Overview
Distribution
Range Description
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Geographic Range
Southern hairy-nosed wombats inhabit the semi-arid and arid grasslands and woodlands that receive 200 to 500 mm of rain per year in southeastern Western Australia, southern South Australia, and southwestern New South Wales.
Biogeographic Regions: australian (Native )
- Menkhorst, P. 2001. A Field Guide to the Mammals of Australia. South Melbourne: Oxford University Press.
- New South Wales, National Parks and Wildlife Service. Southern Hairy-Nosed Wombat. 1. Hurstville, NSW, Australia: NSW National Parks and Wildlife Services. 1999. Accessed March 20, 2006 at http://www.nationalparks.nsw.gov.au/PDFs/tsprofile_southern_hairynosed_wombat.pdf.
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Physical Description
Morphology
Physical Description
Southern hairy-nosed wombats are stocky, robust animals, with a lot of power for digging. The body is close to one meter long with short, powerful legs. They are plantigrade and have flattened claws on each of the 5 digits; these are used for digging. On the hind feet, the second and third toes have fused, creating a digit with two claws that the wombats use for grooming (Cronin, 2000). The robust head is flattened and has narrow, pointed ears that extend over its top (Menkhorst, 2001). The incisors are rodent-like. The palate between the molars is wide (Wells, 1987). All the teeth grow continuously throughout their lives, perhaps an adaptation to accommodate their harsh diet. Since hairy-nosed wombats eat a coarser diet than that of common wombats (Vombatus ursinus), they need to masticate their food more thoroughly than that species, so their temporalis muscle is better developed and the masseteric muscle is more reduced (Nakajima and Townsend, 1994). Lasiorhinus latifrons can be distinguished from other Lasiorhinus species because the nasal bone is longer than the frontal bone. The tail is short and usually hidden by fur. The silky fur ranges in color from grey to tan. Soft, usually white, fur covers the rhinarium, giving the animal its name.
Range mass: 19 to 32 kg.
Range length: 772 to 934 mm.
Other Physical Features: endothermic ; heterothermic ; bilateral symmetry
Sexual Dimorphism: male larger
Average basal metabolic rate: 16.001 W.
- Hamilton, R., P. Stanton, L. O'Donnell, V. Steele, D. Taggart, P. Temple-Smith. 2000. Determination of seasonality in southern hairy-nosed wombats (Lasiorhinus latifrons) by analysis of fecal androgens. Biology of Reproduction, 63: 526-531.
- Nakajima, K., G. Townsend. 1994. A morphometric study of the skulls of two species of wombats (vombatus ursinus and lasiorhinus latifrons). Australian Mammalogy, 17: 65-72.
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Wells, R. 1987.
- Vombatidae
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Ecology
Habitat
Habitat and Ecology
Systems
- Terrestrial
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Habitat
Lasiorhinus latifrons require sturdy soil that is able to support the digging of large burrows. They also require perennial grasses and bluebush scrub (Maireana and Chenopodium), essential parts of their diet.
Habitat Regions: temperate ; tropical ; terrestrial
Terrestrial Biomes: savanna or grassland ; forest
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Trophic Strategy
Food Habits
Southern hairy-nosed wombats are herbivorous, grazing primarily on grasses and herbs. When available, they prefer new green shoots of grass, especially species in the genus Stipa, and will eat them until they are no longer available (“Southern Hairy-Nosed Wombat”, 1997). During droughts, wombats have been known to eat leaves and stems of short bushes, particularly the bluebush species in the genus Maireana (Wells, 1987). There is some evidence that wombats feed on roots during dry periods, with some preference shown for Eucalyptus species (Wells, 1987).
When feeding, southern hairy-nosed wombats choose green shoots if they are present to maximize their water and nutrient uptake. The digestive system has two adaptations that aid wombats in the uptake of water and energy from their food sources. Wombats use fermentation by bacterial colonies in their intestines to help them digest cellulose (Ride, 1970). Additionally, their feces is very dry, containing only 50% water (Ride, 1970).
Plant Foods: leaves; roots and tubers; wood, bark, or stems
Primary Diet: herbivore (Folivore )
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Associations
Ecosystem Roles
When southern hairy-nosed wombats graze in front of their burrows, they may create an area with a higher density of new green shoots, a sign of delayed growth of individual grass plants (Wells, 1987). Additionally, other animals, such as introduced rabbits and foxes, may use wombat burrows. Southern hairy-nosed wombats are known to harbor a number of parasites, but seemingly with no effect on the wombats. Also, they may contract bacterial and fungal diseases.
