Wombat burrows are hotspots for small vertebrates in a landscape subject to gigafire

Abstract

Ecosystem engineers modify their environment and influence the availability of resources for other organisms. Burrowing species, a subset of allogenic engineers, are gaining recognition as ecological facilitators. Burrows created by these species provide habitat for a diverse array of other organisms. Following disturbances, burrows could also serve as ecological refuges, thereby enhancing ecological resistance to disturbance events. We explored the ecological role of Common Wombat (Vombatus ursinus) burrows using camera traps in forests of southeastern Australia. We compared animal activity at paired sites with and without burrows, from the same fire severity class and habitat. We examined how animal activity at Common Wombat burrows was affected by the 2019–20 Black Summer bushfires in Australia. We predicted that burrows would serve as hotspots for animal activity and as refuges in burned areas. The activity of several species including Bush Rat (Rattus fuscipes), Agile Antechinus (Antechinus agilis), Lace Monitor (Varanus varius), Painted Button-quail (Turnix varius), and Grey Shrike-thrush (Colluricincla harmonica) increased at sites where Common Wombat burrows were present, while other species avoided burrows. Species that were more active at burrows tended to be smaller mammal and bird species that are vulnerable to predation, whereas species that avoided burrows tended to be larger mammals that might compete with Common Wombat for resources. Species composition differed between sites with and without burrows, and burrow sites had higher native mammal species richness. The association of several species with burrows persisted or strengthened in areas that burned during the 2019–20 Black Summer bushfires, suggesting that Common Wombat burrows may act as ecological refuges for animals following severe wildfire. Our findings have relevance for understanding how animals survive, persist, and recover following extreme wildfire events.

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Journal of Mammalogy, 2024, 105, 752–764
https://doi.org/10.1093/jmammal/gyae034
Advance access publication 16 May 2024
Research Article
© The Author(s) 2024. Published by Oxford University Press on behalf of the American Society of Mammalogists.
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Received: April 24, 2023; Editorial Decision: March 13, 2024; Accepted: March 20, 2024
Research Article
Wombat burrows are hotspots for small vertebrates
in a landscape subject to gigare
Grant D. Linley,1,2,*, William L. Geary,3,4,5, Chris J. Jolly,1,6, Emma E. Spencer,7, Kita R. Ashman,1,7, Damian R. Michael,1, Dylan
M. Westaway,1,2, and Dale G. Nimmo1,2,
1Gulbali Institute for Agriculture, Water and Environment, Charles Sturt University, Thurgoona, NSW 2640, Australia
2School of Agricultural, Environmental and Veterinary Sciences, Charles Sturt University, Thurgoona, NSW 2640, Australia
3Biodiversity Strategy and Planning Branch, Biodiversity Division, Department of Environment, Land, Water and Planning, East Melbourne, VIC 3002, Australia
4School of Life and Environmental Sciences, Deakin University, Burwood, VIC 3125, Australia
5School of Agriculture, Food and Ecosystem Sciences, University of Melbourne, Parkville, VIC 3010, Australia
6School of Natural Sciences, Macquarie University, Macquarie Park, NSW 2109, Australia
7WWF-Australia, Suite 3.01, Level 3/45 Clarence Street, Sydney, NSW 2000, Australia
*Corresponding author: Gulbali Institute for Agriculture, Water and Environment, Charles Sturt University, Thurgoona, NSW 2640, Australia. Email: grant.linley@gmail.com
Associate Editor was Aaron Greenville
Abstract
Ecosystem engineers modify their environment and inuence the availability of resources for other organisms. Burrowing species, a
subset of allogenic engineers, are gaining recognition as ecological facilitators. Burrows created by these species provide habitat for a
diverse array of other organisms. Following disturbances, burrows could also serve as ecological refuges, thereby enhancing ecological
resistance to disturbance events. We explored the ecological role of Common Wombat (Vombatus ursinus) burrows using camera traps
in forests of southeastern Australia. We compared animal activity at paired sites with and without burrows, from the same re severity
class and habitat. We examined how animal activity at Common Wombat burrows was affected by the 2019–20 Black Summer bushres
in Australia. We predicted that burrows would serve as hotspots for animal activity and as refuges in burned areas. The activity of several
species including Bush Rat (Rattus fuscipes), Agile Antechinus (Antechinus agilis), Lace Monitor (Varanus varius), Painted Button-quail (Turnix
varius), and Grey Shrike-thrush (Colluricincla harmonica) increased at sites where Common Wombat burrows were present, while other
species avoided burrows. Species that were more active at burrows tended to be smaller mammal and bird species that are vulnerable to
predation, whereas species that avoided burrows tended to be larger mammals that might compete with Common Wombat for resources.
Species composition differed between sites with and without burrows, and burrow sites had higher native mammal species richness. The
association of several species with burrows persisted or strengthened in areas that burned during the 2019–20 Black Summer bushres,
suggesting that Common Wombat burrows may act as ecological refuges for animals following severe wildre. Our ndings have rele-
vance for understanding how animals survive, persist, and recover following extreme wildre events.
