Content uploaded by Patricia Anne Fleming
Author content
All content in this area was uploaded by Patricia Anne Fleming on Nov 13, 2022
Content may be subject to copyright.
Foraging activity by the southern brown bandicoot
(Isoodon obesulus) as a mechanism for soil turnover
Leonie E. Valentine
A,B,C
, Hannah Anderson
A
, Giles E. StJ. Hardy
A
and Patricia A. Fleming
A
A
Western Australia Centre of Excellence for Climate Change, Woodland and Forest Health,
School of Veterinary and Life Sciences, Murdoch University, Perth, WA 6150, Australia.
B
Present address: ARC Centre of Excellence for Environmental Decisions, School of Plant Biology,
University of Western Australia, Crawley, WA 6009, Australia.
C
Corresponding author. Email: leonie.valentine@uwa.edu.au
Abstract. Mammals that forage for food by biopedturbation can alter the biotic and abiotic characteristics of their habitat,
influencing ecosystem structure and function. Bandicoots, bilbies, bettongs and potoroos are the primary digging marsupials
in Australia, although most of these species have declined throughout their range. This study used a snapshot approach to
estimate the soil turnover capacity of the southern brown bandicoot (Isoodon obesulus, Shaw 1797), a persisting digging
Australian marsupial, at Yalgorup National Park, Western Australia. The number of southern brown bandicoots was
estimated using mark–recapture techniques. To provide an index of digging activity per animal, we quantified the number of
new foraging pits and bandicoot nose pokes across 18 plots within the same area. The amount of soil displaced and physical
structure of foraging pits were examined from moulds of 47 fresh foraging pits. We estimated that an individual southern
brown bandicoot could create ~45 foraging pits per day, displacing ~10.74 kg of soil, which extrapolates to ~3.9 tonnes of soil
each year. The digging activities of the southern brown bandicoots are likely to be a critical component of soil ecosystem
processes.
Additional keywords: biopedturbation, ecosystem engineering, soil movement.
Received 8 October 2012, accepted 7 May 2013, published online 22 May 2013
Introduction
Mammals that move or manipulate soil for food or to create shelter
(biopedturbation) can act as ecosystem engineers (Whitford
1999), creating disturbances that may be essential for maintaining
ecosystem health (Eldridge and James 2009; Eldridge et al.2009).
Mammalian biopedturbation creates small-scale disturbances via
soil turnover (Whitford 1999; Eldridge et al.2012) and can
subsequently alter the physical properties of soil, including soil
compaction and water infiltration (Garkaklis et al.1998,2000,
2003). Several Australian marsupials dig, though the bettongs
(Bettongia spp., Aepyrymnus rufescens), potoroos (Potorous
spp.), bilbies (Macrotis spp.) and bandicoots (Perameles spp.,
Isoodon spp. and Echymipera rufescens) are the main marsupials
in Australia responsible for creating foraging pits (Martin 2003).
These marsupials are adapted to digging in soil, and use their
strong forefeet and claws to create foraging pits while searching
for food, such as invertebrates, tubers, seeds and fungi. The soil-
turnover capacity of these digging marsupials is impressive, with
individual woylies (Bettongia penicillata) estimated to displace
~4.8 tonnes of soil each year (Garkaklis et al.2004).
Australian digging marsupials (here defined as bettongs,
potoroos, bilbies and bandicoots) are all within the critical weight
range and considered most at risk from introduced predators
(Johnson and Isaac 2009), and most of these species have suffered
drastic declines in mainland populations and substantial range
contractions (Van Dyck and Strahan 2008). Of the 16 extant
digging marsupial species, 11 are considered to be of
conservation concern, while a third (5 species) are considered
critically endangered or endangered (Environment Protection
and Biodiversity Conservation Act 1999). Despite the grim
conservation status of most Australian digging marsupials,
several species (e.g. Isoodon macrourus,I. obesulus and
Perameles nasuta) persist within parts of their former range on
mainland Australia, sometimes in highly modified environments
(e.g. Hughes and Banks 2010). However, the potential ecosystem
role of these species has not been investigated.