Ecosystem Impact: creates habitat
Commensal/Parasitic Species:
- Progamotaenia diaphana
- Eimeria wombati
- Eimeria ursini
- Ileocystis wombati
- Macropostrongyloides lasiorhini
- Phascolostrongylus turleyi
- Oesophagostomoides longispicularis
- Oesophagostomoides gilteneri
- Oesophagostomoides stirtoni
- Ixodes victoriensis
- Ixodes tasmani
- Ixodes holocyclus
- Ixodes phascolomyis
- Aponomma auruginans
- Nowak, R. 1991. Walker's Mammals of the World. Baltimore, Maryland: Johns Hopkins University Press.
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Predation
Since southern hairy-nosed wombats live in burrows, they can easily escape from predators. No predators are known for Lasiorhinus latifrons, but the closely related Lasiorhinus krefftii are preyed upon by dingoes and feral dogs (Banks, et al., 2003). Predation does not appear to be the main limiting factor in wombat longevity. Wombats are extremely dependent on the amount of rainfall in their habitat, which affects their life-span more than predators.
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Life History and Behavior
Behavior
Communication and Perception
Southern hairy-nosed wombats communicate with each other through vocalizations and scents. A direct encounter between wombats is rare; they rely more heavily on scent to communicate. Wombats spend considerable time investigating scents left recently by other wombats (Wells, 1978). When two wombats do encounter each other, they make a rough coughing noise (Wells, 1978). When the animal is alarmed, it will use a more strident call to alert others (Wells, 1987). However, most communication between Lasiorhinus latifrons individuals occurs through olfaction and scent marking.
Communication Channels: acoustic ; chemical
Other Communication Modes: scent marks
Perception Channels: visual ; acoustic
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Life Expectancy
Lifespan/Longevity
Because southern hairy-nosed wombats have continuously growing teeth, it is difficult to non-invasively collect data about wombat age. The best means to make an estimate of an animal’s age is to tag and track it over the course of its life. In a study that tracked and tagged adult wombats, individuals lived for 14 years (Wells, 1987). The main limit on the lifespan of Lasiorhinus latifrons is the amount of rainfall in a season rather than predation (Wells, 1987). Wombats are able to escape predators because they live in burrows.
Average lifespan
Status: wild: 14 years.
Range lifespan
Status: captivity: 20 (high) years.
Average lifespan
Status: captivity: 17 years.
Average lifespan
Status: captivity: 18.0 years.
Average lifespan
Status: captivity: 24.5 years.
Average lifespan
Status: captivity: 20.0 years.
- Finlayson, G., G. Shimmin, P. Temple-Smith, K. Handasyde, D. Taggart. 2005. Burrow use and ranging behaviour of the southern hairy-nosed wombat (lasiorhinus latifrons) in the Murraylands, South Australia. Journal of Zoology, 265: 189-200.
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Lifespan, longevity, and ageing
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Reproduction
Reproduction
Little is known about the mating system of Lasiorhinus latifrons. During the breeding season, males are aggressive towards each other, biting the ears, rump, and flanks of other male wombats (Wells, 1995). Females ready for mating may emit a coughing call when they are pursued by males. However, much of the process of selecting a mate is unknown.
Mating System: polygynandrous (promiscuous)
Southern hairy-nosed wombats breed in synchrony with highest growth rates of the native grasses that make up their diet. Because of their dependence on the germination of grasses, their reproduction is affected by the amount of winter rainfall (Wells, 1987). During periods of drought, native grasses do not grow, and wombats do not breed. When the grasses begin to germinate, female wombats become ready for reproduction. Males also reach their peak reproductive activity when females are active (Temple-Smith et al., 2000). During the breeding season, males are aggressive towards each other and maintain dominance hierarchies (Cronin, 2000). Mating occurs in the burrow, with males usually remaining in one warren, while females move among them. When wombats copulate, the male lengthens the time of intromission by turning the female on her side and mounting her from behind (Wells, 1995). Female wombats usually give birth between September and December. A single young is born after 21 days of gestation. The juvenile then moves to the pouch where it completes its development. Young begin to leave the pouch between six and eight months of age, and leave permanently by nine months. Young wombats may continue to suckle at the pouch until they have reached 15 months of age (Cronin, 2000). Females are able to reproduce again after weaning is complete. At three years of age, both sexes reach sexual maturity.
