orders: Notoryctemorphia (marsupial moles in Notoryctidae), Dasyuromorphia (carnivorous marsupials in Dasyuridae, Myrmecobiidae, and the extinct Thylacinidae), Peramelemorphia (bandicoots and bilbies in Peramelidae, Thylacomyidae, and the extinct Chaeropodidae), and Diprotodontia.^[16] Diprotodontia is subdivided into Vombatiformes (koalas and wombats), Phalangeriformes (cuscuses and possums), and Macropodiformes (kangaroos, wallabies, potoroos, and bettongs). Extinct orders recognized within Marsupialia or stem metatherians include Yalkaparidontia, Polydolopimorphia, and Sparassodonta.^[32][33]

Evolutionary history

The divergence between metatherians and eutherians is estimated from molecular clocks and fossil discoveries to have occurred during the Jurassic or Early Cretaceous, between 125 and 193 million years ago.^[34][35][36] The oldest metatherian fossils, dating to roughly 110 million years ago, are known from western North America.^[37][38] Sinodelphys szalayi, discovered in the Yixian Formation of Liaoning, China, and dated to 125 million years ago, was long described as the earliest metatherian fossil.^{[39][40][41][42]} However, subsequent reassessments, including comparisons with the eutherian Ambolestes zhoui, reinterpreted Sinodelphys as a member of Eutheria.^[37][38][43] Stem metatherian fossils are distinguished from eutherians primarily by dental characteristics, notably the presence of four molar pairs per jaw quadrant in metatherians compared to a maximum of three in eutherians.^[44][45][46] Basal Cretaceous metatherian radiations included Deltatheroida (such as Oklatheridium and Atokatheridium) and early Marsupialiformes such as Kokopellia and Asiatherium.^{[34][47][48][49]}

During the Late Cretaceous, metatherians were common and morphologically diverse across Laurasia, particularly in North America, where families like Stagodontidae (including the bone-crushing Didelphodon) and 'Alphadontidae' were abundant.^{[34][47][50][51][52]} Metatherians suffered severe extinctions at the Cretaceous-Paleogene boundary, undergoing greater declines than placentals or multituberculates.^[50][51][53] Stem metatherians survived into the Cenozoic in Europe, Asia, and North America, including members of Herpetotheriidae and Peradectidae, and briefly colonized northern Africa with Peratherium africanum during the Oligocene before disappearing from the Northern Hemisphere by the Miocene.^{[53][54][55][56][57]}

Metatherians spread to South America from North America, possibly via the Aves Ridge, though whether this happened in the Late Cretaceous or the early Paleocene is debated.^[58][59] The oldest metatherian fossils there date to the early Paleocene, about 64.5 to 63 million years ago.^[59][60] During its prolonged Cenozoic geographic isolation as an 'island continent', South America became a major radiation center.^[59][61] Opossums (Didelphimorphia) and shrew opossums (Paucituberculata) diversified alongside distinct lineages such as Polydolopimorphia.^[32][62] The family Argyrolagidae within Polydolopimorphia produced bipedal, hopping forms like Argyrolagus that converged anatomically with desert rodents such as jerboas.^[32] Large mammalian predatory niches in South America were occupied by Sparassodonta, including Borhyaena, the large Proborhyaena, and the sabertooth-like Thylacosmilus.^[61][63] Metatherians were restricted from large herbivorous niches in South America by native placental ungulates and xenarthrans.^[64][65] When the Isthmus of Panama formed approximately 3 million years ago during the Pliocene, the Great American Interchange initiated faunal mixing.^[61][66][67] Sparassodonts went extinct, while didelphid opossums successfully migrated northward, leading to the establishment of the Virginia opossum (Didelphis virginiana) across North America as far north as Canada.^{[62][66][68][69]}

Marsupials reached Australia via an Antarctic land bridge during the Early Eocene, around 50 to 55 million years ago, prior to the opening of the Drake Passage and the deep Tasman Gateway.^{[70][71][72][73][74]} At that time, Antarctica maintained a warm-temperate climate with Nothofagus forests.^[71][75] The oldest confirmed Australian marsupial fossil is Djarthia murgonensis from the 54.6-million-year-old Tingamarra local fauna in southeastern Queensland.^[76][77][78] Djarthia possesses plesiomorphic australidelphian tarsal morphology, supporting the hypothesis of an australidelphian radiation across Gondwana.^[76][79] A roughly 30-million-year gap occurs in the Australian fossil record between Tingamarra and the late Oligocene deposits at Riversleigh and central Australia.^[80][81] By the late Oligocene and early Miocene, all modern Australian orders, along with extinct groups like Yalkaparidontia, were already established in rainforested ecosystems.^[80][82]

As Australia drifted northward through the Neogene, the circumpolar current formed around Antarctica, leading to continental drying and the retreat of mesothermal rainforests from the late Miocene onward.^[83][84] The expansion of open grasslands stimulated the radiation of grazing Macropodidae.^[85] Repeated drops in sea level formed land bridges uniting Australia, New Guinea, and Tasmania into the continent Sahul, permitting marsupial dispersals into New Guinea and across the Weber Line into parts of Wallacea such as Sulawesi and the Moluccas.^[86][87] During the Pleistocene, Australia supported a megafauna including the rhinoceros-sized Diprotodon optatum (weighing up to 2.7 to 2.8 tons), giant short-faced kangaroos such as Procoptodon (reaching 2 to 3 meters in height), and the apex carnivore Thylacoleo carnifex ('marsupial lion'), which attained lengths of 1.5 meters and weights of 160 kilograms.^{[88][89][90][91][92]} This megafauna largely went extinct between 51,000 and 38,000 years ago, at roughly the time humans arrived in Australia; the cause is disputed, with some researchers blaming human hunting and others climatic change.^[90][93][94]

Anatomy and physiology

Marsupials vary widely in size and structural proportions. Modern forms range from the long-tailed planigale (Planigale ingrami), measuring 55 to 65 millimeters in head-body length and weighing approximately 4 to 5 grams, to the red kangaroo (Osphranter rufus), which reaches 1.4 to 1.8 meters in head-body length and weighs up to 85 to 90 kilograms.^{[95][96][97][98]} Extinct forms reached greater weights, exemplified by Diprotodon optatum at 2,700 to 2,800 kilograms.^[88][90][92] Sexual dimorphism is common, with males of many dasyurid and macropod species weighing up to twice as much as females; in contrast, female honey possums (Tarsipes rostratus) are larger than males.^[99] Convergence with placental body plans is observed across diverse families, producing marsupial equivalents of wolves (Thylacinidae), moles (Notoryctidae), flying squirrels (Petauridae and Acrobatidae), and burrowing rodents (Vombatidae).^{[89][100][101][102]}

Marsupial body temperatures average lower than those of similarly sized placentals, typically around 34 to 36 °C (with 35 °C or 35.5 °C frequently reported), representing a difference of 1.5 to 2.5 °C below placental norms.^{[103][104][105]} Daily torpor is widespread among small dasyurids, pygmy possums, and feathertail gliders to manage energy expenditure during cold and food shortages.^[106] The mountain pygmy possum (Burramys parvus) undergoes true seasonal hibernation in alpine eastern Australia for up to seven months, entering multiday bouts of deep torpor where its body temperature drops to roughly 2 °C.^[107] Physiological adaptations to aridity include water-conservation mechanisms in desert dwellers and the tolerance of saline sea water by the quokka (Setonix brachyurus) on Rottnest Island.^[108]

Skull and dentition

The marsupial skull possesses several unique cranial features distinguishing it from placental skulls. The braincase is relatively small and narrow.^{[3][109][110]} The nasal bones are expanded posteriorly, and the jugal bone is enlarged, extending backward to participate directly with the squamosal in forming the glenoid fossa for the mandibular articulation.^[109][110] The lacrimal foramen (foramen lacrimale) is positioned externally on the anterior rim of the orbit.^[109] The auditory bullae, when ossified, are formed predominantly by the alisphenoid bone rather than by the tympanic bone as in placentals.^{[110][111][112]} The bony hard palate consistently features large fenestrae or vacuities, except in the koala and the honey possum.^{[109][110][113]} The mandibular angle (processus angularis) is medially inflected beneath the jaw base, serving as an attachment site for the medial pterygoid muscle, a characteristic found in almost all marsupials except koalas and honey possums.^[109][110]

Dental architecture and replacement in marsupials differ from placental patterns. The primitive metatherian dental formula is 5.1.3.4 / 4.1.3.4, totaling 50 teeth per quadrant, characterized by five upper and four lower incisors, one canine, three premolars, and four molars.^{[109][114][115]} In contrast, the primitive placental formula is 3.1.4.3 / 3.1.4.3, totaling 44 teeth.^[114] Many marsupials retain 40 to 50 teeth, though tooth counts are reduced in specialized lineages.^{[109][114][115]} Tooth replacement is restricted entirely to the third premolar; all teeth anterior to the third premolar erupt directly as permanent teeth without a deciduous precursor.^[109][115]

Dental morphotypes reflect trophic specialization. Opossums retain generalized tribosphenic and dilambdodont molars suited for omnivory.^[116] Shrew opossums possess enlarged, procumbent lower incisors and specialized shearing blades.^[117] In Dasyuromorphia, dentition is modified for carnivory or insectivory: the extinct thylacine had a shearing dental set of 4.1.3.4 / 3.1.3.4 (46 teeth), while the termite-eating numbat has degenerate, simplified postcanines with the dental count increased to between 48 and 52 teeth.^[112][118] Diprotodontia is named for possessing a single pair of elongated, procumbent lower incisors matched against up to three upper incisor pairs.^{[110][119][120]} Grazing diprotodonts show continuous wear adaptations: wombats possess rootless, continuously growing (hypselodont) incisors and molars, while macropods display molar progression (polyphyodonty), where cheek teeth slowly drift forward along the jawline and shed as they wear down.^{[110][112][119][121]}

Postcranial skeleton and locomotion

The marsupial postcranial skeleton typically features epipubic bones (ossa epipubica) projecting forward from the pubic element of the pelvis.^[3][4][122] Present in both sexes and in pouchless taxa, these bones provide anchorage for abdominal muscles that support the trunk and aid hindlimb mechanics, rather than functioning primarily to support the marsupium.^[3][4][123] Epipubic bones are also present in monotremes and basal fossil mammals, representing a plesiomorphic mammalian feature that modern placentals have lost.^[3][4][123] Epipubic bones are reduced or absent in certain taxa, such as the extinct thylacine.^[124] Ossified patellae are generally absent in modern marsupials, although cartilaginous structures occur.^[125][126]

Locomotor adaptations correlate with foot architecture. Arboreal opossums, cuscuses, and pygmy possums retain a pentadactyl manus and pes, featuring an opposable hallux without a claw that aids grasping.^[122][127] In bandicoots and diprotodonts, the pes exhibits syndactyly, in which the second and third digits are bound within a common sheath of skin, projecting only small claws that function as a grooming comb.^[122][128] In terrestrial cursorial and saltatorial groups, the fourth toe is lengthened to form the primary axis of the foot, while the hallux is reduced or absent.^{[122][129][130]} Macropods exhibit narrow hindfeet with an elongated fourth metatarsal, elastic leg tendons, and a heavy muscular tail used as a counterweight during high-speed bipedal hopping, as well as a fifth support limb during slow pentapedal locomotion.^{[129][131][132]} Burrowing marsupial moles have spade-like claws on the third and fourth manual digits, whereas the extinct pig-footed bandicoot (Chaeropus ecaudatus) had reduced the manual digits to two functional hooves resembling those of artiodactyls.^[112][133] Gliding membranes (patagia) between the limbs evolved independently in three arboreal diprotodont lineages: Petauridae (sugar gliders), Acrobatidae (feathertail gliders), and Pseudocheiridae (greater gliders).^[101][134]