The southern brown bandicoot (I. obesulus, Shaw 1797) is a
medium-sized omnivorous marsupial that occurs in scattered
areas across parts of eastern, southern and south-western
Australia (Van Dyck and Strahan 2008). Home-range estimates
for the southern brown bandicoot vary from 0.5 to 6.0 ha (Lobert
1990), with males typically having larger home ranges than
females (Heinsohn 1966), and in areas of high density (and
correspondingly high food supply) home ranges are likely to
Journal compilation ÓCSIRO 2012 www.publish.csiro.au/journals/ajz
CSIRO PUBLISHING
Australian Journal of Zoology, 2012, 60, 419–423 Short communication
http://dx.doi.org/10.1071/ZO13030
overlap (Broughton and Dickman 1991). Although the eastern
subspecies (I. obesulus obesulus) is listed as endangered
(Environment Protection and Biodiversity Conservation Act
1999), in south-western Australia the southern brown bandicoot
(I. obesulus fusciventer) is the only persisting commonly
occurring digging marsupial, especially within the
urban–wildland interface. Foraging pits are created by
bandicoots when digging with their strong forefeet for fungal
fruiting bodies, invertebrates and subterranean plant material
(Van Dyck and Strahan 2008). Previous observations have
indicated that southern brown bandicoots may be prolific
‘diggers’(Heinsohn 1966; Quin 1985).
The southern brown bandicoot occur in two distinct habitats in
south-western Australia –open forest, and dense vegetation
around swamps and watercourses (Cooper 2000a,2000b)–and
this mammal has consequently been identified as susceptible to
declining groundwater and rainfall (Wilson et al.2012). In the
urban–wildland interface surrounding Perth, populations of the
southern brown bandicoot persist in the bush fragments and
conservation reserves, often without predator control. In this
study, we quantified the physical structure of southern brown
bandicoot foraging pits and estimated soil turnover in a small area,
to compare with other digging marsupial species and to assist in
determining the potential role of the southern brown bandicoot in
maintaining ecosystem processes.
Materials and methods
Study site
This study was conducted at Martin’s Tank at the edge of Martin’s
Lake, Yalgorup National Park on the Swan Coastal Plain IBRA
region (Thackway and Cresswell 1995) in south-western
Australia (3250054.5200S, 1154008.7200 E). Yalgorup National
Park (~12 888 ha) has high regional biodiversity values based
around the chain of 10 coastal lakes, swamps and tuart
(Eucalyptus gomphocephala) forests (Portlock et al.1993).
Although sections of the national park are baited with 1080
(sodium fluoroacetate) to assist in the control of the introduced red
fox (Vulpes vulpes), the area surrounding Martin’s Lake is not
currently baited. The region has a Mediterranean-type climate
with hot dry summers and mild wet winters and an average annual
rainfall of 864 mm (Bureau of Meteorology, Lake Preston Lodge
2 Comp., #009679). Yalgorup National Park contains three major
dune systems: the Quindalup, Spearwood and Bassendean Dunes
(Portlock et al.1993). Our research focussed on foraging activity
and soil turnover of bandicoots within a small section of the
National Park, consisting of a 2-ha area (200 m 100 m) in the
vegetation running parallel to Martins Lake. Our study site was
located on Spearwood Dunes, where soils were predominantly
yellow-phase Karrakatta sands. Vegetation in the study area
included lake-fringing vegetation dominated by Melaleuca
preissiana and M. rhaphiophylla and interspersed with tuarts,
with a dense understorey of sedges (mostly Gahnia trifida)
transitioning to a combination of tuart trees, peppermint (Agonis
flexuosa) and paperbark (M. rhaphiophylla), and a tuart, jarrah
(E. marginata) and marri (Corymbia calophylla) overstorey with
a mid-storey layer of scattered Banksia grandis,B. attenuata and
grasstrees (Xanthorrhoea spp.), and an open understorey of
zamia palms (Zamia spp.) and various herbaceous species (e.g.
Jacksonia sternbergiana,Hibbertia hypericoides) (Portlock
et al.1993).
Estimating soil turnover by the southern brown bandicoot
Bandicoot foraging activity was assessed for 18 plots (each
10 m 10 m), with plots haphazardly stratified along the
vegetation gradient described above, with each plot separated
from each other by a minimum of 30 m. We counted the number of
new foraging pits and nose pokes created within each plot during a
24-h period in June and in August 2011. A bandicoot ‘foraging
pit’was defined as having a clear point at the bottom of the pit and
a spoil heap adjacent to the pit (where displaced soil was
accumulated via the digging activities of the bandicoots). A ‘nose
poke’was defined as an obvious movement of the ground debris
and soil but without a defined point or adjacent spoil heap. Due to
rain occurring in the days before examining foraging activity (but
not during the sample period), new foraging pits and nose pokes
were easily identified during both sampling sessions (as rain in the
previous day had left impressions in the spoil of existing foraging
pits).