Key words: allogenic engineer, Black Summer bushres, ecological facilitation, eucalypt forests, re refuge, re severity.
Ecosystem engineers modify their environment by inuencing
resources available to other organisms (Jones et al. 1997; Coggan
et al. 2018). Modication by ecosystem engineers occurs via 2
pathways: autogenic engineers, which alter the environment by
modifying themselves (e.g., trees and coral; Jones et al. 1994); and
allogenic engineers, which alter the environment by mechani-
cally changing the form of biotic and abiotic materials (e.g., dam-
building beavers; Jones et al. 1994). Ecosystem engineers are
distributed across terrestrial, freshwater, and marine environments
(Jones et al. 1994), and include species from an array of taxonomic
groups, including invertebrates, reptiles, birds, and mammals.
Species that displace soil through burrowing, digging, and for-
aging—known as bioturbators—play an important role as ecosys-
tem engineers and often provide resources for co-occurring species
(Whitford and Kay 1999; Coggan et al. 2018). Bioturbation modies
soil by changing water run-off and erosion (Halstead et al. 2020) and
increasing water inltration (Fleming et al. 2014; Davies et al. 2019),
which alters the chemical properties of soils (Guy and Kirkpatrick
2021) and can enhance seed germination (Eldridge and James 2009;
Fleming et al. 2014). For example, in Australia, the bioturbation
activity of Quenda (Isoodon fusciventer) increases soil nutrients and
microbial activity, promoting seedling growth (Valentine et al. 2018).
In addition, well-digging by feral equids in North American deserts
increases water availability and vertebrate richness (Lundgren et al.
2021). Such engineering activities can provide important resources
and enhance local biodiversity.
Burrow engineers are a subset of bioturbators that combine
bioturbation with the creation of often large and complex burrow

Journal of Mammalogy, 2024, Vol, 105, Issue 4 | 753
systems. Burrow creation is a facilitative interaction whereby com-
mensal species exploit resources offered from burrows, including
the provision of shelter from predators and extreme conditions,
as well as foraging and breeding opportunities. For instance,
American Badger (Taxidea taxus) burrows are exploited by 31 spe-
cies, including numerous mammal species that use the bur-
rows for shelter (Andersen et al. 2021). Similarly, Gopher Tortoise
(Gopherus polyphemus) burrows are used by over 60 vertebrate spe-
cies (Dziadzio and Smith 2016), and Giant Armadillo (Priodontes max-
imus) burrows are visited by over 50 vertebrate species (Desbiez and
Kluyber 2013). In northern Australia, deep nesting warrens created
by Sand Goanna (Varanus gouldii) provide shelter for at least 28 ver-
tebrate species (Doody et al. 2021). Hence, the habitat created by
burrow engineers can result in hotspots of animal activity through
ecological facilitation.
Refuges are features that facilitate survival or persistence of spe-
cies during and following disturbance (Keppel et al. 2012; Reside et
al. 2019). There is some evidence that burrow engineers can create
refuges for other species: for instance, burrows of Southern Hairy-
nosed Wombat (Lasiorhinus latifrons) and Desert Tortoise (Gepherus
agassizzi) are used by birds to escape extreme heat (Attwood 1982;
Walde et al. 2009). The increased use of burrows by commensal spe-
cies during disturbance events is consistent with the stress-gradi-
ent hypothesis, which predicts that facilitative interactions between
species are most important in harsh ecological conditions (Bertness
and Callaway 1994; Lowney and Thomson 2021).
Fire is a global driver of environmental change (Pausas and
Keeley 2021) and has shaped the evolution of species over millions
of years (Pausas and Parr 2018; Nimmo et al. 2021). Retreating to
“re refuges”—features that allow for the survival, persistence,
and reestablishment of populations during and following re
(Robinson et al. 2013)—is 1 strategy that animals deploy dur-
ing and after re (Nimmo et al. 2019; Jolly et al. 2022). The occur-
rence of refuges created by ecosystem engineers has rarely been
considered within this context (Dawson et al. 2019; Reside et al.
2019), but could theoretically enhance the resistance of wild-
life to a range of disturbances, while resources provided by these
engineers could hasten recovery. Knapp et al. (2018) provides
some support for the importance of burrow engineers in re-
prone landscapes, showing that vertebrate use of Gopher Tortoise
(G. polyphemus) burrows increased >8-fold at sites that experienced
prescribed burns compared to unburned sites. Given that the fre-
quency of large, severe wildres is predicted to increase as a result
of climatic change (Wu et al. 2021), re refuges created by ecosys-
tem engineers could play a particularly critical role in the future.
The 2019–20 Australian Black Summer bushres were a series of
megares and gigares (res >10,000 and 100,000 ha, respectively;
Linley et al. 2022) that were unprecedented in their scale (Nolan
et al. 2020) and severity (Collins et al. 2021). These res burnt >10
million ha of southeastern Australian forests (Wintle et al. 2020),
including a record amount (1.8 million ha) burnt at high severity
(Collins et al. 2021), impacting nearly 3 billion native vertebrate ani-
mals (van Eeden et al. 2020). During these bushres, viral stories
emerged on social media of wombats herding native wildlife into
their burrows to protect them from re. While these stories were
ultimately dismissed (Nimmo 2020), an element of truth could still
be gleaned from them.