Nervous and sensory systems

Marsupial brains lack a corpus callosum, the principal commissural fiber tract connecting the neocortical hemispheres in placental mammals.^{[4][135][136]} Interhemispheric communication is instead mediated by an enlarged anterior commissure, whose fibers route through the internal capsule in diprotodonts and along the external margin of the striatum in bandicoots and dasyurids.^{[135][137][138]} Brain size relative to body mass is generally smaller than in comparably sized placentals, though relative neocortical volume is elevated in arboreal diprotodonts; the highest encephalization quotient among marsupials occurs in the striped possum (Dactylopsila trivirgata).^{[138][139][140]}

Marsupial photoreceptor architecture includes ancestral mammalian traits such as colored oil droplets within retinal cones and double cones, features that placentals have lost.^[141] Trichromatic color vision, historically considered unique to primates among mammals, has evolved independently in several marsupials, including the fat-tailed dunnart (Sminthopsis crassicaudata) and the honey possum (Tarsipes rostratus), based on three distinct cone photopigments sensitive to short, medium, and long wavelengths.^[142][143] Marsupial trichromacy is not linked to the X chromosome.^[143] In contrast, fossorial marsupial moles (Notoryctes) have vestigial, subcutaneously buried eyes that completely lack retinas, optic nerves, and oculomotor musculature.^[144] The sense of smell is macrosmatic in most species, accompanied by a functional vomeronasal organ and connected nasopalatine ducts.^[145][146] An accessory olfactory bulb is present in pygmy possums for detecting pheromonal cues.^[145] Hearing frequency sensitivity is generally narrower and exhibits higher minimum thresholds than in placentals, although northern quolls (Dasyurus hallucatus) demonstrate acute auditory detection.^[145]

Genetics and genomics

The marsupial karyotype typically comprises a low number of large chromosomes, exhibiting a bimodal distribution across taxa with diploid numbers of either 2n = 14 or 2n = 22.^[135][147] The ancestral metatherian karyotype is reconstructed as 2n = 14, a complement conserved among many living dasyurids and didelphids.^{[135][147][148]} Chromosomal reorganizations, fusions, and translocations occur among macropods; the swamp wallaby (Wallabia bicolor) possesses an atypical derived chromosome count of 2n = 10 in females (XX) and 2n = 11 in males (XY1Y2).^[148][149] Meiotic recombination frequencies in marsupials are among the lowest recorded across vertebrates, and females exhibit lower crossover rates than males.^[148]

Sex determination operates via an XY chromosomal system, but marsupial sex chromosomes lack the pseudoautosomal pairing regions found in placental mammals and do not form a synaptonemal complex during meiosis.^[148][150] Dosage compensation occurs through the non-random, preferential inactivation of the paternally derived X chromosome, mediated by the long non-coding RNA known as RSX rather than the placental Xist mechanism.^[148][150] Several peramelid genera display somatic sex-chromosome elimination, systematically ejecting one sex chromosome (the Y chromosome in males and an X chromosome in females) from cells of the liver, blood, and intestinal epithelium while retaining both within the germline.^[151] Marsupial nuclear genomes average 3.2 to 3.4 gigabases in size and harbor substantial proportions of transposable elements.^[135][152] The gray short-tailed opossum (Monodelphis domestica) was the first marsupial to have its complete nuclear genome sequenced.^[153]

Reproductive system and early development

The marsupial urogenital tract is bifurcated in both sexes.^{[4][154][155][156]} During embryonic development, the ureters pass medially between the Müllerian ducts, preventing their fusion along the midline into a single uterus and vagina as occurs in placentals.^[155] Females possess two separate lateral vaginas and two distinct uteri, each with its own cervix.^{[4][155][156][157]} The lateral vaginas transport spermatozoa following copulation.^[5][157] Parturition takes place through a central median vagina (pseudovaginal canal) that opens along the midline between the two lateral vaginas directly into the urogenital sinus.^[3][4][5] In most taxa, this birth canal is transitory and closes following delivery, but it persists as a permanent open passage in Macropodidae and the honey possum.^[4][158] Both the urogenital sinus and the rectum empty into a shallow cloaca.^{[155][159][160]}

Male marsupials possess an external scrotum located prepenially, situated on the abdomen anterior to the penis rather than posterior to it as in placentals.^{[3][161][162]} In fossorial marsupial moles, the testes remain permanently inside the abdominal cavity or inguinal canal.^[161] The penis is retracted in an S-shaped curve inside the urogenital sinus when flaccid, and curves forward when erect.^[154][163] The glans penis is bifurcated into two prongs corresponding to the female's two lateral vaginas in most families, but remains undivided in macropods and marsupial moles.^{[154][164][165][166][167]} Accessory sex glands are limited to the prostate and one to three pairs of bulbourethral glands; ampullae, seminal vesicles, and coagulating glands are absent.^{[168][169][170][171]} Sperm pairing, where spermatozoa align in pairs along their acrosomal surfaces during epididymal maturation, is unique to American didelphimorphs and paucituberculates, having been lost in Australidelphia.^[172]

Gestation and neonatal heterochrony

Gestation is exceptionally brief, ranging from 9.5 to 11 days in the stripe-faced dunnart (Sminthopsis macroura) to 38 days in the long-nosed potoroo.^{[3][173][174][175][176]} Marsupial blastocysts lack an inner cell mass and compaction, instead forming a hollow protoderm lining a fluid-filled vesicle surrounded by an uncalcified shell membrane that sheds during early development.^[177][178] The embryo is nourished primarily by a choriovitelline (yolk-sac) placenta that absorbs uterine secretions ('uterine milk').^{[5][179][180]} Peramelemorphs, koalas, and wombats develop an additional chorioallantoic placenta, though it lacks the chorionic villi characteristic of eutherian placentas.^{[5][179][180]} The marsupial placenta lacks the trophoblast cell layer that protects the embryo against the mother's immune system, and as a result gestation is very short.^[3][181]

Neonates are born in an altricial state, weighing less than 1% of the mother's body mass.^{[3][174][176][182]} Birth weights range from approximately 4 to 5 milligrams in the honey possum and planigales to roughly 800 milligrams in giant kangaroos.^{[174][176][183]} Neonatal development displays heterochrony: cranial facial bones, jaws, the shoulder girdle, and muscular forelimbs equipped with temporary deciduous claws are precociously developed, whereas the hindlimbs, eyes, ears, and cerebral hemispheres remain rudimentary buds.^{[184][185][186][187][188]} Internal organs are incompletely formed at birth, lacking alveolar lungs, separated ventricular cardiac septa, and glomerular kidneys.^[189] The newborn uses negative gravitropism and olfactory cues from saliva trails licked across the mother's fur to climb toward the pouch and teats.^{[176][190][191][192]} Joeys possess temporary soft-tissue oral shields that restrict the mouth opening to a circular aperture that fits over the teat, which then swells inside the mouth cavity to anchor the joey for continuous suckling.^{[166][193][194]}

Marsupium, lactation, and diapause

A permanent pouch (marsupium) is present in macropods, wombats, and koalas, but it is not a universal synapomorphy of all marsupials.^{[1][166][195]} Many dasyurids, didelphids, and shrew opossums lack a pouch entirely, possessing only lateral skin folds or bare mammary areas where the young cling exposed to maternal fur.^[166][195] Pouch orientation varies: saltatorial macropods have pouches opening anteriorly (upward), whereas burrowing wombats, bandicoots, and arboreal koalas possess pouches that open posteriorly (backward) or downward, preventing dirt entry during digging.^{[3][166][196]} The male water opossum (Chironectes minimus) possesses a functional pouch that encloses and protects his scrotum while swimming.^[3] Teat counts range from 2 in marsupial moles to up to 27 in certain short-tailed opossums.^[189][197]

Lactation represents the primary maternal investment in marsupials, extending over many months.^{[182][198][199]} Milk composition changes across lactation: early milk is dilute, high in carbohydrates, and low in lipids to sustain embryonic organogenesis, whereas late milk is dense, protein-rich, and high in fats.^[182][199] In kangaroos, asynchronous concurrent lactation enables a mother to simultaneously nurse two young of different ages from separate teats: an older out-of-pouch joey drinks high-fat milk from one teat while a newborn sibling drinks carbohydrate-rich milk from another.^[182][199]

Embryonic diapause occurs widely in macropods, pygmy possums, feathertail gliders, and honey possums.^[200][201] Following postpartum mating, the newly fertilized blastocyst arrests development at approximately the 70-to-100-cell stage while the prior joey suckles in the pouch.^[200][201] Suckling induces prolactin secretion, which suppresses the corpus luteum and blocks progesterone synthesis.^[200] When the pouch joey is weaned, dies, or vacates the teat, prolactin levels drop, the corpus luteum reactivates, and embryonic development resumes, allowing rapid production of replacement offspring without another mating event.^[200][201]

Ecology and behavior

Marsupials occupy nearly all terrestrial biomes across their geographic range, from arid deserts and spinifex grasslands to temperate woodlands, alpine peaks, and tropical rainforests.^[6] South and Central American species are mostly arboreal or ground-dwelling forest inhabitants across elevations from sea level to the Andean paramos, including shrew opossums (Caenolestes) on the forest floor, woolly opossums (Caluromys) in the high canopy, and the semiaquatic yapok (Chironectes minimus), which possesses webbed hindfeet and water-resistant fur.^{[6][127][202]} In Australia, dasyurids and bandicoots occupy arid hummock grasslands and heaths, rock wallabies (Petrogale) exploit cliff crevices, and tree-kangaroos (Dendrolagus) inhabit montane rainforests of northeastern Queensland and New Guinea.^{[2][203][204][205]} Feral populations of the brushtail possum (Trichosurus vulpecula), tammar wallaby, and red-necked wallaby have established introduced colonies in New Zealand and Great Britain.^{[206][207][208]}

Dietary strategies span all feeding guilds.^[209][210] American opossums and shrew opossums are dietary generalists, feeding on invertebrates, small vertebrates, fruits, and carrion.^[209][211] In Australia, Dasyuromorphia includes carnivores such as the Tasmanian devil (Sarcophilus harrisii) and quolls (Dasyurus), alongside small insectivorous planigales and dunnarts.^{[212][213][214]} The numbat (Myrmecobius fasciatus) is a specialized diurnal myrmecophage consuming 10,000 to 20,000 termites daily with its elongated vermiform tongue.^[214][215] Within Diprotodontia, specialized folivores include koalas and greater gliders, which subsist on fibrous, toxin-rich Eucalyptus foliage using an enlarged cecum for microbial fermentation; the koala possesses the largest cecum relative to body size of any mammal.^{[216][217][218]} Common ringtail possums (Pseudocheirus peregrinus) engage in caecotrophy to re-ingest nutrient-rich soft feces.^[216] Macropods use foregut fermentation in a chambered stomach akin to ruminants, while wombats rely on hindgut colon fermentation.^[218][219] Nectarivorous honey possums rely on brush-tipped tongues to extract nectar and pollen from Banksia and other blossoms.^[220]

Social organizations are generally solitary, but vary among taxa.^[3][221] Opossums, bandicoots, dasyurids, and koalas are solitary, associating only for breeding.^[3][221] However, greater glider pairs maintain shared home ranges, and feathertail gliders den communally in groups of several dozen.^[221] Large macropods form loose, dynamic groups called mobs, whose composition shifts through immigration and emigration without rigid hierarchies, although adult males establish dominance hierarchies through ritualized upright wrestling and kicking ('boxing') for access to estrous females.^[222][223] Whiptail wallabies (Notamacropus parryi) maintain complex social structures with linear male dominance hierarchies.^[3][222] Monogamy is rare, but occurs in the rock-haunting possum (Petropseudes dahli), in which mated pairs share parental guarding duties.^[224] Communication relies heavily on olfaction via secretions from sternal and cloacal scent glands, paracloacal glands, and urine.^{[136][222][225]} Vocalizations include coughing, hissing, guttural growls, and maternal clicks, while macropods and Tasmanian devils produce seismic foot-thumping warning signals.^{[222][226][227]}

