Tammar Wallaby

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Tammar Wallaby
Conservation status
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Infraclass: Marsupialia
Order: Diprotodontia
Family: Macropodidae
Genus: Macropus
Species: M. eugenii
Binomial name
Macropus eugenii
Desmarest, 1817
Tammar Wallaby range

The tammar wallaby (Macropus eugenii), also known as the dama wallaby or darma wallaby, is a small macropod and the smallest species of wallaby. It is native to South and Western Australia as well as some nearby islands. Its range was much more extensive in the past but has been severely reduced since the arrival of Europeans. Nevertheless, the tammar remains common within its limited range. The tammar has several notable adaptations, including the ability to retain energy while hopping, colour vision and the ability to drink seawater. It is a nocturnal species and spends nighttime in grassland habitat, and daytime in shrub. It is also very gregarious and has promiscuous mating habits. A female tammar can nurse a joey in her pouch while keeping an embryo in its uterus. The tammar is a model species for research on macropods, and on marsupials in general. It is one of many organisms to have had its genome sequenced.

Contents

[edit] Taxonomy and classification

Drawing of tammar wallabies by John Gould (1863).

The tammar wallaby was seen on West Wallabi Island in the Houtman Abrolhos off Western Australia by survivors of the 1628 Batavia shipwreck, and recorded by Francisco Pelsart in his 1629 Ongeluckige Voyagie. This represents the first recorded sighting of a macropod by Europeans,[2] and probably also the first sighting of an Australian mammal.[3] It was first described by French naturalist Anselme Gaëtan Desmarest in 1817. Its specific name comes from Eugene Hamelin, Commander of the ship Naturaliste.[4]:333 Its common name is derived from the thickets of the local shrub Allocasuarina campestris that sheltered it in Western Australia.[5] It is classified together with the kangaroos and larger wallabies in the genus Macropus. Within this, it is placed in the subgenus Notomacropus along with several species of wallaby, all of which have a facial stripe.[6]

The tammar wallabies on Flinders Island have been described as having a greyer coat and thinner head than the tammars on Kangaroo Island[4]:333 and were given a separate name, while the East and West Wallabi Islands tammars are smaller than those on Kangaroo Island. Other early naturalists suspected that all island populations belonged to one species.[4]:333

A more recent study has compared tammar wallaby skulls, in museums around the world, with skulls from extant populations on the Abrolhos Islands, Garden Island, Middle Island and Kangaroo Island, as well as those from New Zealand, and found that there were three distinct groups; one group including populations from mainland Western Australia, East and West Wallabi Islands, Garden Island and Middle Island; a second group comprising populations from Flinders Island and mainland Southern Australia, in the 19th century, as well as populations from New Zealand; and a third group consisting of the Kangaroo Island population.[7]

[edit] Subspecies

Tammars at Budapest Zoo

As such, three subspecies of tammar wallaby are classified:[5]

  • M. e. derbianus (Western Australia and nearby islands)
  • M. e. decres (Kangaroo Island, South Australia)
  • M. e. eugenii (mainland South Australia and New Zealand).

[edit] Evolution

Fossil evidence exists from the late Pleistocene era—remains were found in the Naracoorte Caves.[6] The island dwelling tammar wallabies have been isolated from those on the mainland from between 7,000 and 15,000 years, since the end of the last Ice Age, and each population shows genetic differences from the other.[4]:332 It is likely that tammars originated in South Australia and spread to Western Australia; where they evolved a resistance to sodium fluoroacetate. The lethal dose for mainland Western tammars is greater than 5 mg/kg, while the lethal dose for tammars on East and West Wallabi Islands and Garden Island is less than 2 mg/kg.[8] Plant species that contain sodium fluoroacetate do not occur on the islands, and therefore tammars must have retained their tolerance since their islolation 7000-11,500 years ago.[4]:334 By contrast tammars from the eastern part of their range are highly susceptible to the poison, with the lethal dose for tammars on Kangaroo Island being less than 0.2 mg/kg.[8] There are also differences in serum proteins, iron-building transferrin protein and red cell antigens, between the Garden Island and Kangaroo Island populations, which are at two ends of the species’ continuum.[4]:333

[edit] Population dynamics and conservation

Tammar wallaby on North Island in the Houtman Abrolhos.