Wombats (family Vombatidae) are the largest burrowing marsu-
pials on earth (Evans et al. 2006). There are 3 extant wombat species,
all of which occur in Australia. The Common Wombat (Vombatus
ursinus; Fig. 1), also known as a Bare-nosed Wombat, has the larg-
est distribution of the 3 species (Taylor 1993; Evans 2008; Baker
and Gynther 2023), which coincides with several of the 2019–20
bushres (Fig. 2). Despite being listed as Least Concern by the IUCN
(2008), the species has undergone substantial decline across its
range since European colonization (Roger et al. 2007), with climate
change predicted to cause further range contractions (Graham et
al. 2019). Common Wombat are active in forests after re (Lunney
and O’Connell 1988), thus having an ability to maintain burrows
in burnt habitat. Wombats excavate and use multiple burrows
(Thornett et al. 2017), which consist of a large network of under-
ground tunnels (Swinbourne et al. 2016b). Common Wombat bur-
rows can be upwards of 15 m long, with multiple entrances—and
can be deep, complex, and multichambered (Triggs 2009; Browne et
al. 2021). Soil turnover from burrow construction can vary from 2.8
to 9.8 t/ha (Triggs 2009) and the soil in mounds contain increased
nitrogen, which supports higher herb cover (Guy and Kirkpatrick
2021).
Wombat burrows are exploited by a range of species: a study
showed that Common Wombat burrows were visited by 11 other
species including possums, rodents, and birds (Old et al. 2018).
Similarly, Thornett et al. (2017) recorded 11 vertebrate species that
used burrows of Southern Hairy-nosed Wombat. The internal tem-
peratures of wombat burrows are typically cooler and more stable
than aboveground temperatures (Finlayson et al. 2003; Evans 2008;
Swinbourne et al. 2016b), which can allow them to act as thermal
refuges during heatwaves (Attwood 1982). Fossorial South-eastern
Slider (Lerista bougainvillii) have also been observed exploiting the
soil mounds around burrows, suggesting that burrows might offer
increased thermoregulatory opportunities (Hodgson and Ritchie
2023). This prompts the question: could Common Wombat burrows
also provide refuge to animals from the impacts of re?
In this study, we monitored the activity (i.e., the number of detec-
tion events separated by >30 min) of a range of native vertebrates
around Common Wombat burrows in habitats that burned at vary-
ing severity during the 2019–20 bushres, and at nearby unburned
sites. We hypothesized that: (i) if Common Wombat are ecological
facilitators via their bioturbation, then animal activity will be higher
around burrows compared to nearby sites without burrows—the
afnity of species with burrows will differ depending on their shel-
ter and resource requirements, and hence community composition
will differ between burrows and ecologically similar areas lacking
burrows; and (ii) if Common Wombat burrows are acting as re ref-
uges, then the negative impacts of re on animal activity—which
we expect to be widespread given the severity of the 2019–20 bush-
res—will be reduced at burrows located in burned habitat, and
Fig. 1. An adult and juvenile Common Wombat (Vombatus ursinus).

754 | Linley et al.
these buffering effects should be most apparent at sites that burned
at high severity, consistent with the stress-gradient hypothesis.
Materials and methods.
Study area.
We conducted this study at Woomargama National Park (24,185
ha) and Woomargama State Forest (7,120 ha), New South Wales,
Australia. Geology consists predominantly of steeply sloping
Silurian Koetong granite, with soils consisting of yellow to red
podsolics (NSW National Parks and Wildlife Service 2009). The
average annual rainfall is 886 mm, with the majority of rainfall
occurring in winter and spring (Bureau of Meteorology 2023). The
area consists of continuous eucalypt forest and woodland com-
munities. Dominant canopy species consists of a mix of Eucalyptus
spp., primarily Broad-leaved Peppermint (E. dives), Narrow-leaved
Peppermint (E. radiata), and Brittle Gum (E. mannifera; NSW National
Parks and Wildlife Service 2009). Both mid and ground stories
consist of a mix of shrubs and grasses (NSW National Parks and
Wildlife Service 2009). The area has undergone historical land
use changes including mining, forestry, and agricultural activities
(NSW National Parks and Wildlife Service 2009). During the 2019–20
bushres, the Green Valley/Tunnel Road re complex was part of a
larger gigare (Linley et al. 2022) in which 6 res merged and burned
632,315 ha. The re burned ~18,119 ha of Woomargama National
Park and Woomargama State Forest at mixed severities, occurred
predominantly on the eastern side of the park, started by lightning
strike on 29 December 2019, and was extinguished by 18 February
2020. Fires in these environments can have long-lasting impacts on
ecosystems (Bradstock 2008; Williams et al. 2008).
Site selection.
We used a paired design to assess the use of Common Wombat
burrows in differing re severity treatments, in comparison to
nearby areas with the same re severity but without a burrow.
Common Wombat burrows were selected from ground searching
the study area in conjunction with re severity maps. Ground-
truthing revealed the extreme impact of the res on vegetation
throughout the study area, with many large trees killed and under-
story regrowth and canopy resprouting occurring postre (Fig. 2).