Since the breeding of Lasiorhinus latifrons occurs in the summer, young wombats are weaned in the early spring, making them vulnerable to food stress and dependent on the amount of rain received in April (Wells, 1987). Wombats usually require two to three years with enough rainfall to successfully raise young. Without enough rainfall, there is not enough grass to sustain juveniles through their growth (Wells, 1995). Additionally, overgrazing of the land can destroy perennial grasses in favor of annual grasses that do not meet the nutritional needs of juvenile wombats.
Breeding interval: Southern hairy-nosed wombats breed once yearly, if optimal rainfall conditions prevail.
Breeding season: Breeding occurs from September to December.
Range number of offspring: 1 to 2.
Average number of offspring: 1.
Average gestation period: 21 days.
Range weaning age: 6 to 9 months.
Average time to independence: 1 years.
Average age at sexual or reproductive maturity (female): 3 years.
Average age at sexual or reproductive maturity (male): 3 years.
Key Reproductive Features: iteroparous ; seasonal breeding ; gonochoric/gonochoristic/dioecious (sexes separate); viviparous
Average birth mass: 0.5 g.
Average gestation period: 21 days.
Average number of offspring: 1.
Average age at sexual or reproductive maturity (male)
Sex: male: 547 days.
Average age at sexual or reproductive maturity (female)
Sex: female: 547 days.
Southern hairy-nosed wombat females invest heavily in their young. Since the young of all marsupials are born very early in development and depend on their mother’s milk to complete development, the mother spends a lot of energy producing milk over a long period of time. Additionally, once the young leave the pouch, they return sporadically for more milk while learning how to burrow and graze from their mothers.
Parental Investment: altricial ; pre-fertilization (Provisioning, Protecting: Female); pre-hatching/birth (Provisioning: Female, Protecting: Female); pre-weaning/fledging (Provisioning: Female, Protecting: Female); pre-independence (Provisioning: Female)
- Menkhorst, P. 2001. A Field Guide to the Mammals of Australia. South Melbourne: Oxford University Press.
- Hamilton, R., P. Stanton, L. O'Donnell, V. Steele, D. Taggart, P. Temple-Smith. 2000. Determination of seasonality in southern hairy-nosed wombats (Lasiorhinus latifrons) by analysis of fecal androgens. Biology of Reproduction, 63: 526-531.
- New South Wales, National Parks and Wildlife Service. Southern Hairy-Nosed Wombat. 1. Hurstville, NSW, Australia: NSW National Parks and Wildlife Services. 1999. Accessed March 20, 2006 at http://www.nationalparks.nsw.gov.au/PDFs/tsprofile_southern_hairynosed_wombat.pdf.
- Wells, R. 1995. Southern Hairy-nosed Wombat Lasiorhinus latifrons (Owen, 1845). Pp. 202-203 in R Strahan, ed. Mammals of Australia. Chatswood, N.S.W.: Reed Books.
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Wells, R. 1987.
- Vombatidae
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Conservation
Conservation Status
IUCN Red List Assessment
Red List Category
Red List Criteria
Version
Year Assessed
Assessor/s
Reviewer/s
Justification
History
- 1996Lower Risk/least concern
- 1996Lower Risk/least concern
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Conservation Status
Southern hairy-nosed wombats have withstood massive habitat loss due to human clearing of land. They are considered threatened, but are not listed as an endangered species (Cronin, 2000). Currently, one of the main threats to their survival is the spread of rabbits throughout Australia. Rabbits, and domestic livestock, compete with wombats for forage, leading to overgrazing in many areas. Once the land is overgrazed, dominant grass species shift from perennial ones, the native diet of wombats, to annual species that do not provide wombats with all of their metabolic needs (Wells, 1995). Additionally, human clearing of the land removes the vegetation that the wombats rely on during drought periods (“Southern Hairy-Nosed Wombat”, 1999). Hunting of wombats by aboriginal people is not considered a serious threat. Wombats are valued by aboriginal people, who will often leave their own land to hunt wombats elsewhere so that they can keep their own wombat populations healthy (Davies, 1998). Low wombat reproduction rates means that populations do not quickly recover from disruptions.
