Open main menu

Antechinus /æntɪˈkaɪnəs/ ('ant-echinus') is a genus of small dasyurid marsupial endemic to Australia. They resemble mice with the bristly fur of shrews. They are sometimes also called broad-footed marsupial mice, pouched mice, route rat and/or Antechinus shrews. However, these common names are considered either regional or archaic and the modern common name for the animals is Antechinus.

Antechinus
Agile Antechinus (Antechinus agilis) on cloth, close-up from front.jpg
Scientific classification e
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Infraclass: Marsupialia
Order: Dasyuromorphia
Family: Dasyuridae
Subfamily: Dasyurinae
Tribe: Phascogalini
Genus: Antechinus
Macleay, 1841
Type species
Antechinus stuartii
Macleay, 1841
Species

Antechinus have short fur and are generally greyish or brownish in colour, varying with species.[1] The fur is dense and generally soft. Their tails are thin and tapering and range from slightly shorter to slightly longer than body length.[1] Their heads are conical in shape and ears are small to medium in size.[1] Some species have a relatively long, narrow snout that gives them a shrew-like appearance.[1] Species vary from 12–31cm in length and weigh 16–170 g when fully grown.[1] A. agilis is the smallest known species, and A. swainsonii the largest.

Sexual dimorphism occurs in most species for both weight and skeletal measurements, with males being typically larger and heavier.[2]

Most species nest communally in tree-hollows.[1] They primarily inhabit all forests, woodlands and rainforest as well as heaths and grasslands in some species.[1] The majority of Antechinus species are located on the eastern coast of Australia along the Great Dividing Range.[1] There is a population of A. flavipes in south west Western Australia.[3] A. bellus lives in northern Australia around the Gulf of Carpentaria.[3]

TaxonomyEdit

There are currently 15 recognised species of Antechinus with a number of subspecies.[3] A few species of New Guinean Antechinus were recognised, however, these have been reclassified into the genus Murexia.[4] The interspecific relations of the genus Antechinus are still under review.[3] The currently accepted phylogeny is the presence of four clades within the Antechinus genus.[3][5] The clades generally are formed by species with similar geographic distributions.[3]

Clade 1 (dusky antechinus)

Clade 2

Clade 3 (brown antechinus)

Clade 4

DietEdit

 
Illustration from Brehms Tierleben, showing an unknown species of Antechinus

Antechinus are mainly insectivorous, but the exact composition of their diet can vary by species and habitat.[10][11] Antechinus mainly eat beetles, insect larvae and spiders.[10][12] Amphipods, millipedes and centipedes are also quite common in their diets.[10][12] It has been found that after a fire, ants make up the majority of their diet - this is thought to be because ants are the only insect present in any number after the fire.[13] It has been reported that Antechinus also eat vertebrates, mainly small reptiles, such as skinks, or mammals, such as feathertail gliders.[1][14] This is likely to occur most commonly when the Antechinus are food stressed.[14] The remains of plants have been found in Antechinus stool, likely accidentally ingested while searching for other food.[12] Studies on Antechinus diet usually use faecal samples which may be unreliable in detecting soft bodied prey.[10][12]

Antechinus are usually classified as dietary generalists as they eat a wide variety of invertebrates as well as some vertebrates.[1][12][15]. They are also classified as opportunists because they feed on most of the prey available to them[1][14][16] However, they do show preference for some prey, i.e. beetles, spiders and larvae, especially when they are not food stressed.[10][14]

The microhabitat and foraging techniques vary between species.[11] Smaller species, such as A. stuartii, are scansorial and mainly hunt in trees.[14] They have been observed jumping between branches to catch flying insects.[17] The larger species of Antechinus, such as A. swainsonii, are completely ground dwelling and forage in the leaf litter.[14][17]

The efficiency of Antechinus hunting increases with their age as learn which prey are the best to eat[1]. Antechinus have been observed scraping slugs on rocks and other objects to remove the mucus and make them more palatable[1].