Using the Australian Google Earth Engine Burnt Area Map of the
2019–20 bushres (Aus GEEBAM; Department of Agriculture 2020)
re severity maps were used to classify the change in vegetation.
Aus GEEBAM re classes were derived from changes in a vegeta-
tion index following re, in comparison to nearby ecologically sim-
ilar areas that escaped the re (Department of Agriculture 2020).
Aus GEEBAM re classes are: unburnt, where little or no change
in the vegetation index was observed; low and moderate severity,
where moderate change to reference unburnt area was detected;
high severity, where the vegetation was mostly scorched; and very
high severity, where vegetation was consumed (Department of
Agriculture 2020).
In total, 28 site pairs (i.e., 56 sites in total) were selected. Site
pairs consisted of a Common Wombat burrow (Fig. 2) and a paired
site approximately 50 m away in the same re severity class and
in similar habitat (i.e., vegetation type, topography), to act as an
Fig. 2. (A) The Australian distribution of the Common Wombat (Vombatus ursinus; dark gray; IUCN 2008) in relation to the areas burned in the 2019–20
Black Summer bushres (red; Department of Agriculture 2020), and the study location (black); (B) a Common Wombat burrow; (C) the early stages of
postre regrowth; and (D) landscape impacts of the 2019–20 Black Summer bushres at Woomargama National Park, New South Wales, Australia (images
in C and D courtesy of Kylie Durant).

Journal of Mammalogy, 2024, Vol, 105, Issue 4 | 755
experimental control (Coggan et al. 2018). Fire severity classes
were ground-truthed to ensure the classication from maps
matched the burn severities on the ground and canopy layers.
We reclassied the Aus GEEBAM re severity classes listed above
to include: unburnt areas outside of the re scar; unburnt areas
within the re scar; low and moderate severities; and high sever-
ities that encompasses both high and very high burned areas,
as differences between these 2 classes were not obvious on the
ground. Seven sites were established within each of the re sever-
ity classes. Preliminary analysis suggested few differences in ani-
mal activity in unburned areas outside of and within the re scar,
and so all unburned sites were classied simply as unburned. All
sites were located within Eucalypt woodlands with shrubby or
grassy understory. Vegetation at sites within the re scar exhibited
signicant regrowth, manifesting at various stages depending on
the re severity at each location.
Animal activity.
We surveyed the activity of terrestrial mammals at each site using
wildlife cameras (Reconyx HC600 Hyperre, Reconyx Inc., Holmen,
Wisconsin). Each of the 28 site pairs consisted of 2 cameras (56
sites), 1 placed at a Common Wombat burrow and the other at the
paired control site. To capture smaller mammals, cameras were fas-
tened to a stake at a 20° downward angle. At burrows, cameras were
placed approximately 2 m from the burrow entrance, facing the
entrance. Camera orientation was consistent between burrows and
control sites, with the only difference being that control sites were
oriented toward bare ground. Cameras operated day and night,
recording 5 images per burst, with no time delay between bursts,
and sensitivity set to high. Cameras were deployed 16 months after
the 2019–20 bushres and operated continuously between June 2021
and April 2022, totaling 16,645 trap-days. Our study followed guide-
lines (Sikes et al. 2016) and all applicable international, national,
and/or institutional guidelines for the use of animals. Research was
conducted in accordance with the Charles Sturt University Animal
Ethics Committee (permit number A21049) and permissions from
relevant management authorities.
Image tagging and data extraction.
Images from 56 cameras were tagged and processed using Wildlife
Insights (Ahumada et al. 2020). When present, animals were iden-
tied to species level, if species-level identication was not fea-
sible, identication was made to the highest possible taxonomic
resolution (typically genus level). This was achieved using Wildlife
Insights’ articial intelligence, which automatically detects and
identies species in images (Ahumada et al. 2020), although every
image and species identication was independently reviewed
by wildlife experts. Behaviors evident in images were manually
recorded where possible, including number of individuals and
the occurrence of specic behaviors around burrows includ-
ing: bathing, referring to an animal using water in the burrow
to clean themselves; drinking, referring to an animal consuming
water from the burrow; entering or emerging, referring to an ani-
mal entering or exiting the burrow entirely; inspecting, referring
to an animal investigating the burrow entrance but not entering
the burrow completely; foraging, referring to animals feeding in
or directly around the lip of the burrow; and geophagy, referring
to animals consuming soil from the burrow. Classifying behaviors
was not always possible due to the small size of some species—e.g.,
House Mouse (Mus musculus) and Agile Antechinus (Antechinus agi-
lis), making it difcult to assign a specic behavior to some images.
We also recorded dates when burrows were lled with water to
assess how often Common Wombat burrows provide a water
source to other wildlife. We extracted the tagged image metadata,
including site information, date, and time using camtrapRdelux
package (https://github.com/carlopacioni/camtrapRdeluxe, an
extension of camtrapR (Niedballa et al. 2016) and the exiftool add
on (Harvey 2020). We treated all detections as independent events
when more than 30 min separated detections of the same species
(Cunningham et al. 2018).