After counting foraging pits (described above), we used
mark–recapture trapping (three nights in June and August 2011)
to estimate the number of southern brown bandicoots potentially
responsible for creating the foraging pits in the 2-ha study area. A
transect of 10 cage traps (Sheffields: 20 cm 20 cm 56 cm)
were spread evenly across the study area. All traps were baited
with universal bait (a combination of peanut butter, rolled oats,
sardines and truffle oil). Hessian bags and pieces of tarpaulin were
placed over all cage traps to provide shelter and to prevent rain
entering the cage. The traps were open in the afternoon each day
and checked within 3 h of sunrise the following morning. All
animals captured were weighed, measured (head length and
long pes), sexed and individually marked using ISO FDX-B
microchips (OzMicrochips, NSW) inserted subcutaneously on
the nape of the neck. Retrapped animals were detected using the
RT100 ISO Scanner (Real Trace, NSW). In this study we have not
assessed home-range sizes for the southern brown bandicoot,
although previous work in south-western Australia indicates that
home ranges are ~2.3 ha for males and ~1.8 ha for females, but
they may overlap (Broughton and Dickman 1991). As we did not
estimate the spatial range of the animals at Martins Tank, we used
the total number of animals captured (both trapping sessions
combined) as our estimate of the number of bandicoots creating
foraging pits within the 2-ha area.
The number of foraging pits was quantified by averaging the
number of new foraging pits per plot counted in June and August
2011 and extrapolating this value to a per-hectare estimate. Plaster
of Paris (Diggers Plaster of Paris, South Australia) was poured
into 47 fresh bandicoot diggings that were representative of the
range of foraging pit sizes observed in plots. We measured the
width (at soil surface) and depth of the plaster moulds, and the
volume of each mould (in millilitres) was estimated by water
displacement (1200 mL graduated cylinder). Measurements
reported are the average standard error. Soil density
(1.25 g cm
–3
) was estimated as the average density obtained from
four soil core samples of known volume (~1021 cm
3
) that were
420 Australian Journal of Zoology L. E. Valentine et al.
oven-dried for 72 h (K. Ruthrof, unpubl. data). The amount of soil
displaced by one bandicoot in a night was calculated as:
Soil displaced ðg individual1ð24-h periodÞ1Þ
¼ðno:of new foraging pits bandicoot1ð24-h periodÞ1Þ
ðforaging pit volumeÞðsoil densityÞ
This figure was also then expressed as tonnes
individual
–1
year
–1
.
Limitations to this study
Our study provides a snapshot approach at estimating the soil-
turnover capacity of the southern brown bandicoot, and has
several limitations that should be considered. First, we used a
single location, Martins Tank, to obtain our estimates of foraging
activity and foraging pit dimensions for the southern brown
bandicoot. These values may vary depending on location, habitat,
soil type and bandicoot density. Second, to estimate the number of
bandicoots creating the foraging pits, we have used the total
number of bandicoots captured within the 2-ha area. Given our
uncertainty of the spatial range of foraging bandicoots, the
foraging pits within our study area may have been created by one
or several bandicoots. Using the total number of captured
bandicoots may overestimate the number of bandicoots creating
the foraging pits and thus could represent a conservative estimate
of the soil-turnover capacity of this species. Third, our estimates
of foraging activity are based on two nights’data collection and
the extrapolation to an annual estimate of soil turnover does not
reflect seasonal differences in foraging behaviour and intensity.
Results
In total, eight bandicoot individuals were captured in the 2-ha area
over 60 trap-nights (June and August sessions combined). Six
bandicoots (two female, four male) were captured in June and
recaptured in August, along with an additional two individuals
(one male, one escaped before it was sexed). Males were typically
larger and heavier (n= 5, mean s.e.: body mass = 1724 107 g;
head length = 93.2 2.1 mm, pes length = 65.0 1.3 mm) than
females (n= 2, mean s.e.: body mass = 1165 15 g; head
length = 85.1 6.0 mm; pes length = 60.6 2.0 mm). The eight
individuals were all in visibly good condition, with no fur loss,
scratches or other signs of fighting.