Life histories span extreme r-selection to K-selection.^[228] Males of small dasyurids of the genus Antechinus mate with as many females as possible during a short, highly synchronized breeding season and die of exhaustion before reaching one year of age.^[229] In Stuart's antechinus, all males die within a two-week period after mating, as a sharp rise in testosterone during mating disrupts their immune system.^[229][230] In contrast, herbivorous marsupials typically give birth to a single young and most live five years or more, while larger herbivores nurse their young for more than a year.^[231][232] Kangaroos can live more than 20 years and the common wombat up to 30 years, although lifespans in the wild are usually much shorter.^[3][233]

Diseases and pathology

Captive and wild macropods are susceptible to necrobacillosis, commonly termed 'lumpy jaw', an osteomyelitis of the mandible and surrounding tissues caused by oral bacterial infection with agents including Fusobacterium necrophorum; if untreated, systemic dissemination causes mortality.^{[234][235][236]} White muscle disease, a degenerative nutritional myopathy linked to vitamin E or selenium deficiencies, causes progressive limb paralysis and mortality in captive macropods, particularly quokkas.^[237]

Wild koala populations are heavily afflicted by Chlamydia pecorum and Chlamydia pneumoniae infections, which cause keratoconjunctivitis leading to blindness, urinary tract infections, and reproductive tract inflammation that causes female infertility.^[238] Tasmanian devils suffer from Devil Facial Tumour Disease (DFTD), an infectious, transmissible clonal cancer spread through biting during social and reproductive encounters; the tumors cause 100% mortality in affected individuals within a year of symptom onset, devastating wild populations.^{[239][240][241][242]} Bare-nosed wombats (Vombatus ursinus) and southern hairy-nosed wombats (Lasiorhinus latifrons) suffer high mortality from sarcoptic mange caused by the mite Sarcoptes scabiei, which causes severe hyperkeratosis, parakeratosis, emaciation, and secondary infection.^[243][244]

Human interactions and conservation

Aboriginal Australians and Papuans have interacted with marsupials for at least 45,000 to 60,000 years, depicting them in rock art and integrating them into Dreamtime traditions, while utilizing them as staple sources of meat, skins, and tools.^{[245][246][247]} European encounters began in 1500 when Vicente Yáñez Pinzón collected a female opossum with pouch young on the coast of Brazil and presented it to Queen Isabella I and King Ferdinand II of Spain.^{[245][248][249]} In Australasia, the Portuguese administrator António Galvão documented the northern common cuscus (Phalanger orientalis) on the Moluccas between 1536 and 1540, while a 1606 record of an animal killed on the southern coast of New Guinea described what appears to be the dusky pademelon (Thylogale brunii).^{[248][249][250]} Systematic European scientific collection commenced following James Cook's 1770 expedition.^[245][251] Early colonists frequently classified marsupials under placental names (such as 'marsupial cat', 'native cat', and 'Tasmanian wolf'), and the German naturalist Johann Christian Polycarp Erxleben initially described the eastern grey kangaroo as a jumping rodent in 1777.^[251][252]

Following European settlement of Australia, agricultural development and predator introduction triggered widespread mammalian declines.^{[253][254][255]} European colonists viewed marsupials as agricultural pests or livestock predators: the New South Wales government passed the Pastures and Stock Protection Act in 1880, classifying all kangaroos, wallaroos, wallabies, and pademelons as vermin, leading to the bounty slaughter of an estimated 21.4 million macropods in New South Wales alone between 1888 and 1900.^[256] Persecution by bounty hunting eliminated the thylacine (Thylacinus cynocephalus), the last known captive individual dying at the Hobart Zoo in 1936.^{[254][255][257]} The introduction of placental predators, namely red foxes (Vulpes vulpes) and feral domestic cats (Felis catus), and of ecological competitors such as European rabbits, sheep, and cattle devastated native populations.^{[253][258][259][260]} Species in the critical weight range of 35 to 5,500 grams in arid and semi-arid zones suffered the highest extinction rates.^[253]

According to assessments published by the International Union for Conservation of Nature (IUCN), 349 living and recently extinct marsupial species are classified across threat categories: 194 species (55.6%) are Least Concern, 41 (11.8%) are Near Threatened, 43 (12.3%) are Vulnerable, 22 (6.3%) are Endangered, 17 (4.9%) are Critically Endangered, 13 (3.7%) are Extinct, and 19 (5.4%) are Data Deficient.^[261] In Australia, extinctions include the pig-footed bandicoot (Chaeropus ecaudatus), desert bandicoot (Perameles eremiana), lesser bilby (Macrotis leucura), crescent nail-tail wallaby (Onychogalea lunata), and broad-faced potoroo (Potorous platyops).^[253][255] In South America, the Red-bellied gracile opossum (Cryptonanus ignitus) from Jujuy Province, Argentina, represents the only modern extinct marsupial recognized by the IUCN, having vanished following agricultural destruction of its forest habitat.^[262] Modern conservation strategies rely heavily on predator-free mainland fenced reserves and translocations to offshore islands.^[263] Translocation efforts assisted the bridled nail-tail wallaby (Onychogalea frenata), which was rediscovered in 1973 and stabilized as a Vulnerable species by 2016.^[263][264]