Data analysis.
To test our hypothesis that animal activity will be higher around
burrows, we t Bayesian mixed-effects models using the package
brms (Bürkner 2017), which ts Bayesian models in stan which
compiles models to sample from posterior distributions (Stan
Development Team 2022). We included site pairs as random effects
and used 2,000 iterations and 4 chains, with a 2,000-iteration
warm-up. We used default priors, which are weakly informative,
to improve convergence and sampling efciency, with minimal
impact on estimations. Model parameters are regarded as show-
ing evidence of an effect when the 89% credible interval did not
overlap zero (McElreath 2020). We used 89% credible intervals
as they are commonly used in Bayesian statistics (Samaš et al.
2021), and due to their relative stability compared to 95% cred-
ible intervals (Kruschke 2014). Our response variables were: the
number of independent events for each species; species richness
(i.e., count of all species detected at a site) of all species (total
richness); native mammal species only; and native bird species
only. Individual species were modeled if recorded by at least 10
camera traps.
We modeled our response variables as a function of whether the
site was a burrow or a control (“site type”). Control sites were used
as the reference category. If the 89% credible intervals of the mixed
models did not overlap zero, it was considered evidence that the
response variable at burrows was substantially different from the
control sites (McElreath 2020). To test for compositional differences
between species assemblages at burrows and control sites, we used
a permutational multivariate analysis of variance (PerMANOVA).
We used the Bray–Curtis index to calculate dissimilarities based on
a square root transformation of our activity data. PerMANOVA was
t using the adonis2 function from the vegan package (Oksanen et al.
2013). We permuted dissimilarities 999 times to assess signicance.
Community composition was visualized using nonmetric multidi-
mensional scaling (nMDS: Legendre and Legendre 1998). To deter-
mine which species most inuenced dissimilarities at burrows and
control sites, we performed indicator species analysis using the mul-
tipatt function from the indicspecies package (De Cáceres et al. 2016).
Finally, to examine if associations were most pronounced in
areas that experienced high-severity res, we again used Bayesian
models in brms (Bürkner 2017) to model response variables as a
function of site type and re severity class. We combined site type
and re severity to create a 6-level categorical variable (unburned
burrow, unburned control, low–moderate burned burrow, low–
moderate burned control, high severity burrow, high severity
control) and used unburned controls as the reference category.
Because some categories had few detections for some species, we
used weakly informative priors with a Student’s t-distribution to
aid in model convergence. Specifying unburned controls as the
reference allows both a comparison between burrows and con-
trol sites, which reveal whether burrows experience increased or
decreased activity relative to controls—as well as a comparison
between unburned and burned controls, which reveal the impacts
of re severity on each response variable. This allows the impact
of burrows to be contextualized in terms of the background inu-
ence of re.

756 | Linley et al.
Results
Camera traps recorded 746,674 images containing 370,845 wildlife
images. Images were compressed into 14,580 events that repre-
sented 15,723 individual animals. Fifty-six species were identied
(47 native species, 9 introduced), consisting of 19 mammal, 33
bird, and 4 reptile species. Common Wombat were present at all
sampled burrows, which had an average burrow opening height
of 19.18 cm (±2.77 SE) and width of 22.66 cm (±3.21 SE). Excluding
Common Wombat, 48 species were identied at burrows and 43
at control sites (Supplementary Data SD1). Species found only at
burrows included Common Ringtail Possum (Pseudocheirus peregri-
nus), Grey Fantail (Rhipidura albiscapa), Spotted Pardalote (Pardalotus
punctatus), and White-throated Treecreeper (Cormobates leuco-
phaea). Species only observed at control sites included Australian
Raven (Corvus coronoides), Common Bronzewing (Phaps chalcoptera),
and Eastern Rosella (Platycercus eximius). A total of 31 species were
recorded interacting with burrows (Supplementary Data SD2),
including several species inspecting (30 of 31 species), foraging
(11/31), and entering and emerging (10/31) from burrows—while
less common behaviors included drinking (4/31) and bathing (1/31)
in ooded burrows, and geophagy (1/31; Supplementary Data SD2;
Fig. 3). From 16,645 trap-days, images containing animals occurred
over 8,384 trap-days, of which burrows contained pooled water for
207 days. From 28 burrows, 19 lled with water at least once for an
average total time of 10.89 days (±3.74 SE). From 207 days where
burrows were lled with water, 109 days occurred in spring, 54 in
winter, and 44 in summer.