Across the 18 survey plots there were 36 new foraging pits and
88 new nose pokes in June and 32 new foraging pits and 122 new
nose pokes in August, with a range of 0–6 foraging pits and 0–21
nose pokes observed per plot in both sampling periods. The mean
number of new foraging pits day
–1
averaged 1.8 plot
–1
(10 10 m), which extrapolated to 180 new foraging pits ha
–1
in a
24-h period. For the purposes of this study, we have assumed that
all eight individual southern brown bandicoots created the
foraging pits (i.e. 4 individual bandicoots ha
–1
), which equates to
45 foraging pits day
–1
(individual bandicoot)
–1
.
Moulds of 47 fresh foraging pits indicated that foraging pits
were fairly consistent in their physical size. Foraging pits were
conical in shape, measuring 100.9 3.9 mm across at the soil
surface with a mean depth of 69.6 3.2 mm (depth range
35–135 mm). The mean volume of these foraging pits was
191 15 mL. In a single night of our study, the soil displaced
by one bandicoot at Martins Tank was therefore estimated
as 8595 cm
3
or 10.74 kg (calculated as follows: 10 743.75 g
soil displaced individual
–1
(24-h period)
–1
= 45 foraging
pits bandicoot
–1
(24-h period)
–1
191 mL soil displaced
1.25 g cm
–3
soil density). Assuming no seasonal differences in
foraging activity, this value can then be extrapolated to an annual
turnover of 3.14 m
3
or 3.92 tonnes for each individual.
Discussion
Southern brown bandicoots are opportunistic omnivores that
forage for a variety of food, consuming invertebrates, fungi, plant
material and occasionally small vertebrates, with diets reflecting
seasonally and locally abundant food items (Heinsohn 1966;
Quin 1988; Van Dyck and Strahan 2008). Foraging of bandicoots
via nose pokes may assist bandicoots in detecting subterranean
prey items (Quin 1992) and/or target invertebrates (e.g.
cockroaches, crickets, spiders) that commonly occur in the leaf
litter layer (Hattenschwiler et al.2005). In Tasmania, a single wild
bandicoot was observed digging 21 foraging pits within 36 min
(Heinsohn 1966), while bandicoots in captivity have been
observed digging up to 32 foraging pits in an evening (Quin
1985). In our study, we estimated that a single bandicoot dug ~45
foraging pits each day, representing a considerable impact in
terms of soil turnover.
Bettongs and potoroos forage principally on fruiting bodies of
underground fungi (Van Dyck and Strahan 2008) and may create
higher numbers of foraging pits while searching for food (e.g.
woylie: 38–114 foraging pits individual
–1
(Garkaklis et al.2004);
southern brown bandicoot: ~45 foraging pits individual
–1
).
Although we did not examine the density of foraging pits
throughout seasons, previous research has indicated that the
densities of foraging pits of digging marsupials may vary
throughout the year, potentially in relation to the availability of
hypogeal fungal fruiting bodies (Claridge et al.1993). As the diet
of the southern brown bandicoot varies seasonally (Quin 1988),
the number of foraging pits created by this species is also likely to
vary seasonally. Foraging pits created by the greater bilby and
burrowing bettong are ~80 mm deep (James and Eldridge 2007),
similar in size to those of the southern brown bandicoot (~70 mm).
The long-nosed potoroo (P. tridactylus) creates foraging pits that
vary in depth from 56 to 120 mm (Claridge et al.1993), while the
woylie creates deeper foraging pits (100–115 mm: Garkaklis et al.
2004).
Although our study is restricted to a small area and represents a
‘snapshot’of foraging activities of the southern brown bandicoot,
it is the first to estimate soil turnover rates of the southern brown
bandicoot, with an individual bandicoot (average body mass
1.6 kg) turning over ~10.74 kg day
–1
. This equates to ~3.9 tonnes
of soil bandicoot
–1
year
–1
and falls within the range of soil
displaced (2.7–9.7 tonnes year
–1
) by the similar-sized woylie
(body mass: 1.0–1.5 kg) (Garkaklis et al.2004). Marsupials that
burrow for food and live underground produce even greater soil
turnover. For example, in predator-free enclosures in arid zones,
where bilbies and burrowing bettongs are held together (therefore
values are for both species combined), these animals excavate
~30 tonnes of soil individual
–1
year
–1
(Newell 2008).
Soil turnover by the southern brown bandicoot Australian Journal of Zoology 421
The loss of once widespread digging mammals in Australia is
likely to have major ramifications for ecosystem processes.