References

1. Pallardy, Richard (4 April 2016). "List of Marsupials". Britannica. Also written and edited by Britannica Editors. Retrieved 6 April 2026.
2. Nowak 2005, s. 21–22. (NOWAK, Ronald, 2005. Walker's Marsupials of the World. Baltimore: Johns Hopkins University Press. Dostupné online. ISBN 9780801882227, ISBN 9780801882111. (anglicky))
3. Wund, M. & P. Myers. 2006 Infraclass Metatheria - marsupial mammals, Animal Diversity Web (engelska), läst 29 november 2010
4. Nowak 1999. (Nowak RM (7 April 1999). Walker's Mammals of the World. JHU Press. ISBN 978-0-8018-5789-8.)
5. POUGH, F. Harvey; JANIS, Christine M. Vertebrate life. 10. vyd. New York Oxford: Sinauer Associates, Oxford University Press 552 s. ISBN 978-1-60535-721-8, ISBN 978-1-60535-607-5. S. 498, 500.
6. Armati, Dickman a Hume 2006, s. 205–207. (ARMATI, P. J.; DICKMAN, C. R.; HUME, I. D, 2006. Marsupials. New York: Cambridge University Press,. Dostupné online. (anglicky))
7. Nowak, Marsupials of the World, 2. (Nowak, Ronald M. (2005). Walker's Marsupials of the World (in English). Baltimore: Johns Hopkins University Press. ISBN 0-8018-8222-2.)
8. Martin WC (1841). A General Introduction to the Natural History of Mammiferous Animals. London, UK: Wright and Co. Printers. pp. 182–4.
9. JACKSON, S.; GROVES, C. Taxonomy of Australian Mammals. Melbourne: CSIRO Publishing, 2015. ISBN 978-1-4863-0013-6, ISBN 978-1-4863-0014-3. S. 30. (anglicky)
10. ILLIGER, Johann Karl Wilhelm. Caroli Illigeri ... Prodromus systematis mammalium et avium additis terminis zoographicis utriusque classis, eorumque versione germanica. Berolini [Berlin]: sumptibus C. Salfeld 330 s. Dostupné online. doi:10.5962/bhl.title.106965. (latinsky)
11. Tyndale-Biscoe, H. (2005) sid.5
12. Zima a Macholán 2021, s. 95. (ZIMA, J.; MACHOLÁN, M, 2021. Systém a fylogeneze savců. 1. vyd. Praha: Academia. 570 s. ISBN 978-80-200-3215-7, ISBN 80-200-3215-0.)
13. Jackson S, Groves C (2015). Taxonomy of Australian Mammals. Australia: CSIRO Publishing. pp. 82–3. ISBN 978-1-4863-0014-3.
14. Nowak 2005, s. 2–3. (NOWAK, Ronald, 2005. Walker's Marsupials of the World. Baltimore: Johns Hopkins University Press. Dostupné online. ISBN 9780801882227, ISBN 9780801882111. (anglicky))
15. Zima a Macholán 2021, s. 101–103. (ZIMA, J.; MACHOLÁN, M, 2021. Systém a fylogeneze savců. 1. vyd. Praha: Academia. 570 s. ISBN 978-80-200-3215-7, ISBN 80-200-3215-0.)
16. Feldhamer 2015, s. 251. (FELDHAMER, George A. & kol, 2015. Mammalogy – Adaptation, Diversity, Ecology. 4. vyd. Baltimore: Johns Hopkins University Press. (anglicky))
17. Moeller, 1990; Nowak, 1991; Vaughan, Ryan, and Czaplewski, 2000
18. Kemp 2005, s. 193. (KEMP, Thomas Stainforth, 2005. The Origin and Evolution of Mammals. Oxford ; New York: Oxford University Press. (anglicky))
19. KIRSCH, J. A. W., 1977. The comparative serology of Marsupialia, and a classification of marsupials. Australian Journal of Zoology Supplementary Series. Roč. 25, čís. 52, s. 1–152. Dostupné online [cit. 2023-07-10]. doi:10.1071/ajzs052. (anglicky)
20. Zima a Macholán 2021, s. 119. (ZIMA, J.; MACHOLÁN, M, 2021. Systém a fylogeneze savců. 1. vyd. Praha: Academia. 570 s. ISBN 978-80-200-3215-7, ISBN 80-200-3215-0.)
21. SZALAY, F. S, 1982. A new appraisal of marsupial phylogeny and classification. In: ARCHER, M. Carnivorous Marsupials. Sydney: Royal Zoological Society of New South Wales. Dostupné online. S. 621–640. (anglicky)
22. Benton 2015, s. 347. (BENTON, Michael, 2015. Vertebrate palaeontology. 4. vyd. Chichester, West Sussex ; Hoboken: Blackwell Publishers. Dostupné online. (anglicky))
23. Gaisler a Zima 2007, s. 572. (GAISLER, Jiří; ZIMA, Jan, 2007. Zoologie obratlovců. 2. vyd. Praha: Academia. ISBN 978-80-200-1484-9.)
24. Amrine-Madsen H.; Scally M.; Westerman M.; Stanhope M.J.; Krajewski C.; Springer M.S. (2003) Nuclear gene sequences provide evidence for the monophyly of australidelphian marsupials. Molecular Phylogenetics and Evolution, 28(2), 186-196
25. Zima a Macholán 2021, s. 107–108. (ZIMA, J.; MACHOLÁN, M, 2021. Systém a fylogeneze savců. 1. vyd. Praha: Academia. 570 s. ISBN 978-80-200-3215-7, ISBN 80-200-3215-0.)
26. Burgin C. J., Widness J., Upham N. S. Introduction to Illustrated Checklist of the Mammals of the World // Illustrated Checklist of the Mammals of the World (англ.) / ed. by Burgin C. J., Wilson D. E., Mittermeier R. A., Rylands A. B., Lacher T. E., Sechrest W. — Lynx Edicions, 2020. — P. 27. — ISBN 978-84-16728-36-7.
27. Horovitz I., Sánchez-Villagra M. R. A morphological analysis of marsupial mammal higher-level phylogenetic relationships (англ.) // Cladistics : journal. — 2003. — Vol. 19, iss. 3. — P. 181—212. — ISSN 1096-0031. — doi:10.1111/j.1096-0031.2003.tb00363.x. Архивировано 13 июня 2021 года.
28. Szalay F (1982). Archer M (ed.). "A new appraisal of marsupial phylogeny and classification". Carnivorous Marsupials. 2: 621–40.
29. Asher, R.J.; Horovitz, I.; Sanchez-Villagra, M.R. (2004) First combined cladistic analysis of marsupial mammal interrelationships. Molecular Phylogenetics And Evolution, 33(1), 240-250.
30. "Mammals". vertlife.org. Retrieved 12 November 2024.
31. Gardner, A. (2005). Wilson, D.E.; Reeder, D.M. (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. pp. 3–21. ISBN 978-0-8018-8221-0. OCLC 62265494.
32. Benton 2015, s. 349. (BENTON, Michael, 2015. Vertebrate palaeontology. 4. vyd. Chichester, West Sussex ; Hoboken: Blackwell Publishers. Dostupné online. (anglicky))
33. Kemp 2005, s. 210–212. (KEMP, Thomas Stainforth, 2005. The Origin and Evolution of Mammals. Oxford ; New York: Oxford University Press. (anglicky))
34. WILLIAMSON, Thomas E.; BRUSATTE, Stephen L.; WILSON, Gregory P. The origin and early evolution of metatherian mammals: the Cretaceous record. ZooKeys. 2014-12-17, roč. 465, s. 1–76. Dostupné online [cit. 2023-07-11]. ISSN 1313-2970. doi:10.3897/zookeys.465.8178. PMID 25589872.
35. DOS REIS, Mario; INOUE, Jun; HASEGAWA, Masami. Phylogenomic datasets provide both precision and accuracy in estimating the timescale of placental mammal phylogeny. Proceedings of the Royal Society B: Biological Sciences. 2012-09-07, roč. 279, čís. 1742, s. 3491–3500. Dostupné online [cit. 2023-07-11]. ISSN 0962-8452. doi:10.1098/rspb.2012.0683. PMID 22628470. (anglicky)
36. van Rheede T., Bastiaans T., Boone D.N. et al. The platypus is in its place: nuclear genes and indels confirm the sister group relation of monotremes and Therians // Molecular Biology and Evolution. — 2005. — doi:10.1093/molbev/msj064. — PMID 16291999.
37. Bi S, Zheng X, Wang X, Cignetti NE, Yang S, Wible JR (June 2018). "An Early Cretaceous eutherian and the placental-marsupial dichotomy". Nature. 558 (7710): 390–395. Bibcode:2018Natur.558..390B. doi:10.1038/s41586-018-0210-3. PMID 29899454. S2CID 49183466.
38. BI, Shundong; ZHENG, Xiaoting; WANG, Xiaoli. An Early Cretaceous eutherian and the placental–marsupial dichotomy. Nature. 2018-06, roč. 558, čís. 7710, s. 390–395. Dostupné online [cit. 2023-07-11]. ISSN 1476-4687. doi:10.1038/s41586-018-0210-3. (anglicky)
39. Rincon P (12 December 2003). "Oldest Marsupial Ancestor Found". BBC News. Retrieved 16 March 2010.
40. Luo ZX, Ji Q, Wible JR, Yuan CX (December 2003). "An Early Cretaceous tribosphenic mammal and metatherian evolution". Science. 302 (5652): 1934–1940. Bibcode:2003Sci...302.1934L. doi:10.1126/science.1090718. PMID 14671295. S2CID 18032860.
41. LUO, Zhe-Xi; JI, Qiang; WIBLE, John R. An Early Cretaceous Tribosphenic Mammal and Metatherian Evolution. Science. 2003-12-12, roč. 302, čís. 5652, s. 1934–1940. Dostupné online [cit. 2023-07-11]. ISSN 0036-8075. doi:10.1126/science.1090718. (anglicky)
42. Ji et al, “The earliest known eutherian mammal”. (Ji, Qiang; Luo, Zhe-Xi; Yuan, Chong-Xi; Wible, John R.; Zhang, Jian-Ping; Georgi, Justin A. (2002). "The earliest known eutherian mammal". Nature (in English). 416: 816–822. doi:10.1038/416816a.)
43. Phys.Org
44. Benton MJ (1997). Vertebrate Palaeontology. London: Chapman & Hall. p. 306. ISBN 978-0-412-73810-4.
45. Cáceras a Dickman 2023, s. 31–32. (CÁCERAS, Nilton C.; DICKMAN, C. R, 2023. American and Australasian Marsupials – An Evolutionary, Biogeographical, and Ecological Approach. Cham: Springer. ISBN 978-3-031-08418-8, ISBN 978-3-031-08419-5. (anglicky))
46. Benton, Michael J. (1997). Vertebrate Palaeontology. London: Chapman & Hall. p. 306. ISBN 0-412-73810-4.
47. 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.
48. CIFELLI, Richard L.; DE MUIZON, Christian. Dentition and Jaw of Kokopellia juddi, a Primitive Marsupial or Near-Marsupial from the Medial Cretaceous of Utah. Journal of Mammalian Evolution. 1997-12-01, roč. 4, čís. 4, s. 241–258. Dostupné online [cit. 2023-07-13]. ISSN 1573-7055. doi:10.1023/A:1027394430433. (anglicky)
49. Rose 2006, s. 76. (ROSE, Kenneth D, 2006. The Beginning of the Age of Mammals. Baltimore: The Johns Hopkins University Press. (anglicky))
50. Bennett, C. Verity; Upchurch, Paul; Goin, Francisco J.; Goswami, Anjali (6 February 2018). "Deep time diversity of metatherian mammals: implications for evolutionary history and fossil-record quality". Paleobiology. 44 (2): 171–198. Bibcode:2018Pbio...44..171B. doi:10.1017/pab.2017.34. hdl:11336/94590. ISSN 0094-8373. S2CID 46796692.
51. Zima a Macholán 2021, s. 104. (ZIMA, J.; MACHOLÁN, M, 2021. Systém a fylogeneze savců. 1. vyd. Praha: Academia. 570 s. ISBN 978-80-200-3215-7, ISBN 80-200-3215-0.)
52. Kemp 2005, s. 196–198. (KEMP, Thomas Stainforth, 2005. The Origin and Evolution of Mammals. Oxford ; New York: Oxford University Press. (anglicky))
53. Benton 2015, s. 346. (BENTON, Michael, 2015. Vertebrate palaeontology. 4. vyd. Chichester, West Sussex ; Hoboken: Blackwell Publishers. Dostupné online. (anglicky))
54. Feldhamer 2015, s. 249–250. (FELDHAMER, George A. & kol, 2015. Mammalogy – Adaptation, Diversity, Ecology. 4. vyd. Baltimore: Johns Hopkins University Press. (anglicky))
55. Russel, E. M. (1996) sid. 96
56. Vicente D. Crespo, Francisco J. Goin und Martin Pickford: The last African metatherian. Fossil Record 25 (1), 2022, S. 173–186, doi:10.3897/fr.25.80706.