Several species showed increased activity around burrows
when compared to control sites (i.e., 89% CIs did not overlap
zero; Fig. 4; Supplementary Data SD3), including several native
species such as Bush Rat (Rattus fuscipes), Agile Antechinus, Lace
Monitor (Varanus varius), Painted Button-quail (Turnix varius), Grey
Shrike-thrush (Colluricincla harmonica), and introduced Black Rat
(Rattus rattus; Fig. 4; Supplementary Data SD3). Several native
species including Swamp Wallaby, Red-necked Wallaby, Eastern
Grey Kangaroo, Australian Magpie (Gymnorhina tibicen), Satin
Bowerbird (Ptilonorhynchus violaceus), Crimson Rosella (Platycercus
elegans), White-winged Chough (Corcorax melanorhamphos), and
introduced Feral Pig (Sus scrofa) were more active at control sites
(Fig. 4; Supplementary Data SD3). The activity of all other spe-
cies did not differ substantially between burrows and control
sites. Native mammal richness was greater at burrows than con-
trol sites (Fig. 4; Supplementary Data SD3), whereas total rich-
ness and bird richness did not substantially differ (i.e., 89% CIs
overlapped zero). For mammals, there was a tendency for smaller
species to be positively associated with burrows and for larger
species to be negatively associated with burrows (Fig. 5). A linear
regression comparing the effect size from Bayesian mixed-effects
models to the natural logarithm of body mass revealed a strong,
negative relationship for mammals (Coefcient ± 89% CI = −0.494
[−0.638 to −0.350], t = −6.026, R2 = 0.784). This relationship was far
weaker when considering birds and reptiles in addition to mam-
mals (Coefcient ± 89% CI = −0.295 [−0.502 to −0.089], t = −0.621,
R2 = 0.052), suggesting that this size-dependent relationship with
burrows was specic to mammals.
Species composition varied between burrow and control sites
(PerMANOVA, F1,55 = 2.188, P = 0.003; Fig. 6). Indicator species analy-
sis identied several species associated with burrows: Bush Rat (stat
= 0.849, P = 0.001); Agile Antechinus (stat = 0.773, P = 0.001); Lace
Monitor (stat = 0.602, P = 0.003); White-throated Treecreeper (stat
= 0.463, P = 0.023); and Yellow-footed Antechinus (Antechinus avi-
pes; stat = 0.463, P = 0.026). Indicator species for control sites were
the Red-necked Wallaby (stat = 0.799, P = 0.022) and Feral Pig (stat =
0.639, P = 0.037; Fig. 6).
The association between Agile Antechinus and burrows was
most apparent at burrows burned at high severity, remained
evident at burrows burned at low–moderate severity, and was
least evident (89% CIs overlapping zero) at unburned burrows
(Fig. 7; Supplementary Data SD4). Bush Rat were more active
at burrows burned at high severity, low–moderate severity, and
unburned burrows. Painted Button-quail were more active around
burrows that burned at high severity compared to unburned
control sites (Fig. 7; Supplementary Data SD4). The association
of Lace Monitor with burrows was only evident in unburned
sites (Fig. 7; Supplementary Data SD4). Feral Pig, Spotted Quail-
thrush, Common Brushtail Possum, and Australian Magpie were
Fig. 3. Examples of animals and behaviors observed at Common Wombat burrows: (A) a Lace Monitor foraging in the burrow entrance; (B) a Short-beaked
Echidna inspecting the lip of the burrow; (C) a Swamp Wallaby drinking from a burrow full of water; (D) a Red-necked Wallaby inspecting the burrow
entrance; (E) a Grey Shrike-thrush foraging at the burrow entrance; and (F) a Pied Currawong drinking from a partially lled burrow.

Journal of Mammalogy, 2024, Vol, 105, Issue 4 | 757
all less active around unburnt burrows (Fig. 7; Supplementary
Data SD4). Native mammal richness was higher at burrows that
were either burned at high severity or remained unburnt (Fig. 7;
Supplementary Data SD4). Swamp Wallaby, Feral Pig, and feral
Domestic Cat (Felis catus) were the only species for which activity
varied depending on re severity, all being more active in areas
that burned at high severity during the 2019–20 bushres (Fig. 7;
Supplementary Data SD4).
Discussion
Ecological facilitation by Common Wombat increases the activity
of numerous native species, alters community composition, and
boosts native mammal richness. Body size determined whether
mammals were positively or negatively associated with burrows,
with smaller species being more active and larger species being
less active around burrows. The association of several species with
burrows persisted—and in some instances was strengthened—after
severe wildre, consistent with the stress-gradient hypothesis. Our
results add to a growing literature underscoring the signicance of
burrowing engineers as ecological facilitators.
Wombat burrows affect animal community
composition.
The presence of Common Wombat burrows alters the local envi-
ronment sufciently to elicit clear changes in local animal com-
munities. Of particular interest is an apparent size dependency
in species interactions with burrows, with smaller mammals
being more active and larger mammals less active at burrows. The
capacity of a species to enter Common Wombat burrows appears
to be linked to the size of the species, with larger animals being
excluded from utilizing the available shelter and foraging poten-
tial due to the size of the burrow openings. We detected numer-
ous examples of smaller species using Common Wombat burrows,
including Bush Rat, Black Rat, Short-beaked Echidna, Grey Shrike-
thrush, White-throated Treecreeper, and Lace Monitor. Our survey
method may have even underestimated the activity of some of
these species, as camera traps are often less effective at captur-
ing smaller species of mammals, birds, and reptiles (Jumeau et al.