Further research on the foraging activities of the southern brown
bandicoot, preferably over a longer time frame and across several
sites, is necessary to elucidate the soil-turnover capacity of this
digging marsupial. Although the range and population of the
southern brown bandicoot has declined since European
settlement (Abbott 2008), these animals persist in urban,
periurban and rural regions of south-western Australia, where
they are likely to be playing an important role in ecosystem
processes, contributing to the health and function of the
woodlands and forests. Understanding the role of these animals
may therefore contribute towards conservation management
decisions. Since the southern brown bandicoot appears to be more
resilient to human-mediated disturbances compared with other
digging marsupials (e.g. woylie), they provide us with an ideal
opportunity to reintroduce them into landscapes where soil
turnover is required for ecosystem health and function.
Acknowledgements
We gratefully thank the Department of Environment and Conservation –Swan
Coastal District for their support with this project, especially Craig Olejnik,
Paul Tholen and Alan Wright. We also thank three anonymous reviewers for
substantially improving this manuscript. Our work was funded by the WA
State Centre of Excellence for Climate Change, Woodland and Forest
Health and the ARC Centre of Excellence for Environmental Decisions, and
was carried out with a Murdoch University Animal Ethics Committee permit
(W2341/10) and a WA Department of Environment and Conservation permit
(Regulation 17: SF001280).
References
Abbott, I. (2008). Historical perspectives of the ecology of some conspicuous
vertebrate species in south-west Western Australia. Conservation Science
Western Australia 6,1–214.
Broughton, S. K., and Dickman, C. R. (1991). The effect of supplementary
food on home range of the southern brown bandicoot, Isoodon obesulus
(Marsupialia: Peramelidae). Australian Journal of Ecology 16,71–78.
doi:10.1111/j.1442-9993.1991.tb01482.x
Claridge, A. W., Cunningham, R. B., and Tanton, M. T. (1993). Foraging
patterns of the long-nosed potoroo (Potorous tridactylus) for hypogeal
fungi in mixed-species and regrowth eucalypt forest stands in southeastern
Australia. Forest Ecology and Management 61,75–90. doi:10.1016/
0378-1127(93)90191-O
Cooper, M. L. (2000a). Random amplified polymorphic DNA analysis of
southern brown bandicoot (Isoodon obesulus) populations in Western
Australia reveals genetic differentiation related to environmental
variables. Molecular Ecology 9, 469–479. doi:10.1046/j.1365-294x.20
00.00883.x
Cooper, M. L. (2000b). Temporal variation in skull size and shape in the
southern brown bandicoot, Isoodon obesulus (Peramelidae: Marsupialia)
in Western Australia. Australian Journal of Zoology 48,47–57. doi:10.
1071/ZO99047
Eldridge, D. J., and James, A. I. (2009). Soil-disturbance by native animals
plays a critical role in maintaining healthy Australian landscapes.
Ecological Management & Restoration 10, S27–S34. doi:10.1111/j.14
42-8903.2009.00452.x
Eldridge, D. J., Whitford, W. G., and Duval, B. D. (2009). Animal
disturbances promote shrub maintenance in a desertified grassland.
Journal of Ecology 97, 1302–1310. doi:10.1111/j.1365-2745.2009.01
558.x
Eldridge, D. J., Koen, T. B., Killgore, A., Huang, N., and Whitford, W. G.
(2012). Animal foraging as a mechanism for sediment movement and soil
nutrient development: evidence from the semi-arid Australian woodlands
and the Chihuahuan Desert. Geomorphology 157–158, 131–141. doi:10.
1016/j.geomorph.2011.04.041
Garkaklis, M. J., Bradley, J. S., and Wooller, R. D. (1998). The effects of
woylie (Bettongia penicillata) foraging on soil water repellency and water
infiltration in heavy textured soils in southwestern Australia. Australian
Journal of Ecology 23, 492–496. doi:10.1111/j.1442-9993.1998.tb00
757.x
Garkaklis, M., Bradley, J., and Wooller, R. D. (2000). Digging by vertebrates
as an activity promoting the development of water-repellent patches in
sub-surface soil. Journal of Arid Environments 45,35–42. doi:10.1006/
jare.1999.0603
Garkaklis, M. J., Bradley, J. S., and Wooller, R. D. (2003). The relationship
between animal foraging and nutrient patchiness in south-west Australian
woodland soils. Australian Journal of Soil Research 41, 665–673.
doi:10.1071/SR02109
Garkaklis, M. J., Bradley, J. S., and Wooller, R. D. (2004). Digging and soil
turnover by a mycophagous marsupial. Journal of Arid Environments 56,
569–578. doi:10.1016/S0140-1963(03)00061-2
Hattenschwiler, S., Tiunov, A. V., and Scheu, S. (2005). Biodiversity and litter
decomposition in terrestrial ecosystems. Annual Review of Ecology
Evolution and Systematics 36, 191–218. doi:10.1146/annurev.ecolsys.