57. Marc Furió, Francisco J. Ruiz-Sánchez, Vicente D. Crespo, Matthijs Freudenthal und Plinio Montoya: The southernmost Miocene occurrence of the last European herpetotheriid Amphiperatherium frequens (Metatheria, Mammalia). Comptes Rendus Palevol 11 (5), 2012, S. 371–377, doi:10.1016/j.crpv.2012.01.004.
58. Kemp, Thomas Stainforth (2005). The origin and evolution of mammals (PDF). Oxford: Oxford University Press. p. 217. ISBN 0-19-850760-7.
59. Black et al. 2012, s. 987. (BLACK, K. H.; ARCHER, M.; HAND, S. L., 2012. The Rise of Australian Marsupials: A Synopsis of Biostratigraphic, Phylogenetic, Palaeoecologic and Palaeobiogeographic Understanding. In: TALENT, J. A. Earth and Life. [s.l.]: Springer. Dostupné online. (anglicky))
60. Armati, Dickman and Hume, Marsupials, 17. (Armati, Patricia; Dickman, Chris; Hume, Ian, eds. (2006). Marsupials (in English). Cambridge University Press. ISBN 978-0-521-65074-8.)
61. Dickman, A Fragile Balance, 28. (Dickman, Christopher (2008). A Fragile Balance: The Extraordinary Story of Australian Marsupials (in English). University Of Chicago Press. ISBN 978-0226146300.)
62. SÁNCHEZ-VILLAGRA, Marcelo; LADEVÈZE, Sandrine; HOROVITZ, Inés. Exceptionally preserved North American Paleogene metatherians: adaptations and discovery of a major gap in the opossum fossil record. Biology Letters. 2007-06-22, roč. 3, čís. 3, s. 318–322. Dostupné online [cit. 2023-07-14]. ISSN 1744-9561. doi:10.1098/rsbl.2007.0090. PMID 17426007. (anglicky)
63. Vaughan, Ryan a Czaplawski 2015, s. 85–86. (VAUGHAN, Terry A.; RYAN, James M.; CZAPLAWSKI, Nicholas J, 2015. Mammalogy. 6. vyd. Burlington, MA: Jones & Bartlett Learning. (anglicky))
64. Simpson GG (July 1950). "History of the Fauna of Latin America". American Scientist. 38 (3): 361–389, see p. 368. JSTOR 27826322. Archived from the original on 15 June 2009. Retrieved 21 January 2020.
65. Stonehouse and Gilmore, The Biology of Marsupials, 62. (Stonehouse, Bernard; Gilmore, Desmond (1977). The Biology of Marsupials (in English). University Park Press. ISBN 0-8391-0852-4.)
66. Prevosti FJ, Forasiepi A, Zimicz N (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. hdl:11336/2663. S2CID 15751319.
67. Feldhamer 2015, s. 250. (FELDHAMER, George A. & kol, 2015. Mammalogy – Adaptation, Diversity, Ecology. 4. vyd. Baltimore: Johns Hopkins University Press. (anglicky))
68. Anděra 1997, s. 22. (ANDĚRA, Miloš, 1997. Savci (1). Praha: Albatros. (Svět zvířat; sv. 1). ISBN 80-00-00541-7.)
69. Cuarón, A.D. et al. 2008 Didelphis virginiana Från: IUCN 2010. IUCN Red List of Threatened Species. Version 2010.4. <www.iucnredlist.org>. Läst 30 januari 2011.
70. Bond M, Reguero MA, Vizcaíno SF, Marenssi SA (2006). "A new 'South American ungulate' (Mammalia: Litopterna) from the Eocene of the Antarctic Peninsula". In Francis JE, Pirrie D, Crame JA (eds.). Cretaceous-tertiary high-latitude palaeoenvironments: James Ross Basin, Antarctica. Geological Society, London, Special Publications. Vol. 258. The Geological Society of London. pp. 163–176. Bibcode:2006GSLSP.258..163B. doi:10.1144/GSL.SP.2006.258.01.12. S2CID 140546667.
71. Kemp 2005, s. 218. (KEMP, Thomas Stainforth, 2005. The Origin and Evolution of Mammals. Oxford ; New York: Oxford University Press. (anglicky))
72. Benton 2015, s. 346–347. (BENTON, Michael, 2015. Vertebrate palaeontology. 4. vyd. Chichester, West Sussex ; Hoboken: Blackwell Publishers. Dostupné online. (anglicky))
73. Sanmartín I. & Ronquist F.(2004)Southern Hemisphere Biogeography Inferred by Event-Based Models: Plant versus Animal Patterns. Systematic Biology, 53(2),216–243
74. Lillegraven, J.A.(1974). Biological Considerations of the Marsupial-Placental Dichotomy. Annual Review of Ecology and Systematics, 5, 263-283
75. Inman, M. (2010). Jumping Genes Reveal Kangaroos' Origins. PLoS Biology 8 (7): e1000437. DOI:10.1371/journal.pbio.1000437.
76. Beck RM, Godthelp H, Weisbecker V, Archer M, Hand SJ (March 2008). Hawks J (ed.). "Australia's oldest marsupial fossils and their biogeographical implications". PLOS ONE. 3 (3) e1858. Bibcode:2008PLoSO...3.1858B. doi:10.1371/journal.pone.0001858. PMC 2267999. PMID 18365013.
77. Black et al. 2012, s. 988. (BLACK, K. H.; ARCHER, M.; HAND, S. L., 2012. The Rise of Australian Marsupials: A Synopsis of Biostratigraphic, Phylogenetic, Palaeoecologic and Palaeobiogeographic Understanding. In: TALENT, J. A. Earth and Life. [s.l.]: Springer. Dostupné online. (anglicky))
78. Karen H. Black, Michael Archer, Suzanne J. Hand, Henk Godthelp: The Rise of Australian Marsupials: A Synopsis of Biostratigraphic, Phylogenetic, Palaeoecologic and Palaeobiogeographic Understanding. In: J. A. Talent (Hrsg.): Earth and Life. International Year of Planet Earth, 2012, S. 983–1078, doi:10.1007/978-90-481-3428-1_35, ISBN 978-90-481-3427-4.
79. Beck R.M.D. et al. (2008) Australia’s Oldest Marsupial Fossils and their Biogeographical Implications. PLoS ONE 3(3): e1858. doi:10.1371/journal.pone.0001858
80. Black et al. 2012, s. 998. (BLACK, K. H.; ARCHER, M.; HAND, S. L., 2012. The Rise of Australian Marsupials: A Synopsis of Biostratigraphic, Phylogenetic, Palaeoecologic and Palaeobiogeographic Understanding. In: TALENT, J. A. Earth and Life. [s.l.]: Springer. Dostupné online. (anglicky))
81. Armati, P. (2006) sid. 19
82. Black et al. 2012, s. 1004. (BLACK, K. H.; ARCHER, M.; HAND, S. L., 2012. The Rise of Australian Marsupials: A Synopsis of Biostratigraphic, Phylogenetic, Palaeoecologic and Palaeobiogeographic Understanding. In: TALENT, J. A. Earth and Life. [s.l.]: Springer. Dostupné online. (anglicky))
83. Kemp 2005, s. 221. (KEMP, Thomas Stainforth, 2005. The Origin and Evolution of Mammals. Oxford ; New York: Oxford University Press. (anglicky))
84. Black et al. 2012, s. 1028. (BLACK, K. H.; ARCHER, M.; HAND, S. L., 2012. The Rise of Australian Marsupials: A Synopsis of Biostratigraphic, Phylogenetic, Palaeoecologic and Palaeobiogeographic Understanding. In: TALENT, J. A. Earth and Life. [s.l.]: Springer. Dostupné online. (anglicky))
85. CELIK, Mélina; CASCINI, Manuela; HAOUCHAR, Dalal. A molecular and morphometric assessment of the systematics of the Macropus complex clarifies the tempo and mode of kangaroo evolution. Zoological Journal of the Linnean Society. 2019-06-25, roč. 186, čís. 3, s. 793–812. Dostupné online [cit. 2023-07-19]. ISSN 0024-4082. doi:10.1093/zoolinnean/zlz005. (anglicky)
86. Zima a Macholán 2021, s. 111. (ZIMA, J.; MACHOLÁN, M, 2021. Systém a fylogeneze savců. 1. vyd. Praha: Academia. 570 s. ISBN 978-80-200-3215-7, ISBN 80-200-3215-0.)
87. Anděra 1997, s. 19. (ANDĚRA, Miloš, 1997. Savci (1). Praha: Albatros. (Svět zvířat; sv. 1). ISBN 80-00-00541-7.)
88. Wroe, S.; Crowther, M.; Dortch, J.; Chong, J. (2004). "The size of the largest marsupial and why it matters". Proceedings of the Royal Society B: Biological Sciences. 271 (Suppl 3): S34–S36. Bibcode:2004PBioS.271.0095W. doi:10.1098/rsbl.2003.0095. PMC 1810005. PMID 15101412.
89. Benton 2015, s. 348. (BENTON, Michael, 2015. Vertebrate palaeontology. 4. vyd. Chichester, West Sussex ; Hoboken: Blackwell Publishers. Dostupné online. (anglicky))
90. LONG, John. Původ zvláštní australské fauny. In: BENTON, Michael. Sedmdesát velkých záhad světa přírody. Praha: Slovart, 2009. ISBN 978-80-7391-123-2. S. 162–166.
91. Ice Age Marsupial Topped Three Tons, Scientists Say, National Geographic, 2003-09-17. läst 2003-09-17.
92. Nowak R. M. Walker's marsupials of the world. — JHU Press, 2005. — С. 1. — ISBN 0801882222. (англ.)
93. GRUBER, K. Humans killed most of Australia's megafauna: study. Australian Geographic [online]. 2017-01-24 [cit. 2023-08-12]. Dostupné online. (anglicky)
94. WESTAWAY, M.; OLLEY, J.; GRUN, R. Aboriginal Australians co-existed with megafauna for at least 17,000 years. Australian Geographic [online]. 2017-01-12 [cit. 2023-08-12]. Dostupné online. (anglicky)
95. Baker, Andrew M. (27 April 2021). "Meet 5 of Australia's tiniest mammals, who tread a tightrope between life and death every night". The Conversation. Retrieved 29 May 2024.
96. McDade 2003a, s. 298. (MCDADE, M. C. & kol, 2003a. Grzimek’s Animal Life Encyclopedia – Volume 12 Mammals I. 2. vyd. Farmington Hills, MI: Gale. (anglicky))
97. McDade 2003b, s. 95. (MCDADE, M. C. & kol, 2003b. Grzimek’s Animal Life Encyclopedia – Volume 13 Mammals II. 2. vyd. Farmington Hills, MI: Gale. (anglicky))
98. Nowak, R. M. (1999) s. 119 online
99. Nowak 2005, s. 27. (NOWAK, Ronald, 2005. Walker's Marsupials of the World. Baltimore: Johns Hopkins University Press. Dostupné online. ISBN 9780801882227, ISBN 9780801882111. (anglicky))
100. Todorov OS, Blomberg SP, Goswami A, Sears K, Drhlík P, Peters J, Weisbecker V (March 2021). "Testing hypotheses of marsupial brain size variation using phylogenetic multiple imputations and a Bayesian comparative framework". Proceedings. Biological Sciences. 288 (1947) 20210394. doi:10.1098/rspb.2021.0394. PMC 8059968. PMID 33784860.
101. Todorov OS (2019). "Marsupial Cognition". In Vonk J, Shackelford T (eds.). Encyclopedia of Animal Cognition and Behavior. Cham: Springer International Publishing. pp. 1–8. doi:10.1007/978-3-319-47829-6_1167-1. ISBN 978-3-319-47829-6. S2CID 242256517.
102. Zima a Macholán 2021, s. 112–113. (ZIMA, J.; MACHOLÁN, M, 2021. Systém a fylogeneze savců. 1. vyd. Praha: Academia. 570 s. ISBN 978-80-200-3215-7, ISBN 80-200-3215-0.)
103. White (1999). "Thermal Biology of the Platypus". Davidson College. Archived from the original on 6 March 2012. Retrieved 14 September 2006.
104. "Control Systems Part 2" (PDF). Archived from the original (PDF) on 8 October 2016. Retrieved 6 July 2016.
105. Tyndale-Biscoe, H. (2005) sid.9
106. JONES, Caroline J.; GEISER, Fritz. Prolonged and daily torpor in the feathertail glider, Acrobates pygmaeus (Marsupialia: Acrobatidae). Journal of Zoology. 1992-05, roč. 227, čís. 1, s. 101–108. Dostupné online [cit. 2023-08-07]. ISSN 0952-8369. doi:10.1111/j.1469-7998.1992.tb04347.x. (anglicky)
107. Nowak 2018, s. 239–240. (NOWAK, Ronald, 2018. Walker's Mammals of the World : Monotremes, Marsupials, Afrotherians, Xenarthrans, and Sundatherians. Baltimore; London: Johns Hopkins University Press. Dostupné online. (anglicky))
108. TURNER, James. Mammals of Australia: An Introduction to Their Classification, Biology & Distribution. Sofia-Moscow: Pensoft, 2004. ISBN 954-642-198-7. S. 149.