ReproductionEdit

Antechinus have an extremely unusual reproductive system. The females are synchronously monoestrous with mating occurring over a short 3 week period.[2][18] The males experience mass mortality after mating, with male survival only observed in very rare cases.[18] Females often mate twice and in some cases three times in their lives.[19] The gestation period varies by species between 25 and 35 days.[18] The offspring are independent after about 90-100 days, depending on the species.[2] This development period is rather long compared with other, similarly sized marsupials.[2]

Timing of the breeding seasonEdit

The timing of the breeding season differs between species and also with the location of populations.[18][20] The breeding season is in winter or early spring, from July to September.[21] The timing of the breeding season changes very little at the same location between different years.[21] The rate of change of photoperiod, the length of day, determines the commencement of the breeding season.[18] Different species respond to different critical rates of change, for example 80% of A. stuartii ovulate when the photoperiod is increasing by a rate of 97-117s/day.[18] The critical rate also varies by location within species.

Antechinus live in the relatively stable environments along the east coast of Australia.[18] Responding to the photoperiod, rather than temperature or rainfall, allows Antechinus to have young developing before the seasonal increase of food experienced rather than waiting for the increase of food and possibly missing the window of opportunity. The timing of mating ensures that the offspring are weaning when there is a large amount of food available in the environment.[21] The timing of this increase in food changes throughout the different ranges of the various species of Antechinus, a reason why different species have different mating seasons.[18]

Many species of Antechinus occur sympatrically.[18] Whenever two or more species of Antechinus occur together, the critical rate of change of photoperiod which triggers ovulation differs between the species.[18][22] This reproductive isolation may have lead to sympatric speciation.[18] There is a possible genetic link that would allow for the selection of specific responses to different rates of change of photoperiod.[18] The slightly different timings of reproduction by these sympatric species may also mean that they can take advantage of different increases of food in the spring and summer[18]. Larger species of Antechinus usually mate first meaning that the smaller species may have evolved later mating times to reduce competition and capitalise on later increases in food[18].

A. swainsonii and A. minimums occur in Tasmania and their reliance on the rate of change of the photoperiod for reproduction is less certain.[18] This is because in higher latitudes the photoperiod changes much faster. Animals that mate when the photoperiod is changing by 35-90s/day would only have 2-3 days in Tasmania compared with 2 weeks in New South Wales.[18] 

Females control the synchrony of mating, with males reproductively mature and ready to mate 4-5 weeks before the breeding season.[21][23] Males have been selected to be ready before the females to ensure the maximum amount of time for mating.

 
Pouch of agile antechinus (A. agilis), showing offspring inside

Reproductive anatomyEdit

Antechinus do not have a complete pouch as in other marsupials but simply a flap of skin covering the teats.[1] The number of teats in Antechinus species varies between different populations of the same species.[2][19] The number of teats can be anywhere from 6 to 13.[19] Populations in which females have 6 teats are found in lower latitudes where the environment is more stable. In higher latitudes, there are populations with 8 to 12 nipples. Odd numbers of teats only occur in transition zones and probably occur when populations with a different number of teats mate.[19] The number of nipples is thought to correlate with the number of young that can be supported. In more seasonal environments, where more nipples occur, there is a larger increase in the availability of food for supporting the young.[19] Females with fewer nipples are more likely to survive until the next breeding season. This is thought to be because it is less stressful to raise a smaller litter.[19] The rate of survival of females to reproduce twice differs between species. The larger species, e.g. A. swainsonii are more likely to survive to reproduce for a second time.[19]

Mating behaviourEdit

Mating is intense for Antechinus and can last up to 12 hours in some species.[2][24] The males mate with multiple females and the litters have multiple fathers.[2] During the short breeding window males expand their home range and are often active during night and day.[24]

Males disperse from the nests once they are physiologically independent. Mothers initiate this dispersal but are tolerant of unrelated males in the nest. Inbreeding avoidance is likely to explain this behaviour.[20]

Male semelparityEdit

Male die-off occurs because of an increase in free corticosteroids in the blood,[24] which causes a suppression of the immune system and gastrointestinal ulcers and which result in male mortality.[24][25][26] An increase in free corticosteroids is thought to allow males to utilise their reserve energy and maximise their reproductive effort, even though the increase is usually fatal.[24] If there was no male die-off there would still be a small likelihood of males surviving to the next mating period.[18] Thus, it is far better for the males to invest heavily in one breeding season than attempt to survive to the next one.