Since European occupation, tammar wallaby populations on both mainland Australia and some islands have been greatly reduced or even eradicated. The mainland population of tammars have been descripted as "very plentiful in many parts of the south-west, but rapidly disappearing in the cultivated districts, especially towards the northern end of its range."[9] The region was cleared for wheat and sheep which caused the decline to continue.[4]:332 Farming activities led to the extinction of the tammar on Flinders Island and St. Peter’s Island. In the 19th century, tammars were so common on the Eyre Peninsula and around Adelaide that battues were organised to protect crops and pastures by destroying large numbers of the animals. This has caused drastic declines, resulting in the tammar becoming extinct in the areas around Adelaide by the 1920s and in the Eyre Peninsula probably by the 1970s.[4]:332-33 However the genotype of those tammars survived, as they were introduced to New Zealand on Kawau Island and Rotorua.

In 2003, 85 New Zealand tammars were imported to South Australia and housed at the Monarto Zoo before being reintroduced to the Yorke Peninsula in Innes National Park.[10] Tammars have also been successfully translocated to other areas, such as Nambung National Park and Avon Valley National Park.[1] Since their introduction to New Zealand, tammars have flourished and their over-grazing has damaged indigenous plants. Biologists have used sodium fluoroacetate to control their populations.[11] In 1985, tammars were introduced to the North Island of the Houtman Abrolhos and have made similar impacts on native vegetation.[12] Their numbers have grown up to 450 individauls, but culling efforts appear to have reduced their numbers to 25 individuals by 2008.[12] The tammar wallaby is listed as of Least Concern by the IUCN, due to "its wide distribution, presumed large population, occurrence in a number of protected areas, and because it is unlikely to be declining at nearly the rate required to qualify for listing in a threatened category".[1]

[edit] Description

Taxidermy of tammar wallaby and joey

The tammar wallaby is the smallest wallaby species. It has a small head and large ears; the tail is long and thick at the base.[13] In colour, it is a dark, grizzled grey-brown above, becomes rufous on the sides of the body and the limbs, particularly in males, and becomes pale grey-buff below.[5] The tammar wallaby exhibits significant sexual dimorphism, with the maximum recorded weight in males being 9.1 kg (20 lb) and maximum recorded weight in females is 6.9 kg (15 lb). The body length is 59–68 cm (23–27 in) in males and 52–63 cm (20–25 in) in females. Both males and females are about 45 cm (18 in) in height. The tails of males range from 34–45 cm (13–18 in) and that of females from 33–44 cm (13–17 in).[13]

[edit] Locomotion

As with most macropods, the tammar wallaby moves around by hopping. This species has a hopping frequency of 3.5 strides per second, with a stride length of 0.8–2.4 m (2.5–8 ft).[14] When hopping, the proximal muscles produce most of the power for the jump and that power is delivered by the ankle through multi-joint muscles.[15] Leaping forward represents kinetic energy and landing back on the ground represents potential energy. As its feet hit the ground, the energies are converted into the elastic strain energy of its stretching leg tendons.[16] The elastic recoil that propels the tammar back off the ground, can recover much of this energy for reuse.[16] As such, a tammar can increase its hopping speed without losing more energy. In fact, the amount of energy stored in the tendons increases with the speed and the weight of the load the animal is carrying. This is particularly helpful for mothers carrying young.[17]

[edit] Senses

The tammar has 324 degree peripheral vision with 50 degree binocular vision.[4]:312 Its ability to discriminate black/white gratings of different widths and in different light intensities is stronger than those of most other small mammals such as rabbits.[18] It is nevertheless slightly weaker than a cat’s and much weaker than a human’s. In addition, it appears that the tammar has some colour vison. Its eyes have only the blue sensitive and green sensitive photoreceptor cones.[19] It can thus see colour in the blue-green band of the colour spectrum, but cannot see higher wavelengths in the red-yellow band. Nevertheless, in the band where it can see colour, it can discriminate between two monochromatic colours that are only 20 nanometres apart.[19] The pinna of the tammar is mobile and it relies on its directional position when detecting sounds. When a wallaby points its pinna at a sound source, the eardrum’s acoustic pressure increases 25-fold with an optimum frequency of 3–8 kHz.[20] When the pinna is not pointed at a sound source, the amplification falls sharply. As such, the tammar can detect where a sound is coming from when it moves its pinna, and can hear sounds of different frequencies and intensities from different parts of its surroundings.[20] A tammar's olfactory system is already developed at birth; this allows the nearborn to find its mother's pouch by the scent.[21]

[edit] Ecology and life history

The tammar is mostly nocturnal.