US Federal List: no special status
CITES: no special status
IUCN Red List of Threatened Species: least concern
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Trends
Population
Population Trend
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Threats
Threats
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Management
Conservation Actions
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Relevance to Humans and Ecosystems
Benefits
Economic Importance for Humans: Negative
Southern hairy-nosed wombats are sometimes agricultural pests. When they dig their burrows, they can destroy crops. The coexistence of wombats and livestock pose a resource competition problem (Nowak, 1991). Additionally, livestock can sometimes break their legs when they break through into a wombat burrow (Nowak, 1991). Burrows also provide problems for farmers because they are good habitats for other pest species such as rabbits (Nowak, 1991). Many farmers also incorrectly believe that wombats destroy fences on their properties (Nowak, 1991).
Negative Impacts: crop pest
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Economic Importance for Humans: Positive
Wombats are hunted throughout their region by aboriginal people. These native cultures consider wombats a part of their culture and enjoy eating their meat. However, they are cautious not to hunt them too frequently, as it takes a lot of time and energy to capture a wombat (Davies, 1998). Wombats have also been captured and domesticated as pets (Nowak, 1991).
Positive Impacts: food
- Davies, J. 1998. "Who Owns the Animals? Sustainable Commercial use of Wildlife and Indigenous Rights in Australia" (On-line pdf). Accessed April 17, 2006 at http://www.indiana.edu/~iascp/Final/davies.pdf.
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Wikipedia
Southern hairy-nosed wombat
The southern hairy-nosed wombat (Lasiorhinus latifrons) is one of three species of wombats. It is found in scattered areas of semi-arid scrub and mallee from the eastern Nullarbor Plain to the New South Wales border area. It is the smallest of all three wombat species. The young often do not survive dry seasons. It is the state animal of South Australia.[3]
The oldest southern hairy-nosed wombats ever documented were a male and a female from Brookfield Zoo just outside of Chicago. Their names were Carver, who lived to be 34, and his mother, Vicky, who lived to be 24.
Contents |
Physical description
The southern hairy-nosed wombat is adapted to digging; it has a stocky and robust build, flatten claws and 5 digits.[4] It is also plantigrade. The body length ranges 772–934 millimetres (30.4–36.8 in) with a body mass ranging 19–32 kilograms (42–71 lb).[4] It has a short tail that is hidden by its fur. The pelage is silky and is typically greyish or tan in colour. The wombat grooms itself with its second and third, which are fused together, expect at the tips.[5][6] The head is robust and flattened and the ears are pointed.[7] The snout resembles that of a pig.[5] The animal gets its name from the hairs that cover its rhinarium.[4] The wombat's incisors resemble those of rodents and have molars are widely spaced by the palate.[5] The teeth keep growing for the entirely of the animal’s life, which is likely an adaptation to its harsh diet.[4] Compared to the common wombat, the southern hairy-nosed wombat has a larger temporalis muscle and a smaller masseter muscle.[8] Also, unlike the northern hairy-nosed wombat, the southern hairy-nosed wombat’s nasal bone is longer than the frontal bone.[4]
Biology and ecology
Southern hairy-nosed wombats range though Western Australia, southern South Australia, and south-western New South Wales. They live in semi-arid to arid grasslands and woodlands.[7]
Feeding and energy
Southern hairy-nosed wombats, along with other wombat species, select native perennial grasses and sedges but do consume introduced pasture species, forbs and the leaves of woody shrubs if its favoured food isn't available.[9] Much of the southern hairy-nosed wombat's diet is Stipa nitida, which grows around its warren complex and are trimmed as it grazes.[9] This creates an area with a higher density of new green shoots, a sign of delayed growth of individual grass.[5] The teeth of the wombat are more effective in grinding food into small particles than the Western grey kangaroo.[5] The digestive tract of the wombat has a tiny caecum and a colon divided into parts.[9] The anterior part is relatively small and serves as the site for fermentation while the posterior part is larger and is where water is reabsorbed. The wombat conserves water by recycling more urea to the colon rather than releasing it as urine. Wombats release less than other herbivorous mammals.[10] As such the southern hairy-nosed wombat produces very dry faeces with water content as low as 40%.[9]
The harsh environment in which the southern hairy-nosed wombat lives is further reflected in its energetics. In captivity, their standard metabolic rate (SMR) was found to be 130 kl/kg^0.75 per day which is very low compared to most placental animals and other marsupials.[10][11] They also have the lowest thyroid hormone levels among mammals.[9] The food wombats eat provide more than enough energy.[10] As long as there is enough food, the forage consumed by the wombat can support it during late lactation. It is more effective than a donkey at maintaining its weight on low quailty food.[12]