36.112904.151932
Heinsohn, G. E. (1966). Ecology and reproduction of the Tasmanian
bandicoots (Perameles gunni and Isoodon obesulus). University of
California Publications in Zoology 80,1–96.
Hughes, N. K., and Banks, P. B. (2010). Heading for greener pastures?
Defining the foraging preferences of urban long-nosed bandicoots.
Australian Journal of Zoology 58, 341–349. doi:10.1071/ZO10051
James, A. I., and Eldridge, D. J. (2007). Reintroduction of fossorial native
mammals and potential impacts on ecosystem processes in an Australian
desert landscape. Biological Conservation 138, 351–359. doi:10.1016/j.
biocon.2007.04.029
Johnson, C. N., and Isaac, J. L. (2009). Body size and extinction risk in
Australian mammals: back to the Critical Weight Range. Austral Ecology
34,35–40. doi:10.1111/j.1442-9993.2008.01878.x
Lobert, B. (1990). Home range and activity period of the southen brown
bandicoot (Isoodon obesulus) in a Victorian heathland. In ‘Bandicoots and
Bilbies’. (Eds J. H. Seebeck, P. R. Brown, R. L. Wallis and C.
M. Kemper.) pp. 319–325. (Surrey Beatty: Sydney.)
Martin, G. (2003). The role of small ground-foraging mammals in topsoil
health and biodiversity: implications to management and restoration.
Ecological Management & Restoration 4, 114–119. doi:10.1046/j.1442-
8903.2003.00145.x
Newell, J. (2008). The role of the reintroduction of greater bilbies (Macrotis
lagotis) and burrowing bettongs (Bettongia lesueur) in the ecological
restoration of an arid ecosystem: foraging diggings, diet and soil seed
banks. Ph.D. Thesis, School of Earth and Environmental Sciences,
University of Adelaide.
Portlock, C., Koch, A., Wood, S., Hanly, P., and Dutton, S. (1993). Yalgorup
National Park Management Plan. Department of Conservation and Land
Management, Perth, Western Australia.
Quin, D. G. (1985). Aspects of the feeding ecology of the bandicoots
Perameles gunnii (Gray 1838) and Isoodon obesulus (Shaw and Nodder
1797) (Marsupialia: Peramelidae) in southern Tasmania. B.Sc.(Honours)
Thesis. University of Tasmania, Hobart.
Quin, D. G. (1988). Observations on the diet of the southern brown bandicoot,
Isoodon obesulus (Marsupialia: Peramelidae), in southern Tasmania.
Australian Mammalogy 11,15–25.
Quin, D. G. (1992). Observations of prey detection by the bandicoots, Isoodon
obesulus and Perameles gunnii (Marsupialia: Peramelidae). Australian
Mammalogy 15, 131–133.
422 Australian Journal of Zoology L. E. Valentine et al.
Thackway, R., and Cresswell, I. D. (1995). ‘An Interim Biogeographic
Regionalisation of Australia: a Framework for Establishing the National
System of Reserves.’(Australian Nature Conservation Agency:
Canberra.)
Van Dyck, S., and Strahan, R. (2008). ‘The Mammals of Australia.’3rd edn.
(Reed New Holland: Sydney.)
Whitford, W. G. (1999). Biopedturbation by mammals in deserts: a review.
Journal of Arid Environments 41, 203–230. doi:10.1006/jare.1998.0482
Wilson, B. A., Valentine, L. E., Reaveley, A., Isaac, J., and Wolfe, K. M.
(2012). Terrestrial mammals of the Gnangara Groundwater System,
Western Australia: history, status and the possible impacts of a drying
climate. Australian Mammalogy 34, 202–216. doi:10.1071/AM11040
Handling Editor: Steven Cooper
Soil turnover by the southern brown bandicoot Australian Journal of Zoology 423
www.publish.csiro.au/journals/ajz