109. Beck, Robin M.D.; Voss, Robert S.; Jansa, Sharon A. (28 June 2022). "Craniodental Morphology and Phylogeny of Marsupials". Bulletin of the American Museum of Natural History. 457 (1). Bibcode:2022BAMNH.4577.1.1B. doi:10.1206/0003-0090.457.1.1. ISSN 0003-0090.
110. Nowak 2005, s. 9–10. (NOWAK, Ronald, 2005. Walker's Marsupials of the World. Baltimore: Johns Hopkins University Press. Dostupné online. ISBN 9780801882227, ISBN 9780801882111. (anglicky))
111. Vaughan, Ryan a Czaplawski 2015, s. 80. (VAUGHAN, Terry A.; RYAN, James M.; CZAPLAWSKI, Nicholas J, 2015. Mammalogy. 6. vyd. Burlington, MA: Jones & Bartlett Learning. (anglicky))
112. Nowak, Ronald. (1999). Walker's Mammals of the World. Johns Hopkins University Press ISBN 978-0-8018-5789-8. (kontsulta data: 2023-05-29).
113. Russel, E. M. (1996) sid. 98
114. Berkovitz a Shellis 2018, s. 57–58. (BERKOVITZ, Barry; SHELLIS, Peter, 2018. The Teeth of Mammalian Vertebrates. 1. vyd. London, United Kingdom ; San Diego, CA, United States: Academic Press, an imprint of Elsevier. ISBN 978-0-12-802818-6. (anglicky))
115. Russel, E. M. (1996) sid. 99
116. Berkovitz a Shellis 2018, s. 59. (BERKOVITZ, Barry; SHELLIS, Peter, 2018. The Teeth of Mammalian Vertebrates. 1. vyd. London, United Kingdom ; San Diego, CA, United States: Academic Press, an imprint of Elsevier. ISBN 978-0-12-802818-6. (anglicky))
117. Berkovitz a Shellis 2018, s. 61. (BERKOVITZ, Barry; SHELLIS, Peter, 2018. The Teeth of Mammalian Vertebrates. 1. vyd. London, United Kingdom ; San Diego, CA, United States: Academic Press, an imprint of Elsevier. ISBN 978-0-12-802818-6. (anglicky))
118. Berkovitz a Shellis 2018, s. 63–64. (BERKOVITZ, Barry; SHELLIS, Peter, 2018. The Teeth of Mammalian Vertebrates. 1. vyd. London, United Kingdom ; San Diego, CA, United States: Academic Press, an imprint of Elsevier. ISBN 978-0-12-802818-6. (anglicky))
119. Berkovitz a Shellis 2018, s. 66–73. (BERKOVITZ, Barry; SHELLIS, Peter, 2018. The Teeth of Mammalian Vertebrates. 1. vyd. London, United Kingdom ; San Diego, CA, United States: Academic Press, an imprint of Elsevier. ISBN 978-0-12-802818-6. (anglicky))
120. (Ingelesez) Themes, U. F. O.. (2016-08-27). «Marsupialia (Marsupials)» Veterian Key (kontsulta data: 2023-05-29).
121. Tyndale-Biscoe, H. (2005) sid.18
122. Nowak 2005, s. 10–11. (NOWAK, Ronald, 2005. Walker's Marsupials of the World. Baltimore: Johns Hopkins University Press. Dostupné online. ISBN 9780801882227, ISBN 9780801882111. (anglicky))
123. NOVACEK, Michael J.; ROUGIER, Guillermo W.; WIBLE, John R. Epipubic bones in eutherian mammals from the Late Cretaceous of Mongolia. Nature. 1997-10, roč. 389, čís. 6650, s. 483–486. Dostupné online [cit. 2023-07-25]. ISSN 0028-0836. doi:10.1038/39020. (anglicky)
124. The Thylacine Museum - Biology: Anatomy: Skull and Skeleton: Post-cranial Skeleton (page 1). www.naturalworlds.org [online]. [cit. 2023-07-13]. Dostupné online.
125. Samuels ME, Regnault S, Hutchinson JR (2017). "Evolution of the patellar sesamoid bone in mammals". PeerJ. 5 e3103. Bibcode:2017PeerJ...5.3103S. doi:10.7717/peerj.3103. PMC 5363259. PMID 28344905.
126. Dickman, C. «Marsupials of the World: An Introduction». A: Nowak, R. M. Walker's Marsupials of the World (en anglès). The Johns Hopkins University Press, 2005, p. 1-67. ISBN 9780801882227.
127. Tyndale-Biscoe, H. (2005) sid.127
128. Hussey, Matt (2011). "Marsúipiach". Fréamh an Eolais. Coiscéim. p. 440.
129. Nowak 2018, s. 311. (NOWAK, Ronald, 2018. Walker's Mammals of the World : Monotremes, Marsupials, Afrotherians, Xenarthrans, and Sundatherians. Baltimore; London: Johns Hopkins University Press. Dostupné online. (anglicky))
130. Dickman, A Fragile Balance, 19. (Dickman, Christopher (2008). A Fragile Balance: The Extraordinary Story of Australian Marsupials (in English). University Of Chicago Press. ISBN 978-0226146300.)
131. Zima a Macholán 2021, s. 129. (ZIMA, J.; MACHOLÁN, M, 2021. Systém a fylogeneze savců. 1. vyd. Praha: Academia. 570 s. ISBN 978-80-200-3215-7, ISBN 80-200-3215-0.)
132. McDade 2003b, s. 83. (MCDADE, M. C. & kol, 2003b. Grzimek’s Animal Life Encyclopedia – Volume 13 Mammals II. 2. vyd. Farmington Hills, MI: Gale. (anglicky))
133. Nowak 2018, s. 192. (NOWAK, Ronald, 2018. Walker's Mammals of the World : Monotremes, Marsupials, Afrotherians, Xenarthrans, and Sundatherians. Baltimore; London: Johns Hopkins University Press. Dostupné online. (anglicky))
134. CANOPY LIFE: flying, gliding, prehensile tails and other adaptations to life at the top [online]. szgdocent.org. Dostupné v archivu pořízeném z originálu dne 2006-05-06.
135. Zima a Macholán 2021, s. 105. (ZIMA, J.; MACHOLÁN, M, 2021. Systém a fylogeneze savců. 1. vyd. Praha: Academia. 570 s. ISBN 978-80-200-3215-7, ISBN 80-200-3215-0.)
136. Nowak, Marsupials of the World, 10. (Nowak, Ronald M. (2005). Walker's Marsupials of the World (in English). Baltimore: Johns Hopkins University Press. ISBN 0-8018-8222-2.)
137. Armati, Dickman a Hume 2006, s. 175. (ARMATI, P. J.; DICKMAN, C. R.; HUME, I. D, 2006. Marsupials. New York: Cambridge University Press,. Dostupné online. (anglicky))
138. Armati, Dickman and Hume, Marsupials, 175. (Armati, Patricia; Dickman, Chris; Hume, Ian, eds. (2006). Marsupials (in English). Cambridge University Press. ISBN 978-0-521-65074-8.)
139. Armati, Dickman a Hume 2006, s. 185. (ARMATI, P. J.; DICKMAN, C. R.; HUME, I. D, 2006. Marsupials. New York: Cambridge University Press,. Dostupné online. (anglicky))
140. Nowak 2018, s. 283. (NOWAK, Ronald, 2018. Walker's Mammals of the World : Monotremes, Marsupials, Afrotherians, Xenarthrans, and Sundatherians. Baltimore; London: Johns Hopkins University Press. Dostupné online. (anglicky))
141. ZEISS, Caroline J.; SCHWAB, Ivan R.; MURPHY, Christopher J. Comparative retinal morphology of the platypus. Journal of Morphology. 2011-08, roč. 272, čís. 8, s. 949–957. PMID: 21567446. Dostupné online [cit. 2023-07-26]. ISSN 1097-4687. doi:10.1002/jmor.10959. PMID 21567446.
142. ARRESE, Catherine A; ODDY, Alison Y; RUNHAM, Philip B. Cone topography and spectral sensitivity in two potentially trichromatic marsupials, the quokka ( Setonix brachyurus ) and quenda ( Isoodon obesulus ). Proceedings of the Royal Society B: Biological Sciences. 2005-04-22, roč. 272, čís. 1565, s. 791–796. Dostupné online [cit. 2023-07-26]. ISSN 0962-8452. doi:10.1098/rspb.2004.3009. PMID 15888411. (anglicky)
143. EBELING, Wiebke; NATOLI, Riccardo C.; HEMMI, Jan M. Diversity of color vision: not all Australian marsupials are trichromatic. PloS One. 2010-12-06, roč. 5, čís. 12, s. e14231. PMID: 21151905 PMCID: PMC2997786. Dostupné online [cit. 2023-07-26]. ISSN 1932-6203. doi:10.1371/journal.pone.0014231. PMID 21151905.
144. Armati, Dickman a Hume 2006, s. 183. (ARMATI, P. J.; DICKMAN, C. R.; HUME, I. D, 2006. Marsupials. New York: Cambridge University Press,. Dostupné online. (anglicky))
145. Armati, Dickman a Hume 2006, s. 184. (ARMATI, P. J.; DICKMAN, C. R.; HUME, I. D, 2006. Marsupials. New York: Cambridge University Press,. Dostupné online. (anglicky))
146. SÁNCHEZ-VILLAGRA, Marcelo R. Ontogenetic and phylogenetic transformations of the vomeronasal complex and nasal floor elements in marsupial mammals. Zoological Journal of the Linnean Society. 2001-04, roč. 131, čís. 4, s. 459–479. Dostupné online [cit. 2023-07-26]. doi:10.1111/j.1096-3642.2001.tb01322.x. (anglicky)
147. Graves JA, Renfree MB (2013)Marsupials in the age of genomics. Annu Rev Genom Hum Genet
148. DEAKIN, Janine. Chromosome Evolution in Marsupials. Genes. 2018-02-06, roč. 9, čís. 2, s. 72. Dostupné online [cit. 2023-08-11]. ISSN 2073-4425. doi:10.3390/genes9020072. PMID 29415454. (anglicky)
149. TODER, Roland; O’NEILL, Rachel J. W.; WIENBERG, Johannes. Comparative chromosome painting between two marsupials: origins of an XX/XY1Y2 sex chromosome system. Mammalian Genome. 1997-06, roč. 8, čís. 6, s. 418–422. Dostupné online [cit. 2023-08-11]. ISSN 0938-8990. doi:10.1007/s003359900459. (anglicky)
150. Zima a Macholán 2021, s. 84–85. (ZIMA, J.; MACHOLÁN, M, 2021. Systém a fylogeneze savců. 1. vyd. Praha: Academia. 570 s. ISBN 978-80-200-3215-7, ISBN 80-200-3215-0.)
151. JOHNSTON, P.G.; WATSON, C.M.; ADAMS, M. Sex chromosome elimination, X chromosome inactivation and reactivation in the southern brown bandicoot Isoodon obesulus (Marsupialia: Peramelidae). Cytogenetic and Genome Research. 2002, roč. 99, čís. 1–4, s. 119–124. Dostupné online [cit. 2023-08-11]. ISSN 1424-8581. doi:10.1159/000071583. (anglicky)
152. GALLUS, Susanne; HALLSTRÖM, Björn M.; KUMAR, Vikas. Evolutionary histories of transposable elements in the genome of the largest living marsupial carnivore, the Tasmanian devil. Molecular Biology and Evolution. 2015-05, roč. 32, čís. 5, s. 1268–1283. PMID: 25633377 PMCID: PMC4408412. Dostupné online [cit. 2023-08-11]. ISSN 1537-1719. doi:10.1093/molbev/msv017. PMID 25633377.
153. MIKKELSEN, Tarjei S.; WAKEFIELD, Matthew J.; AKEN, Bronwen. Genome of the marsupial Monodelphis domestica reveals innovation in non-coding sequences. Nature. 2007-05, roč. 447, čís. 7141, s. 167–177. Dostupné online [cit. 2023-08-11]. ISSN 1476-4687. doi:10.1038/nature05805. (anglicky)
154. Renfree M, Tyndale-Biscoe H (1987). Reproductive Physiology of Marsupials. Cambridge University Press. ISBN 978-0-521-33792-2.
155. Armati, Dickman a Hume 2006, s. 91. (ARMATI, P. J.; DICKMAN, C. R.; HUME, I. D, 2006. Marsupials. New York: Cambridge University Press,. Dostupné online. (anglicky))
156. Reis, Nelio R.; Peracchi, Adriano L.; Fandiño-Mariño, Hernán; Rocha, Vlamir J. (2005). Mamíferos da Fazenda Monte Alegre - Paraná. Londrina: Editora da Universidade Estadual de Londrina (Eduel). p. 15
157. Nowak, Ronald M. (12 September 2005). Walker's Marsupials of the World. JHU Press. ISBN 978-0-8018-8211-1.
158. Armati, Dickman a Hume 2006, s. 92. (ARMATI, P. J.; DICKMAN, C. R.; HUME, I. D, 2006. Marsupials. New York: Cambridge University Press,. Dostupné online. (anglicky))
159. King A (2001). "Discoveries about Marsupial Reproduction". Iowa State University Biology Dept. Archived from the original on 5 September 2012. Retrieved 22 November 2012.
160. Romer, Alfred Sherwood; Parsons, Thomas S. (1977). The Vertebrate Body. Philadelphia, PA: Holt-Saunders International. pp. 396–399. ISBN 978-0-03-910284-5.