Burrow system and activities
Southern hairy-nosed wombats dig and live in burrow which they connect into warrens with many entrances. These warrens are their prime refuges and are shared by up to ten individuals.[9] A wombat digs with its foreclaws while sitting up.[13] It leaves its new burrow backwards and push out dirt with all its paws.[13] The central warren is surrounded by a circle of small simple burrows 100–15 m from it.[11] There are small burrows along the other edge, which is where young wombat go when they are displaced from the central warren.[9] Wombats may favour a certain burrow and not share it with others. However, there is no monopolization of burrows.[14] Wombats move between burrows and even warrens. Male wombats are territorial towards wombats from other warrens, possibly to defend food resources and the warren refuges.[9] Trails of droppings connect the burrows together. They also mark their territory with anal scent secretions by rubbing their backs and rumps on objects.[5] Fights between males over territories or mates do occur and invlove bites the ears, flank, or rump.[4] There is also a dominance hierarchy among males.[4]
The burrows of a southern hairy-nosed wombat can have air temperatures around of 14°C in mid winter to 26°C in mid summer, the wombat's preferred thermo-neutral zone, while the ambivalent temperature are around 2 to 36 °C (36 to 97 °F).[15] Warrens can make surface conditions in habitats of low humidity and high temperatures better for the wombat.[15] A wombat retires deep in the burrow after foraging.[11] The next night, the wombat moves to the entrance to check if conditions are right before emerging again.[11] In the evening, wombats leave their burrows as the ambient temperature and burrow temperature are the same. In the early morning, when the surface temperature is lower, they retire.[16]
Mating and reproduction
The breeding of the southern hairy-nosed wombat occurs when their favoured food is at their peak growth rates. The reproduction of wombats relies on the winter rainfall, which germinates the grasses.[5] Between August and October, when there is enough rainfall, females enter ovulation and the males' testosterone levels and prostate gland size increase. In years of low rainfall, neither of these things occur.[9] When breeding, dominance hierarchies among males are established though aggression.[6] Copulation takes place in the warren with males remaining in one burrow and females moving among them. Mating takes place underground and involes the male mounting the female from behind while she is on her side.[14] The gestation period of the wombat lasts 22 days and most births occur in October. When a young is born, it climb into the pouch and clings to a teat. It stays in the pouch for six months. At this time weighs around 0.45 kg, with a light pelage and open eyes.[9] It soon lives the pouch and starts grazing at the surface. The young is fully weaned when it is a year old and reaches full size at three years old,[9] which is also when it becomes sexually mature.[4]
Communication
Southern hairy-nosed wombats use vocalisations and scents for communication. Wombats rely more on scent to communicate as they don't often encounter each other directly.[4] Wombats will examine they conspecifics leave.[11] While most communication between wombats occurs through olfaction and scent marking, wombats will emit rough coughing noises when they pass each other[11] and will emit a more strident call for alarm.[5]
Status
The southern hairy wombat is listed as Least Concern by the IUCN because "While there are sporadic outbreaks of sarcoptic mange, competition with introduced herbivores, susceptibility to drought, and severe fragmentation in parts of its range, the species has a wide distribution, large population, occurs in a number of protected areas, and it is unlikely to be declining at nearly the rate required to qualify for listing in a threatened category".[2] Wombats were hunted by aboriginal people for their meat. However, capturing a wombat takes a lot of time and energy and so they were not hunted too frequently.[17] The indigenous people of Australia value the wombat culturally and keep their local wombat populations healthy by hunting wombats in other areas.[17]
Wombats have been considered agricultural pests. Their digging can destory crops and can increase the risk livestock of livestock breaking their legs falling though their burrow system.[18] Competition between livestock, rabbits and wombat can lead to overgrazing. Overgrazing leads to annual grass species becoming the dominant species and wombats can't get enough of their metabolic needs from these.[4] The spread of rabbits threaten their survival of wombats.[4]
References
- ^ Groves, Colin P. (16 November 2005). "Order Diprotodontia (pp. 43-70)". In Wilson, Don E., and Reeder, DeeAnn M., eds. Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Baltimore: Johns Hopkins University Press, 2 vols. (2142 pp.). pp. 43–44. ISBN 978-0-8018-8221-0. OCLC 62265494. http://www.bucknell.edu/msw3/browse.asp?id=11000012.