On Kangaroo Island, tammar wallabies shelter under divaricating bushes which cover most of the island. Tammars are nocturnal, being most active after dusk and before dawn. During this time, they forage for acacia seeds in the shrubs and feed on grassy clearings or farmland pasture.[4]:335 Tammars live in well-defined home ranges of 42 ha (100 acres) in summer and 16 ha (40 acres) in winter, and the ranges overlap with each other.[22] During the day tammars occupy a area of scrub for shelter from the sun, and then at night move to an area of grassland. In summer, grass dries off and the tammars move over greater distances than in winter to find higher quality food.[22] They are primarily grazers and have "a high digesta throughput".[23] When grazing, a tammar moves its lower jaw from side to side to grind its food. It appears that the digestion particles in the animal's foregut "expend relatively greater masticatory effort".[23] In addition, "secondary" fermentation may occur the hind gut. Compared to browsing macropods of similar size, tammars have a longer colon and shorter small intestine.[23]

Tammar resting

The tammar rests in two different positions: firstly, the "sitting tail forward" position, where the hind legs are outstreached with tail brought forward between, the body prone and the head above ground; secondly the "lying" position, where animal lays on its side with the head touching the ground. The former is of long duration, and done mostly during the day, while the latter is brief and done mostly at night.[23] Tammars are gregarious and have stable group activities, in which members of a group continue to associate on a long term basis.[24] Gathering in groups lessens the chance of an individual being taken by a predator. As the group size increases, tammars spend more time foraging, grooming and interacting and less time being vigilant and moving about.[25] Predators of the tammar include dingos, feral cats, red foxes and wedge-tailed eagles. It may also have been preyed upon by the now extinct thylacine. Tammars appear to respond more to the sight than the sound of predators. When a predator is detected, a tammar will alert conspecifics by thumping its foot.[26] When lost, young tammars are known to emit a coarse screeching call and adult females may give a similar call in response.[24]

Tammars maintain their body temperature, against high ambient temperatures, by licking their forearms and panting. When the ambient temperature is above 30 degrees Celsius, the respiratory system of the tammar increases steeply, and evaporative water loss also increases. When the ambient temperature is above 40 degrees Celsius, it can no longer control its body temperature and will die unless it can get away from the heat.[27] Tammars pass little water as urine, and can reabsorb water from the distal colon, which causes them to have only about 40 percent water content in their feces. This allows them to avoid dehydration.[4]:335 Tammars live on several islands that have no fresh water and must subsist on seawater or the moisture in plants.[4]:335 Tammars have been observed going to the nearest beach to drink seawater. They can live on seawater because their renal concentrating capacity is far greater than those of other small macropods.[28]

[edit] Breeding and development

Mother wallaby with young in pouch

The tammar wallaby has a promiscuous mating system, with females mating with multiple males.[29] Breeding is strictly seasonal and mating is highly synchronised, with a large proportion of births occurring in late January and early February.[30]:77 Females have a post-partum estrous cycle within hours of giving birth, and all mating activity concludes within four hours of parturition, when the wallabies are undisturbed.[29] The testis size of male tammars have little seasonal variation.[31] Concentrations of sperm and testosterone peak at the onset of the breeding season, but are low the rest of the year.[31] Around two weeks before the females give birth, the males will begin sniffing the uro-gential openings and pouches of the females to evaluate their reproductive status.[30]:78 Females allow males to mate with them soon during post-partum. However, a male that attempts to mate with an estrous female may face aggression from other males, which can delay ejaculations.[29] Mate-guarding is an important contributor to male reproductive success.[30]:83 During the estrous period, a clear dominance hierarchy is established. The more dominant males spend more time guarding, checking and disrupting the consortships of the estrous females.[30]:83 A post-partum female may be pursued by several males; these are known as "mating chases".[29]