161. Armati, Dickman a Hume 2006, s. 83–85. (ARMATI, P. J.; DICKMAN, C. R.; HUME, I. D, 2006. Marsupials. New York: Cambridge University Press,. Dostupné online. (anglicky))
162. Dickman, A Fragile Balance, 13. (Dickman, Christopher (2008). A Fragile Balance: The Extraordinary Story of Australian Marsupials (in English). University Of Chicago Press. ISBN 978-0226146300.)
163. Sadleir R (1973). The Reproduction of Vertebrates. Elsevier Science. ISBN 978-0-323-15935-7.
164. Staker L (30 June 2014). Macropod Husbandry, Healthcare and Medicinals—Volumes One and Two. Lynda Staker. ISBN 978-0-9775751-2-1.
165. On the Habits and Affinities of the New Australian Mammal, Notoryctes typhlops E. D. Cope The American Naturalist Vol. 26, No. 302 (February 1892), pp. 121–128
166. Feldhamer 2015, s. 248–249. (FELDHAMER, George A. & kol, 2015. Mammalogy – Adaptation, Diversity, Ecology. 4. vyd. Baltimore: Johns Hopkins University Press. (anglicky))
167. Armati, Dickman and Hume, Marsupials, 86. (Armati, Patricia; Dickman, Chris; Hume, Ian, eds. (2006). Marsupials (in English). Cambridge University Press. ISBN 978-0-521-65074-8.)
168. Rodger JC, Hughes RL (1973). "Studies of the accessory glands of male marsupials". Australian Journal of Zoology. 21 (3): 303. doi:10.1071/ZO9730303. hdl:1959.4/70011.
169. Vogelnest L, Portas T (1 May 2019). Current Therapy in Medicine of Australian Mammals. Csiro Publishing. ISBN 978-1-4863-0753-1.
170. Rodger JC (January 1976). "Comparative aspects of the accessory sex glands and seminal biochemistry of mammals". Comparative Biochemistry and Physiology. B, Comparative Biochemistry. 55 (1): 1–8. doi:10.1016/0305-0491(76)90164-4. PMID 780045.
171. Hunsaker II D (1977). The Biology of Marsupials. Elsevier Science. ISBN 978-0-323-14620-3.
172. Zima a Macholán 2021, s. 108. (ZIMA, J.; MACHOLÁN, M, 2021. Systém a fylogeneze savců. 1. vyd. Praha: Academia. 570 s. ISBN 978-80-200-3215-7, ISBN 80-200-3215-0.)
173. Drews, Barbara; Roellig, Kathleen; Menzies, Brandon R.; Shaw, Geoff; Buentjen, Ina; Herbert, Catherine A.; Hildebrandt, Thomas B.; Renfree, Marilyn B. (15 March 2013). "Ultrasonography of wallaby prenatal development shows that the climb to the pouch begins in utero". Scientific Reports. 3 (1458): 1458. Bibcode:2013NatSR...3.1458D. doi:10.1038/srep01458. PMC 3597997. PMID 23492830.
174. Smith, Kathleen K.; Keyte, Anna L. (2020). "Adaptations of the Marsupial Newborn: Birth as an Extreme Environment". The Anatomical Record. 303 (2): 235–249. doi:10.1002/ar.24049. PMID 30548826. S2CID 56484546.
175. Nowak 2018, s. 126. (NOWAK, Ronald, 2018. Walker's Mammals of the World : Monotremes, Marsupials, Afrotherians, Xenarthrans, and Sundatherians. Baltimore; London: Johns Hopkins University Press. Dostupné online. (anglicky))
176. Nowak 2005, s. 11–14. (NOWAK, Ronald, 2005. Walker's Marsupials of the World. Baltimore: Johns Hopkins University Press. Dostupné online. ISBN 9780801882227, ISBN 9780801882111. (anglicky))
177. Frankenberg SR, de Barros FR, Rossant J, Renfree MB (2016). "The mammalian blastocyst". Wiley Interdisciplinary Reviews. Developmental Biology. 5 (2): 210–232. doi:10.1002/wdev.220. PMID 26799266. S2CID 22001725.
178. KARDONG, Kenneth V. Vertebrates: Comparative Anatomy, Function, Evolution. 8. vyd. New York, NY: McGraw-Hill Education, 2006. 790 s. Dostupné online. ISBN 978-1-260-09204-2, ISBN 978-1-259-70091-0. Kapitola Life History.
179. Short RV, Balaban E (1994). The Differences Between the Sexes. Cambridge University Press. ISBN 978-0-521-44878-9.
180. Side om reproduksjon Arkivert 28. august 2008 hos Wayback Machine. fra Centennial Museum, Univerity of Texas and El Paso
181. Blackburn, D. G. (2000). Classification of the reproductive patterns of amniotes:" Herpetological Monographs", side 371-377.
182. Dickman, A Fragile Balance, 14. (Dickman, Christopher (2008). A Fragile Balance: The Extraordinary Story of Australian Marsupials (in English). University Of Chicago Press. ISBN 978-0226146300.)
183. Nowak, R. M. (1999) s. 144 online
184. Old JM, Deane EM (2003). "The lymphoid and immunohaematopoietic tissues of the embryonic brushtail possum (Trichosurus vulpecula)". Anatomy and Embryology. 206 (3): 193–197. doi:10.1007/s00429-002-0285-2. PMID 12592570. S2CID 546795.
185. Sears KE (August 2009). "Differences in the timing of prechondrogenic limb development in mammals: the marsupial-placental dichotomy resolved". Evolution; International Journal of Organic Evolution. 63 (8): 2193–2200. Bibcode:2009Evolu..63.2193S. doi:10.1111/j.1558-5646.2009.00690.x. PMID 19453378. S2CID 42635687.
186. Smith KK (2001). "Early development of the neural plate, neural crest and facial region of marsupials". Journal of Anatomy. 199 (Pt 1–2): 121–131. doi:10.1046/j.1469-7580.2001.19910121.x. PMC 1594995. PMID 11523813.
187. Cáceras a Dickman 2023, s. 405–407. (CÁCERAS, Nilton C.; DICKMAN, C. R, 2023. American and Australasian Marsupials – An Evolutionary, Biogeographical, and Ecological Approach. Cham: Springer. ISBN 978-3-031-08418-8, ISBN 978-3-031-08419-5. (anglicky))
188. «Early development of the neural plate, neural crest and facial region of marsupials» (PDF)
189. Vaughan, Ryan a Czaplawski 2015, s. 81–82. (VAUGHAN, Terry A.; RYAN, James M.; CZAPLAWSKI, Nicholas J, 2015. Mammalogy. 6. vyd. Burlington, MA: Jones & Bartlett Learning. (anglicky))
190. Dawkins, Richard; Wong, Yan (9 February 2005). The Ancestor's Tale: A Pilgrimage to the Dawn of Evolution. Houghton Mifflin Harcourt. ISBN 978-0-618-61916-0 – via Google Books.
191. Kangaroos and Other Marsupials: Climb to the Pouch Begins in Utero (in English), ScienceDaily, 15 March 2013, retrieved 12 August 2013
192. Tyndale-Biscoe, H. (2005) sid.70
193. Schneider NY (August 2011). "The development of the olfactory organs in newly hatched monotremes and neonate marsupials". Journal of Anatomy. 219 (2): 229–242. doi:10.1111/j.1469-7580.2011.01393.x. PMC 3162242. PMID 21592102.
194. Ferner, K.; Schultz, J. A.; Zeller, U. (2017). "Comparative anatomy of neonates of the three major mammalian groups (monotremes, marsupials, placentals) and implications for the ancestral mammalian neonate morphotype – PMC". Journal of Anatomy. 231 (6): 798–822. doi:10.1111/joa.12689. PMC 5696127. PMID 28960296.
195. Dickman, A Fragile Balance, 16–17. (Dickman, Christopher (2008). A Fragile Balance: The Extraordinary Story of Australian Marsupials (in English). University Of Chicago Press. ISBN 978-0226146300.)
196. Dickman, A Fragile Balance, 17. (Dickman, Christopher (2008). A Fragile Balance: The Extraordinary Story of Australian Marsupials (in English). University Of Chicago Press. ISBN 978-0226146300.)
197. Gaisler a Zima 2007, s. 570–571. (GAISLER, Jiří; ZIMA, Jan, 2007. Zoologie obratlovců. 2. vyd. Praha: Academia. ISBN 978-80-200-1484-9.)
198. Stannard HJ, Miller RD, Old JM (2020). "Marsupial and monotreme milk – a review of its nutrients and immune properties". PeerJ. 8 e9335. Bibcode:2020PeerJ...8e9335S. doi:10.7717/peerj.9335. PMC 7319036. PMID 32612884.
199. LEFÈVRE, Christophe M.; DIGBY, Matthew R.; WHITLEY, Jane C. Lactation transcriptomics in the Australian marsupial, Macropus eugenii: transcript sequencing and quantification. BMC Genomics. 2007-11-13, roč. 8, čís. 1, s. 417. Dostupné online [cit. 2023-08-01]. ISSN 1471-2164. doi:10.1186/1471-2164-8-417. PMID 17997866.
200. Armati, Dickman a Hume 2006, s. 97. (ARMATI, P. J.; DICKMAN, C. R.; HUME, I. D, 2006. Marsupials. New York: Cambridge University Press,. Dostupné online. (anglicky))
201. Dickman, A Fragile Balance, 16. (Dickman, Christopher (2008). A Fragile Balance: The Extraordinary Story of Australian Marsupials (in English). University Of Chicago Press. ISBN 978-0226146300.)
202. Anděra 1997, s. 24. (ANDĚRA, Miloš, 1997. Savci (1). Praha: Albatros. (Svět zvířat; sv. 1). ISBN 80-00-00541-7.)
203. Nowak 2005, s. 17–21. (NOWAK, Ronald, 2005. Walker's Marsupials of the World. Baltimore: Johns Hopkins University Press. Dostupné online. ISBN 9780801882227, ISBN 9780801882111. (anglicky))
204. POTTER, Sally; CLOSE, Robert L.; TAGGART, David A. Taxonomy of rock-wallabies, Petrogale (Marsupialia: Macropodidae). IV. Multifaceted study of the brachyotis group identifies additional taxa. Australian Journal of Zoology. 2014, roč. 62, čís. 5, s. 401. Dostupné online [cit. 2023-08-04]. ISSN 0004-959X. doi:10.1071/ZO13095. (anglicky)
205. Armati, Dickman a Hume 2006, s. 205–206. (ARMATI, P. J.; DICKMAN, C. R.; HUME, I. D, 2006. Marsupials. New York: Cambridge University Press,. Dostupné online. (anglicky))
206. Trichosurus vulpecula. www.iucngisd.org [online]. [cit. 2023-08-05]. Dostupné online.
207. Wilson & Reeder (2005): Macropus rufogriseus departments.bucknell.edu.
208. Wilson & Reeder (2005): Macropus parma departments.bucknell.edu.
209. McDade 2003a, s. 253. (MCDADE, M. C. & kol, 2003a. Grzimek’s Animal Life Encyclopedia – Volume 12 Mammals I. 2. vyd. Farmington Hills, MI: Gale. (anglicky))
210. Dickman, A Fragile Balance, 50–52. (Dickman, Christopher (2008). A Fragile Balance: The Extraordinary Story of Australian Marsupials (in English). University Of Chicago Press. ISBN 978-0226146300.)
211. Nowak 2018, s. 93. (NOWAK, Ronald, 2018. Walker's Mammals of the World : Monotremes, Marsupials, Afrotherians, Xenarthrans, and Sundatherians. Baltimore; London: Johns Hopkins University Press. Dostupné online. (anglicky))
212. McDade 2003a, s. 289. (MCDADE, M. C. & kol, 2003a. Grzimek’s Animal Life Encyclopedia – Volume 12 Mammals I. 2. vyd. Farmington Hills, MI: Gale. (anglicky))
213. Dickman, A Fragile Balance, 52. (Dickman, Christopher (2008). A Fragile Balance: The Extraordinary Story of Australian Marsupials (in English). University Of Chicago Press. ISBN 978-0226146300.)
214. Dickman, A Fragile Balance, 52–53. (Dickman, Christopher (2008). A Fragile Balance: The Extraordinary Story of Australian Marsupials (in English). University Of Chicago Press. ISBN 978-0226146300.)
215. Anděra 1997, s. 25. (ANDĚRA, Miloš, 1997. Savci (1). Praha: Albatros. (Svět zvířat; sv. 1). ISBN 80-00-00541-7.)
216. Nowak 2005, s. 25–27. (NOWAK, Ronald, 2005. Walker's Marsupials of the World. Baltimore: Johns Hopkins University Press. Dostupné online. ISBN 9780801882227, ISBN 9780801882111. (anglicky))