There are a few possible evolutionary advantages to evolve synchronous mating. It may ensure that as many matings as possible occur during the period of time. It also ensures that males can put all their effort into one short breeding season. Another possible advantage of synchronous mating is to swamp predators with large amounts of offspring after weaning.[23]

TorporEdit

Torpor is periodic lowering of body temperature and metabolic rate to reduce energy consumption.[27] Many marsupials undergo torpor as well as some birds and placental mammals.[27] There are two types of torpor: hibernation which is long term (weeks or months) and daily torpor which is usually only a few hours.[27] Daily torpor involves a less extreme lowering of body temperature and metabolic rate than hibernation. Antechinus undergo daily torpor.[28]

Unlike hibernation, daily torpor is not simply reliant on ambient air temperatures.[29] Antechinus can move into torpor on summer days with temperatures in the range of 25-30°C.[29] This is called spontaneous torpor. Induced torpor occurs when food and water are restricted – this is most easily observed in a laboratory environment as restriction of food in the wild is hard to determine[27]. Torpor of Antechinus in the wild is likely to be the result of a combination of factors.[29]

Antechinus are small mammals and thus have a high surface/volume ratio which results in high heat loss.[29] They also have a high metabolic rate and normothermic temperatures of around 35°C.[30]. Going into torpor allows them to greatly reduce metabolic rates, sometimes up to 80%[27] This reduces both the food and water requirements of the animal.

Torpor in Antechinus usually occurs during rest periods, either in the early morning after nightly foraging or the late afternoon before nightly foraging.[27][31] However, going into torpor during the night is not uncommon.[13] Torpor allows the animals to conserve energy while they are not foraging. The substantial savings in water requirements during torpor may also help Antechinus cope with droughts.[27]

Body mass is an important factor for the induction of torpor, with larger animals less likely to go into torpor and more likely to stay in torpor for shorter periods of time.[29] This means that males go into torpor less frequently than females. Lactating females do not go into torpor.[29] Torpor in Antechinus can last for one to nine hours and in extreme cases nearly 20 hours.[27] The body temperatures of these animals drop significantly.[31] In female A. flavipes the minimum body temperature during torpor can be around 20°C.[30][31] In males it is higher, around 30°C.[30][31] Smaller animals have a lower body temperature during torpor.[27]

An increase in the induction of torpor in Antechinus has been found to occur after intense bushfires.[13] Intense bushfires destroy the dense undergrowth that provide Antechinus with shelter and food. After intense bush fires Antechinus were recorded as eating mainly ants, which usually forms a very small part of their diet.[13] Increasing the occurrences of torpor, in both males and females, is thought to help with survival after an intense bushfire by reducing the need for foraging and thus avoiding predators.[13] It has been shown that smoke, ash and charcoal provide a cue for torpor induction.[32]

Increasing body temperature after torpor is energetically costly and reduces some of the metabolic savings.[27] However, animals have been observed basking to help increase their body temperature without increasing their food intake.[13]

Females that have survived to a second year go into torpor much more frequently than females that are in their first year, even though second-year females are larger.[31] It is thought that this is because first-year females are still growing and thus need more food which requires more time spent foraging.[31] Older females are also more experienced at foraging and may meet their energy requirements faster.[31][1]  

ThreatsEdit

A. arkos and A. argentus are currently listed as endangered.[33][34] The main threats to these species, indeed all species of Antechinus, are habitat destruction and introduced animals.[35]

Deforestation and habitat destruction removes the complex understorey habitat which Antechinus require for protection from predators and for food.[35][36] Antechinus also nest in tree hollows, which only form in senescent trees.[37][38]

Introduced predators also threaten Antechinus populations with foxes and cats predating upon Antechinus.[35] There is also competition between Antechinus and introduced rats and mice for habitat and food which may be detrimental to Antechinus numbers.[39] Pigs, cattle and horses trample their habitats.[1]

Changed fire regimes since European arrival have resulted in more intense bushfires which can negatively impact Antechinus populations by destroying understorey vegetation and removing most of their preferred food.[13][36]

Climate change threatens several high altitude species, especially those in northern Queensland.[1]