- ^ a b Taggart, D. & Robinson, T. (2008). Lasiorhinus latifrons. In: IUCN 2008. IUCN Red List of Threatened Species. Downloaded on 28 December 2008. Database entry includes justification for why this species is of least concern
- ^ "Southern Hairy-nosed Wombat". Unique Australian Animals Website. http://australian-animals.net/wombat-s.htm.
- ^ a b c d e f g h i j k Green, Emily (2006). "Lasiorhinus latifrons southern hairy-nosed wombat". Animal Diversity Web. University of Michigan Museum of Zoology. http://animaldiversity.ummz.umich.edu/site/accounts/information/Lasiorhinus_latifrons.html. ] Animal Diversity
- ^ a b c d e f g h Wells, R. (1989). "Vombatidae". In Walton, D.W.. Fauna of Australia. Canberra, Australia: Australian Government Publishing Service. pp. 1–25. http://www.environment.gov.au/biodiversity/abrs/publications/fauna-of-australia/pubs/volume1b/32-ind.pdf.
- ^ a b Cronin, L. (2000). Australian Mammals. Annandale: Envirobook.
- ^ a b Menkhorst, P. (2001). A Field Guide to the Mammals of Australia. South Melbourne: Oxford University Press.
- ^ Nakajima, K.; Townsend, G. (1994). "A morphometric study of the skulls of two species of wombats (Vombatus ursinus and Lasiorhinus latifrons)". Australian Mammalogy 17: 65–72.
- ^ a b c d e f g h i j k Tyndale-Biscoe, H (2005). Life of Marsupials. CSIRO Publishing. pp. 269–285. ISBN 978-0-643-06257-3. http://books.google.com/books?id=KqtlPZJ9y8EC&pg=PA269.
- ^ a b c Wells, RT; Green, B (1998). "Aspects of water metabolism in the southern hairy-nosed wombat Lasiorhinus latifrons". In Wells, RT; Pridmore, PA. Wombats. Chipping Norton: Surrey Beatty. ISBN 0949324817. http://books.google.com/books?id=xPBFAAAAYAAJ.
- ^ a b c d e f Wells, R.T. (1978). "Thermoregulation and activity rhythms in the hairly-nosed wombats, Laisorhinus latifrons (Owen), (Vombatidae)". Australian Journal of Zoology 26 (4): 639–51. doi:10.1071/ZO9780639.
- ^ Hume ID (1999). Marsupial Nutrition. Cambridge: Cambridge University Press. p. 434. ISBN 052159555X.
- ^ a b Ride, W. (1970). A Guide to the Native Mammals of Australia. Melbourne: Oxford University Press. p. 93.
- ^ a b Wells, R. (1995). "Southern Hairy-nosed Wombat Lasiorhinus latifrons (Owen, 1845)". In Strahan, R.. Mammals of Australia. Chatswood, NSW: Reed Books. pp. 202–3.
- ^ a b Shimmin GA, Skinner J, Baudinette RV (2002). "The warren architecture and environment of the southern hairy-nosed wombat (Lasiorhinus latifrons)". Journal of Zoology 258 (4): 469–77. doi:10.1017/S0952836902001620.
- ^ Tayor GK. "A long term population study of the southern hairy-nosed wombat Laisorhinus latifrons at Moorude Wildlife Reserve, South Australia". pp. 198–205. (Wells & Pridmore 1998)
- ^ a b Davies J. 1998. "Who Owns the Animals? Sustainable Commercial use of Wildlife and Indigenous Rights in Australia" Presented at "Crossing Boundaries" the 7th annual conference of the International Association for the Study of Common Property, Vancouver, British Columbia, Canada, 10–14 June 1998.
- ^ Nowak, R. (1991). Walker's Mammals of the World. Baltimore MA: Johns Hopkins University Press.
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