217. McDade 2003b, s. 38. (MCDADE, M. C. & kol, 2003b. Grzimek’s Animal Life Encyclopedia – Volume 13 Mammals II. 2. vyd. Farmington Hills, MI: Gale. (anglicky))
218. Dickman, A Fragile Balance, 63. (Dickman, Christopher (2008). A Fragile Balance: The Extraordinary Story of Australian Marsupials (in English). University Of Chicago Press. ISBN 978-0226146300.)
219. Armati, Dickman a Hume 2006, s. 149–151. (ARMATI, P. J.; DICKMAN, C. R.; HUME, I. D, 2006. Marsupials. New York: Cambridge University Press,. Dostupné online. (anglicky))
220. Vaughan, Ryan a Czaplawski 2015, s. 98–99. (VAUGHAN, Terry A.; RYAN, James M.; CZAPLAWSKI, Nicholas J, 2015. Mammalogy. 6. vyd. Burlington, MA: Jones & Bartlett Learning. (anglicky))
221. Cáceras a Dickman 2023, s. 948–949. (CÁCERAS, Nilton C.; DICKMAN, C. R, 2023. American and Australasian Marsupials – An Evolutionary, Biogeographical, and Ecological Approach. Cham: Springer. ISBN 978-3-031-08418-8, ISBN 978-3-031-08419-5. (anglicky))
222. RUSSELL, Eleanor M. Social behaviour and social organization of marsupials. Mammal Review. 1984-09, roč. 14, čís. 3, s. 101–154. Dostupné online [cit. 2023-08-08]. ISSN 0305-1838. doi:10.1111/j.1365-2907.1984.tb00343.x. (anglicky)
223. Armati, Dickman and Hume, Marsupials, 277. (Armati, Patricia; Dickman, Chris; Hume, Ian, eds. (2006). Marsupials (in English). Cambridge University Press. ISBN 978-0-521-65074-8.)
224. Nowak 2018, s. 264. (NOWAK, Ronald, 2018. Walker's Mammals of the World : Monotremes, Marsupials, Afrotherians, Xenarthrans, and Sundatherians. Baltimore; London: Johns Hopkins University Press. Dostupné online. (anglicky))
225. Russel, E. M. (1996) sid. 100
226. Armati, Dickman a Hume 2006, s. 236. (ARMATI, P. J.; DICKMAN, C. R.; HUME, I. D, 2006. Marsupials. New York: Cambridge University Press,. Dostupné online. (anglicky))
227. Armati, Dickman and Hume, Marsupials, 236–240. (Armati, Patricia; Dickman, Chris; Hume, Ian, eds. (2006). Marsupials (in English). Cambridge University Press. ISBN 978-0-521-65074-8.)
228. Nowak 2005, s. 27–28. (NOWAK, Ronald, 2005. Walker's Marsupials of the World. Baltimore: Johns Hopkins University Press. Dostupné online. ISBN 9780801882227, ISBN 9780801882111. (anglicky))
229. Cáceras a Dickman 2023, s. 967–968. (CÁCERAS, Nilton C.; DICKMAN, C. R, 2023. American and Australasian Marsupials – An Evolutionary, Biogeographical, and Ecological Approach. Cham: Springer. ISBN 978-3-031-08418-8, ISBN 978-3-031-08419-5. (anglicky))
230. Bradley, “Stress, hormones and mortality in small carnivorous marsupials”. (Bradley, A.J. (2003). "Stress, hormones and mortality in small carnivorous marsupials". In Jones, M.; Dickman, C.; Archer, M. (eds.). Predators with Pouches: The Biology of Carnivorous Marsupials (in English). Sydney: CSIRO Press. p. 250–263.)
231. Nowak 2005, s. 31–33. (NOWAK, Ronald, 2005. Walker's Marsupials of the World. Baltimore: Johns Hopkins University Press. Dostupné online. ISBN 9780801882227, ISBN 9780801882111. (anglicky))
232. Cáceras a Dickman 2023, s. 946–947. (CÁCERAS, Nilton C.; DICKMAN, C. R, 2023. American and Australasian Marsupials – An Evolutionary, Biogeographical, and Ecological Approach. Cham: Springer. ISBN 978-3-031-08418-8, ISBN 978-3-031-08419-5. (anglicky))
233. GALETKA, Benjamin. Vombatus ursinus (coarse-haired wombat). Animal Diversity Web [online]. [cit. 2023-08-10]. Dostupné online. (anglicky)
234. Stonehouse and Gilmore, The Biology of Marsupials, 146. (Stonehouse, Bernard; Gilmore, Desmond (1977). The Biology of Marsupials (in English). University Park Press. ISBN 0-8391-0852-4.)
235. Jackson, Australian Mammals. (Jackson, S.M. (2003). Australian Mammals: Biology and Captive Management (in English). Melbourne: CSIRO Publishing. ISBN 9780643095076.)
236. Antiabong et al, “A molecular survey of a captive wallaby population”. (Antiabong, John F.; Boardman, Wayne; Smith, Ian; Brown, Melissa H.; Ball, Andrew S.; Goodman, Amanda E. (2013). "A molecular survey of a captive wallaby population for periodontopathogens and the co-incidence of Fusobacterium necrophorum subspecies necrophorum with periodontal diseases". Veterinary Microbiology (in English). 163 (3–4): 335–343.)
237. Smith, “Macropod Nutrition”. (Smith, Joseph A. (2009). "Macropod Nutrition". Veterinary Clinics of North America: Exotic Animal Practice (in English). 12 (2): 197–208. doi:10.1016/j.cvex.2009.01.010.)
238. Polkinghorne, Hanger and Timms, “Recent advances in understanding the chlamydial infections in koalas”. (Polkinghorne, Adam; Hanger, Jon; Timms, Peter (2013). "Recent advances in understanding the biology, epidemiology and control of chlamydial infections in koalas". Veterinary Microbiology (in English). 165 (3–4): 214–223. doi:10.1016/j.vetmic.2013.02.026.)
239. MURCHISON, Elizabeth P.; SCHULZ-TRIEGLAFF, Ole B.; NING, Zemin. Genome Sequencing and Analysis of the Tasmanian Devil and Its Transmissible Cancer. Cell. 2012-02, roč. 148, čís. 4, s. 780–791. Dostupné online [cit. 2023-08-11]. doi:10.1016/j.cell.2011.11.065. PMID 22341448. (anglicky)
240. Hawkins, C.E., McCallum, H., Mooney, N., Jones, M. & Holdsworth, M. 2008. Sarcophilus harrisii. The IUCN Red List of Threatened Species 2008: e.T40540A10331066. https://dx.doi.org/10.2305/IUCN.UK.2008.RLTS.T40540A10331066.en. Accessed on 13 August 2023.
241. Deakin, “Marsupial Genome Sequences”. (Deakin, Janine E. (2012). "Marsupial Genome Sequences: Providing Insight into Evolution and Disease". Scientifica (in English): 1–22.)
242. Dickman, A Fragile Balance, 49. (Dickman, Christopher (2008). A Fragile Balance: The Extraordinary Story of Australian Marsupials (in English). University Of Chicago Press. ISBN 978-0226146300.)
243. Borchard, Eldridge and Wright, “Sarcoptes mange increases diurnal activity of bare-nosed wombats”. (Borchard, Philip; Eldridge, David J.; Wright, Ian A. (2012). "Sarcoptes mange (Sarcoptes scabiei) increases diurnal activity of bare-nosed wombats (Vombatus ursinus) in an agricultural riparian environment". Mammalian Biology (in English). 77 (4): 244–248. doi:10.1016/j.mambio.2012.04.004.)
244. Ruykys et al, “Effects and Treatment of Sarcoptic Mange in Southern Hairy-Nosed Wombats”. (Ruykys, Laura; Breed, Bill; Schultz, David; Taggart, David (2013). "Effects and Treatment of Sarcoptic Mange in Southern Hairy-Nosed Wombats (Lasiorhinus latifrons)". Journal of Wildlife Sciences (in English). 49 (2): 312–320. doi:10.7589/2012-10-256.)
245. Cáceras a Dickman 2023, s. 9–11. (CÁCERAS, Nilton C.; DICKMAN, C. R, 2023. American and Australasian Marsupials – An Evolutionary, Biogeographical, and Ecological Approach. Cham: Springer. ISBN 978-3-031-08418-8, ISBN 978-3-031-08419-5. (anglicky))
246. Dickman, A Fragile Balance, 145. (Dickman, Christopher (2008). A Fragile Balance: The Extraordinary Story of Australian Marsupials (in English). University Of Chicago Press. ISBN 978-0226146300.)
247. Dickman, A Fragile Balance, 146–147. (Dickman, Christopher (2008). A Fragile Balance: The Extraordinary Story of Australian Marsupials (in English). University Of Chicago Press. ISBN 978-0226146300.)
248. Tyndale-Biscoe H (2004). Life of Marsupials. Collingwood, Australia: CSIRO. ISBN 978-0-643-06257-3.
249. Dickman, A Fragile Balance, 150. (Dickman, Christopher (2008). A Fragile Balance: The Extraordinary Story of Australian Marsupials (in English). University Of Chicago Press. ISBN 978-0226146300.)
250. Dawson TJ (2012). Kangaroos (2nd ed.). Collingwood, US: CSIRO Publishing. p. 181. ISBN 978-0-643-10625-3.
251. Armati, Dickman a Hume 2006, s. 1–2. (ARMATI, P. J.; DICKMAN, C. R.; HUME, I. D, 2006. Marsupials. New York: Cambridge University Press,. Dostupné online. (anglicky))
252. Marsupials. Cambridge University Press ISBN 978-0-511-24207-6..
253. Armati, Dickman a Hume 2006, s. 301–305. (ARMATI, P. J.; DICKMAN, C. R.; HUME, I. D, 2006. Marsupials. New York: Cambridge University Press,. Dostupné online. (anglicky))
254. Dickman, A Fragile Balance, 155. (Dickman, Christopher (2008). A Fragile Balance: The Extraordinary Story of Australian Marsupials (in English). University Of Chicago Press. ISBN 978-0226146300.)
255. Tyndale-Biscoe, H. (2005) sid.376-382
256. Dickman, A Fragile Balance, 156. (Dickman, Christopher (2008). A Fragile Balance: The Extraordinary Story of Australian Marsupials (in English). University Of Chicago Press. ISBN 978-0226146300.)
257. PIPER, Ross. Extinct Animals: An Encyclopedia of Species That Have Disappeared during Human History. Westport: Greenwood Press, 2009. Dostupné online. ISBN 978-0-313-34987-4. S. 11–13. (anglicky)
258. Kennedy, “Australian Marsupials and Monotremes”, 29. (Kennedy, Michael (1992). Australian Marsupials and Monotremes: An Action Plan for their Conservation (in English). IUCN/SSC Australian Marsupial and Monotreme Specialist Group. ISBN 2-8317-0052-3.)
259. Kennedy, “Australian Marsupials and Monotremes”, 29-30. (Kennedy, Michael (1992). Australian Marsupials and Monotremes: An Action Plan for their Conservation (in English). IUCN/SSC Australian Marsupial and Monotreme Specialist Group. ISBN 2-8317-0052-3.)
260. Kennedy, “Australian Marsupials and Monotremes”, 30. (Kennedy, Michael (1992). Australian Marsupials and Monotremes: An Action Plan for their Conservation (in English). IUCN/SSC Australian Marsupial and Monotreme Specialist Group. ISBN 2-8317-0052-3.)
261. Mammals [online]. IUCN SSC Groups [cit. 2023-08-13]. Dostupné online. (anglicky)
262. Flores, D. 2016. Cryptonanus ignitus. The IUCN Red List of Threatened Species 2016: e.T41320A22177809. https://dx.doi.org/10.2305/IUCN.UK.2016-2.RLTS.T41320A22177809.en. Accessed on 13 August 2023.
263. Cáceras a Dickman 2023, s. 1348–1351. (CÁCERAS, Nilton C.; DICKMAN, C. R, 2023. American and Australasian Marsupials – An Evolutionary, Biogeographical, and Ecological Approach. Cham: Springer. ISBN 978-3-031-08418-8, ISBN 978-3-031-08419-5. (anglicky))
264. Burbidge, A.A., Johnson, C.N. & Zichy-Woinarski, J. 2016. Onychogalea fraenata. The IUCN Red List of Threatened Species 2016: e.T15330A21958130. https://dx.doi.org/10.2305/IUCN.UK.2016-1.RLTS.T15330A21958130.en. Accessed on 08 October 2023.