2017). Other researchers have documented small mammals enter-
ing and exiting burrows: Triggs (2009:33) mentions antechinus
and bush rats seen “scurrying away from burrow entrances,” that
antechinus “emerge from small crevices in the walls of Bare-nosed
Fig. 4. Coefcients and credible intervals from Bayesian models comparing species activity in burrows versus control sites (controls specied as the
reference category), and examples of species activity predicted from those models. In activity plots: colored dot = point estimate; thick bars = 66% CIs; and
thin bars = 89% CIs.

758 | Linley et al.
Wombat burrows,” and that goannas “have also been found in
wombat burrows,” while McIlroy (1973) found Bush Rat in Common
Wombat burrows. One of our authors (DRM) has observed ante-
chinus and bush rats escaping to Common Wombat burrows not
far from our study region. Even relatively larger species, like the
Swamp Wallaby and Red-necked Wallaby, were observed entering
burrows, although these were rare events (relative to other behav-
iors). Common Wombat are large-bodied and stocky mammals
with occasional displays of aggression, and they are known to
compete with macropods for resources (Tamura et al. 2021). Thus,
it is plausible that Common Wombat actively deter larger compet-
itors away from their burrows, or that macropods avoid burrows to
reduce the risk of a negative encounter, as Common Wombat vig-
orously defend their territories against unwanted intruders (Triggs
2009).
Aside from providing shelter, Common Wombat burrows could
affect animal communities via changes in soils, vegetation, and
topography around the burrow. Soil around Common Wombat
burrows has greater nitrogen than surrounding soil due to the
deposition of wombat feces, while vegetation in the vicinity of bur-
rows typically includes more bare ground and herbaceous plants,
with less shrub cover (Guy and Kirkpatrick 2021). These variables
could affect the foraging opportunities for small herbivores and car-
nivores alike. Common Wombat burrows may increase the activity
of ground-dwelling insectivores and omnivores such as Bush Rat,
Agile Antechinus, Grey Shrike-thrush, and Painted Button-quail, by
providing increased foraging opportunities. While little is known of
invertebrate communities present at Common Wombat burrows
(and was not a specic subject of our research here), high diver-
sities of invertebrates have been recorded at the burrows of other
species. For instance, Gopher Tortoise burrows support hundreds of
arthropod species (Jackson and Milstrey 1989), while burrows and
mounds from Gunnison’s Prairie Dog (Cynomys gunnisoni), Black-
tailed Prairie Dog (C. ludovicianus), and Banner-tailed Kangaroo Rat
(Dipodomys spectabilis) increase arthropod abundance and richness
compared to surrounding areas without mounds and burrows
(Davidson and Lightfoot 2007). Additionally, native Australian pred-
ators including Western Quoll (Dasyurus geoffroii), Sand Goanna
Fig. 5. Effect size from Bayesian mixed-effects models of treatment (burrow vs. control) against the natural logarithm of body mass for mammals at
Woomargama National Park: black line = tted relationship from linear regression model; black dots = coefcients ± 89% credible intervals; effect sizes > 0
indicate a positive association with burrows, and effect sizes < 0 indicate negative associations.

Journal of Mammalogy, 2024, Vol, 105, Issue 4 | 759
(V. gouldii), and Mulga Snake (Pseudechis australis) use burrows from
European Rabbit (Oryctolagus cuniculus) that concentrated prey spe-
cies (Dean et al. 2023). The increase in activity of small vertebrates
around Common Wombat burrows may cascade through the sys-
tem, as these species attract and are exploited by larger native
predators such as Lace Monitor. These large, generalist, ectother-
mic carnivores may also derive additional benets from Common
Wombat burrows, because burrows provide a thermal refuge, buff-
ering against extreme environmental conditions (Pike and Mitchell
2013).
Finally, we observed several instances of Common Wombat bur-
rows lling with water including during summer, signaling that
water provision may be another ecological role that they provide.
Macropods including Eastern Grey Kangaroo, Red-necked Wallaby,
and Swamp Wallaby were all captured drinking from burrows, and
Pied Currawong were observed bathing. These observations sug-
gest that burrows could provide an important water resource to
animals, although they may only provide critical services during
periods of low water availability. For example, well-digging by feral
equids in North American deserts increases water availability and,
probably due to the scarcity of water in this system, vertebrate
richness (Lundgren et al. 2021). While water was not scarce during
our study period, or compared to other drier ecosystems, it does
play a vital role in regulating and maintaining ecosystem services,
especially in the drier months (Pepper et al. 2008). As such, water
provision via Common Wombat burrows, especially during warmer
periods, presents an interesting observation that warrants further
investigation.
Wombat burrows as re refuges.
We found that associations between species and burrows
persisted, and often strengthened in re-impacted habitat,
particularly in areas that burned at high severity. Smaller-sized
animals including Agile Antechinus, Bush Rat, and Painted Button-
quail all retained positive associations with burrows in areas that
were subject to high-severity re, and all 3 were most active at
burrows subject to high-severity re. By contrast, the association
of Lace Monitor with burrows was only evident at unburned bur-
rows. These ndings suggest that the association of species with
Common Wombat burrows is highly context-dependent, and that
re plays a signicant role in determining the nature of this rela-
tionship in Australian forests.