ReferencesEdit

  1. ^ a b c d e f g h i j k l m n o p q Baker, Andrew; Dickman, Chris (2018). Secret Lives of Carnivorous Marsupials. doi:10.1071/9781486305155. ISBN 9781486305155.
  2. ^ a b c d e f g Marlow, BJ (1961). "Reproductive Behaviour of the Marsupial Mouse, antechinus flavipes (Waterhouse) (Marsupialia) and the development of the Pouch young". Australian Journal of Zoology. 9 (2): 203. doi:10.1071/zo9610203. ISSN 0004-959X.
  3. ^ a b c d e f Mutton, Thomas Y; Phillips, Matthew J; Fuller, Susan J; Bryant, Litticia M; Baker, Andrew M (2019-04-27). "Systematics, biogeography and ancestral state of the Australian marsupial genus Antechinus (Dasyuromorphia: Dasyuridae)". Zoological Journal of the Linnean Society. 186 (2): 553–568. doi:10.1093/zoolinnean/zly062. ISSN 0024-4082.
  4. ^ Grossek, G. L.; Woolley, P. A.; Menzies, J. I. (2010). "The diet of some New Guinean dasyurid marsupials". Australian Mammalogy. 32 (2): 145. doi:10.1071/am10002. ISSN 0310-0049.
  5. ^ Baker, Andrew; Mutton, T.; Mason, E.; Gray, E. (2015). "A taxonomic assessment of the Australian Dusky Antechinus Complex: a new species, the Tasman Peninsula Dusky Antechinus (Antechinus vandycki sp. nov.) and an elevation to species of the Mainland Dusky Antechinus (Antechinus swainsonii mimetes (Thomas))". Memoirs of the Queensland Museum - Nature. 59: 75–126. doi:10.17082/j.2204-1478.59.2015.2014-10. ISSN 0079-8835.
  6. ^ Baker; Mutton; Mason; Gray (2015). "A taxonomic assessment of the Australian Dusky Antechinus Complex: a new species, the Tasman Peninsula Dusky Antechinus (Antechinus vandycki sp. nov.) and an elevation to species of the Mainland Dusky Antechinus (Antechinus swainsonii mimetes (Thomas))". Memoirs of the Queensland Museum. 59: 75–126. doi:10.17082/j.2204-1478.59.2015.2014-10.
  7. ^ Baker; Mutton; Mason; Gray (2015). "A taxonomic assessment of the Australian Dusky Antechinus Complex: a new species, the Tasman Peninsula Dusky Antechinus (Antechinus vandycki sp. nov.) and an elevation to species of the Mainland Dusky Antechinus (Antechinus swainsonii mimetes (Thomas))". Memoirs of the Queensland Museum. 59: 75–126. doi:10.17082/j.2204-1478.59.2015.2014-10.
  8. ^ Andrew M Baker, Thomas Y Mutton & Harry B Hines (2013). "A new dasyurid marsupial from Kroombit Tops, south-east Queensland, Australia: the silver-headed antechinus, Antechinus argentus sp. nov. (Marsupialia: Dasyuridae)". Zootaxa. 3746 (2): 201–239. doi:10.11646/zootaxa.3746.2.1.
  9. ^ Baker, A.M.; Mutton, T.Y.; Dyck, S. van 2012: A new dasyurid marsupial from eastern Queensland, Australia: the buff-footed Antechinus, Antechinus mysticus sp. nov. (Marsupialia: Dasyuridae). Zootaxa, (3515): 1-37. Preview
  10. ^ a b c d e Hall, S (1980). "The Diets of Two Coexisting Species of Antechinus (Marsupialia: Dasyuridae)". Wildlife Research. 7 (3): 365. doi:10.1071/wr9800365. ISSN 1035-3712.
  11. ^ a b Green, K (1989). "Altitudinal and Seasonal Differences in the Diets of Antechinus-Swainsonii and Antechinus-Stuartii (Marsupialia, Dasyuridae) in Relation to the Availability of Prey in the Snowy Mountains". Wildlife Research. 16 (6): 581. doi:10.1071/wr9890581. ISSN 1035-3712.
  12. ^ a b c d e Allison, Louise M.; Gibson, Lesley A.; Aberton, John G. (2006). "Dietary strategy of the swamp antechinus (Antechinus minimus maritimus) (Marsupialia:Dasyuridae) in coastal and inland heathland habitats". Wildlife Research. 33 (1): 67. doi:10.1071/wr05038. ISSN 1035-3712.
  13. ^ a b c d e f g Matthews, Jaya K.; Stawski, Clare; Körtner, Gerhard; Parker, Cassandra A.; Geiser, Fritz (2016-10-12). "Torpor and basking after a severe wildfire: mammalian survival strategies in a scorched landscape". Journal of Comparative Physiology B. 187 (2): 385–393. doi:10.1007/s00360-016-1039-4. ISSN 0174-1578.
  14. ^ a b c d e f Lunney, Daniel; Matthews, Alison; Grigg, Jan (2001). "The diet of Antechinus agilis and A. swainsonii in unlogged and regenerating sites in Mumbulla State Forest, south-eastern New South Wales". Wildlife Research. 28 (5): 459. doi:10.1071/wr00015. ISSN 1035-3712.
  15. ^ Mason, Eugene D.; Burwell, Chris J.; Baker, Andrew M. (2015). "Prey of the silver-headed antechinus (Antechinus argentus), a new species of Australian dasyurid marsupial". Australian Mammalogy. 37 (2): 164. doi:10.1071/am14036. ISSN 0310-0049.