The female tammar conceives a few hours after giving birth and suckles her young in her pouch for the next seven months. Tammars undergo embryonic diapause. During the first five months of her lactation, the dormant embryo in her uterus will only develop if the young in her pouch, also known as a joey, is lost or removed.[4]:338 However, after the antipodean winter solstice in June, the embryo will not develop if the pouch young is removed, nor does it develop when the joey leaves the pouch in October, but instead continues to remain dormant in the uterus until after the summer solstice in December.[4]:338 Gestation in the tammar lasts 26.5 days. At this time, all four of the female's mammary glands prepare for lactation.[32] After birth, the underdeveloped joey climbs into the pouch and attaches to the teat of one mammary gland and the others regress in lactation. The joey remain sattached to the teat for 100–125 days.[32] After which, it ceases its attachment to the teat, but remains in the pouch and continues to suckle at frequent but intermittent intervals. The joey begins to leave the pouch and eat grass at 200 days.[32] It begins to suckle more vigorously but less frequently. The joey leaves the pouch permanently after 250 days and is fully weaned at around 300–350 days.[32]

[edit] Sudden deaths

In late 1998, about 120-230 tammar wallabies died suddenly in research facilities and zoos in New South Wales and Queensland, perishing less than twelve hours after exhibiting the first symptoms, with most exhibiting no symptoms at all. Post mortem examination revealed haemorrhage of muscle, and abdominal and thoracic organs. The syndrome is known as Tammar Sudden Death Syndrome (TSDS) and the pathogen is an orbivirus of the family Reoviridae. It does not occur south of Sydney. Captive populations of tammar wallabies in New South Wales have subsequently suffered infections in summer months.[33]

[edit] Use in science

[edit] Genome sequencing

Tammars are easy to keep in captivity.

The tammar wallaby was the second marsupial to have its genome sequenced, after the gray short-tailed opossum and the first Australian marsupial to be sequenced (although it was the second published, after the fat-tailed dunnart).[34] The genome of a marsupial is of great interest to scientists studying comparative genomics, because marsupials are at an ideal degree of evolutionary divergence from humans: mice are too close and have not developed many different functions, while birds are genetically too remote.[35] The tammar wallaby is one of the model organisms as it is cheap to maintain and relatively tractable to scientific study.[35] The tammar genome is about 3.6 Gb, with a relatively short genetic map length of 1172 cM.[36] Researchers managed to isolate tammar bacterial artificial chromosomes (BACs) containing key immune genes from the tammar BAC library.[37] Using fluorescence in situ hybridisation (FISH), BACs containing T cell receptor (TCR) and immunoglobulin (Ig) genes were physically mapped in tammar wallaby chromosomes.[37] This research has highlighted the conservation of the genomic context of these important immune genes in marsupials.[37] Human and tammar genome comparisons have led to the discovery of new human genes, such as RBMX and several related genes.[34]

[edit] Antibiotics

Australian scientists, led by Dr. Ben Cocks, have found a compound in the milk of the tammar wallaby called AGG01, which has the potential to be a new and powerful form of antibiotic. AGG01 is a protein and, in laboratory testings, it has been shown to be 100 times more effective than penicillin, killing over 99% of the pathogenic bacteria (both gram-positive and gram-negative) and fungus that it was incubated with, including Salmonella, Proteus vulgaris and golden staphylococcus.[38]

[edit] References

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  2. ^ Pearson, M. (2005). Great Southern Land: The maritime exploration of Terra Australis. Department of the Environment and Heritage, Government of Australia. ISBN 0642551855. Archived from the original on 29 September 2007. http://web.archive.org/web/20070929132830/http://www.environment.gov.au/heritage/publications/pubs/great-southern-land.pdf. Retrieved 21 December 2007. 
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  33. ^ Kirkland, P. D. (2005). "Epidemic Viral Diseases in Wildlife". Microbiology Australia 26 (2): 82–85. ISSN 1324-4272. http://books.google.com.au/books?id=haCr7ztxTrwC&pg=PA82&dq=Tammar+Wallaby#v=onepage&q=Tammar%20Wallaby&f=false. 
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  37. ^ a b c Sanderson, C. E.; Belov, K.; Deakin, J. E. (2009). "Physical mapping of immune genes in the tammar wallaby (Macropus eugenii)". Cytogenet Genome Res 127 (1): 21–25. doi:10.1159/000260372. 
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[edit] External links

Data related to Macropus eugenii at Wikispecies Media related to Macropus eugenii at Wikimedia Commons

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