For small vertebrates that are vulnerable to predators, many
ammable refuges such as logs are consumed by severe res
(Bassett et al. 2015). Wombat burrows—which survive re and can
persist for decades (Taggart and Temple-Smith 2008)—potentially
provide important postre shelter. Our study suggested that the
2019–20 bushres had relatively negligible impacts on the activity
of species that were relatively similar between burnt and unburnt
habitat. However, it did indicate that the impacts of re might
concentrate the activity of small vertebrates within the vicinity of
burrows, providing a locus from which foraging and intraspecic
interactions can occur in relative proximity to predictable shel-
ter and protection from predators. Disturbances in other regions
likely shape associations between other ecosystem engineers and
the co-occurring species. Allospecic use of Gopher Tortoise bur-
rows dramatically increases during and after a re (Knapp et al.
2018), and European Rabbit warrens support greater densities of
Mediterranean lizards in unfavorable habitat (Gálvez Bravo et al.
2009), further highlighting the context-dependent importance of
burrows in disturbed landscapes.
Our results have important implications for conservation. The
geographic range of Common Wombat has declined since European
colonization of Australia (Buchan and Goldney 1998; Triggs 2009;
Fig. 6. The difference in species composition at burrows and control sites in Woomargama National Park. Ordinations show nMDS plots of square root
transformed activity indices: (A) sites (points) colored by treatment (burrow vs. control)—distances between sites indicate community dissimilarity, and
shaded areas are ellipses based on a multivariate t-distribution; and (B) the location of species in ordination space with indicator species highlighted in
bold.

760 | Linley et al.
Baker and Gynther 2023), and is predicted to substantially contract
from climate change (Graham et al. 2019). The 2 other species of
wombat are vulnerable to extinction (Swinbourne et al. 2016a), and
numerous other burrowing Australian marsupials have been driven
to extinction since European colonization (Woinarski et al. 2019).
Throughout the forests of eastern Australia, Common Wombat are
the only extant species of native mammal capable of excavating
wide, deep, and elaborate burrows, and our research suggests that
they play a critical role for a host of other species. The importance of
ecosystem engineers for providing critical shelter for other species
highlights the often-overlooked risk of co-extinctions due to the loss
of burrowers. For example, endangered pygmy Blue-tongued Skink
(Tiliqua adelaidensis)—presumed extinct for 33 years (Armstrong and
Reid 1992; Armstrong et al. 1993)—are entirely reliant on burrows
constructed by lycosid and mygalomorph spiders (Hutchinson et
al. 1994). Changed land use has caused declines in these ecosystem
engineering spiders and the near extinction of the lizards (Souter et
al. 2007). The value of burrowers as providers of shelter and refuge
likely increases after disturbance across a range of ecosystems. The
occurrence of larger and more severe wildres around the world has
placed a premium on understanding how animals survive, persist,
and recover following extreme wildre events. Our research has
revealed that Common Wombat burrows play a valuable and under-
appreciated role in the re-prone forests of Australia.
Supplementary data
Supplementary data are available at Journal of Mammalogy online.
Fig. 7. Coefcients and credible intervals from Bayesian models comparing species activity in burrows versus control sites in sites that vary in re severity
(controls specied as the reference category), and examples of species activity predicted from those models. In activity plots: colored dot = point estimate;
thick bars = 66% CIs; and thin bars = 89% CIs.

Journal of Mammalogy, 2024, Vol, 105, Issue 4 | 761
Supplementary Data SD1.—Species which were identied at
burrows and controls.
Supplementary Data SD2.—Number of images per behavior
observed from species interacting with wombat burrows.
Supplementary Data SD3.—Results of coefcients, with 89%
condence intervals, from Bayesian models comparing species
activity in burrows versus controls.
Supplementary Data SD4.—Results of coefcients, with 89%
condence intervals, from Bayesian models comparing species
activity in burrows versus controls in sites that vary in re severity.
Acknowledgments
We acknowledge the Wiradjuri People, the Traditional Owners of
the land upon which this study took place, and we pay respects to
Elders both past, present, and emerging. Dave Pearce from NSW
National Parks and Wildlife Service provided access and insights
into Woomargama National Park. Kylie Durant from Holbrook
Landcare Group provided information and insights into the
regional area. Kylee Imlach provided administrative and logisti-
cal support. Caroline Maurel and Mitch Cowan assisted with eld-
work. Alana de Laive helped with image identication. The Gulbali
Institute’s GAPS funding was instrumental in the completion of
this manuscript.
Author contributions
GDL helped conceptualize the study, collect data, curate the data
set, wrote the original draft, and led the analysis; DGN concep-
tualized the study, provided analysis assistance, and assisted
on the original draft; WLG played a pivotal role in the analysis
of the data and editing and revising the manuscript; CJJ assisted
with the analysis and played a signicant role in editing and revis-
ing the manuscript; EES, KA, DRM, and DMW helped edit and revise
the manuscript.
Funding
Funding for this project was provided by the WWF-Australia’s
Regenerate Australia scheme and Charles Sturt University’s Gulbali
Institute.
Conict of interest
None declared.
Data availability
Data are available upon reasonable request.
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