  16. ^ Fox, BJ; Archer, E (1984). "The Diets of Sminthopsis Murina and Antechinus Stuartii (Marsupialia: Dasyuridae) in Sympatry". Wildlife Research. 11 (2): 235. doi:10.1071/wr9840235. ISSN 1035-3712.
  17. ^ a b Dickman, Chris R. (1991), "Use of trees by ground-dwelling mammals: implications for management", Conservation of Australia's Forest Fauna, Royal Zoological Society of New South Wales, pp. 125–136, doi:10.7882/rzsnsw.1991.012, ISBN 0959995153
  18. ^ a b c d e f g h i j k l m n o p q McAllan, Bronwyn M.; Dickman, Christopher R; Crowther, Mathew S. (2006-02-24). "Photoperiod as a reproductive cue in the marsupial genus Antechinus: ecological and evolutionary consequences". Biological Journal of the Linnean Society. 87 (3): 365–379. doi:10.1111/j.1095-8312.2006.00571.x. ISSN 0024-4066.
  19. ^ a b c d e f g Cockburn, Andrew; Lee, Anthony K.; Martin, Roger W. (1983). "Macrogeographic Variation in Litter Size in Antechinus (Marsupialia: Dasyuridae)". Evolution. 37 (1): 86. doi:10.2307/2408177. ISSN 0014-3820. JSTOR 2408177.
  20. ^ a b Cockburn, Andrew; Scott, Michelle P.; Scotts, David J. (1985). "Inbreeding avoidance and male-biased natal dispersal in Antechinus spp. (Marsupialia: Dasyuridae)". Animal Behaviour. 33 (3): 908–915. doi:10.1016/s0003-3472(85)80025-7. ISSN 0003-3472.
  21. ^ a b c d Dickman, Chris R. (1985). "Effects of photoperiod and endogenous control on timing of reproduction in the marsupial genus Antechinus". Journal of Zoology. 206 (4): 509–524. doi:10.1111/j.1469-7998.1985.tb03555.x. ISSN 0952-8369.
  22. ^ McAllan, B. M.; Dickman, C. R. (1986). "The role of photoperiod in the timing of reproduction in the Dasyurid Marsupial Antechinus stuartii". Oecologia. 68 (2): 259–264. doi:10.1007/bf00384797. ISSN 0029-8549.
  23. ^ a b Scott, M. P. (1986-08-08). "The Timing and Synchrony of Seasonal Breeding in the Marsupial, Antechinus stuartii: Interaction of Environmental and Social Cues". Journal of Mammalogy. 67 (3): 551–560. doi:10.2307/1381287. ISSN 1545-1542. JSTOR 1381287.
  24. ^ a b c d e Scott, M. P. (1987-08-28). "The Effect of Mating and Agonistic Experience on Adrenal Function and Mortality of Male Antechinus stuartii (Marsupialia)". Journal of Mammalogy. 68 (3): 479–486. doi:10.2307/1381584. ISSN 1545-1542. JSTOR 1381584.
  25. ^ McDonald, I.R.; Lee, A.K.; Bradley, A.J.; Than, K.A. (1981). "Endocrine changes in dasyurid marsupials with differing mortality patterns". General and Comparative Endocrinology. 44 (3): 292–301. doi:10.1016/0016-6480(81)90004-6. ISSN 0016-6480.
  26. ^ BRADLEY, A. J.; McDONALD, I. R.; LEE, A. K. (1975). "Effect of Exogenous Cortisol on Mortality of a Dasyurid Marsupial". Journal of Endocrinology. 66 (2): 281–282. doi:10.1677/joe.0.0660281. ISSN 0022-0795.
  27. ^ a b c d e f g h i j Geiser, Fritz; Körtner, Gerhard (2010). "Hibernation and daily torpor in Australian mammals". Australian Zoologist. 35 (2): 204–215. doi:10.7882/az.2010.009. ISSN 0067-2238.
  28. ^ Wallis, Robert L. (1976). "Torpor in the dasyurid marsupial Antechinus stuartii". Comparative Biochemistry and Physiology Part A: Physiology. 53 (4): 319–322. doi:10.1016/s0300-9629(76)80147-8. ISSN 0300-9629.
  29. ^ a b c d e f Geiser, Fritz (1988). "Daily torpor and thermoregulation in antechinus (Marsupialia): influence of body mass, season, development, reproduction, and sex". Oecologia. 77 (3): 395–399. doi:10.1007/bf00378050. ISSN 0029-8549.
  30. ^ a b c Geiser, F (1994). "Hibernation and Daily Torpor in Marsupials - a Review". Australian Journal of Zoology. 42 (1): 1. doi:10.1071/zo9940001. ISSN 0004-959X.
  31. ^ a b c d e f g Rojas, A. Daniella; Körtner, Gerhard; Geiser, Fritz (2014-01-18). "Torpor in free-ranging antechinus: does it increase fitness?". Naturwissenschaften. 101 (2): 105–114. doi:10.1007/s00114-013-1136-0. ISSN 0028-1042.
  32. ^ Stawski, Clare; Nowack, Julia; Körtner, Gerhard; Geiser, Fritz (2017-01-15). "A new cue for torpor induction: charcoal, ash and smoke". The Journal of Experimental Biology. 220 (2): 220–226. doi:10.1242/jeb.146548. ISSN 0022-0949.
  33. ^ Baker, ANDREW M; Mutton, THOMAS Y; Hines, Harry B (2013-12-11). "

    A new dasyurid marsupial from Kroombit Tops, south-east Queensland, Australia: the Silver-headed Antechinus, Antechinus argentus sp. nov. (Marsupialia: Dasyuridae)

    ". Zootaxa. 3746 (2): 201. doi:10.11646/zootaxa.3746.2.1. ISSN 1175-5334.
  34. ^ Gray, Emma L.; Burwell, Chris J.; Baker, Andrew M. (2016). "Benefits of being a generalist carnivore when threatened by climate change: the comparative dietary ecology of two sympatric semelparous marsupials, including a new endangered species (Antechinus arktos)". Australian Journal of Zoology. 64 (4): 249. doi:10.1071/zo16044. ISSN 0004-959X.
  35. ^ a b c Stokes, Vicki L; Pech, Roger P; Banks, Peter B; Arthur, Anthony D (2004). "Foraging behaviour and habitat use by Antechinus flavipes and Sminthopsis murina (Marsupialia: Dasyuridae) in response to predation risk in eucalypt woodland". Biological Conservation. 117 (3): 331–342. doi:10.1016/j.biocon.2003.12.012. ISSN 0006-3207.
  36. ^ a b Lada, Hania; Mac Nally, Ralph (2008). "Decline and potential recovery of Yellow-footed Antechinus in parts of south-eastern Australia: A perspective with implications for management". Ecological Management & Restoration. 9 (2): 120–125. doi:10.1111/j.1442-8903.2008.00403.x. ISSN 1442-7001.
  37. ^ Dickman, Chris R.; Steeves, Tracey E. (2004), "Use of habitat by mammals in eastern Australian forests: are common species important in forest management?", Conservation of Australia's Forest Fauna, Royal Zoological Society of New South Wales, pp. 761–773, doi:10.7882/fs.2004.046, ISBN 095860858X
  38. ^ Naylor, R.; Richardson, S. J.; McAllan, B. M. (2008-01-22). "Boom and bust: a review of the physiology of the marsupial genus Antechinus". Journal of Comparative Physiology B. 178 (5): 545–562. doi:10.1007/s00360-007-0250-8. ISSN 0174-1578.
  39. ^ Glen, Alistair S.; Dickman, Chris R. (2005-04-13). "Complex interactions among mammalian carnivores in Australia, and their implications for wildlife management". Biological Reviews. 80 (3): 387. doi:10.1017/s1464793105006718. ISSN 1464-7931.