Malleefowl Leipoa ocellata incubation mounds as habitat for other vertebrates
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Australian Field Ornithology 2021, 38, 99–106
http://dx.doi.org/10.20938/afo38099106
Malleefowl Leipoa ocellata incubation mounds as habitat
for other vertebrates
Heather Neilly1* , David E. Wells2, Tim Pascoe3, Craig Gillespie4 and Peter Cale1
1
Australian Landscape Trust, P.O. Box 955, Renmark SA 5341, Australia
2
North Calperum Volunteer Group, 14 Schaefer Drive, Loxton SA 5333, Australia
3
BirdLife Australia Gluepot Reserve, via Lunn Road, Waikerie SA 5330, Australia
4
Murraylands and Riverland Landscape Board, 110A Mannum Road, Murray Bridge SA 5253, Australia
*Corresponding author. Email: heathern@alt.org.au
Abstract. Ecosystem engineers change the availability of resources for other species by forming new habitat or modifying
existing habitat but, despite the diversity of avian ecosystem engineers, 80% of current literature focuses on mammals and
invertebrates. Malleefowl Leipoa ocellata build large incubation mounds of soil and leaf-litter that are likely to provide habitat
for invertebrates and vertebrates but use of their mounds by other vertebrates has never been quantified. Here, we examine
vertebrate fauna visitation rates at Malleefowl mounds and non-mounds using camera-trap data collected by two citizen
science projects. From 2012 to 2018, 20 active Malleefowl mounds and 16 non-mound sites were monitored over 31,913 hours
and 225,144 hours, respectively. In total, we identified visits by 1724 birds, reptiles and mammals from 36 species. The mean
number of vertebrate visits per 1000 hours of surveillance was around one and a half times and species richness five times that
at mounds compared with non-mounds. Malleefowl mounds may enhance the availability of invertebrate prey for insectivorous
birds and mammals, provide a favourable microclimate for reptiles to thermoregulate, and be signalling/social communication
locations. Our results show that further research is warranted and suggest that conservation of Malleefowl may be important
not only for the Malleefowl itself, but also for a suite of mallee birds and reptiles.
Introduction engineers, which include burrowers, colonising seabirds,
hollow-excavators and mound-builders, have been largely
Ecosystem engineers change the availability of resources overlooked. Considering the ubiquity and diversity of birds,
for other species by forming new habitat or modifying however, research into their role as ecosystem engineers
existing habitat (Jones et al. 1994). Globally, ecosystem requires more attention.
engineering by fauna is a facilitative process, increasing Malleefowl Leipoa ocellata (Megapodiidae) are large,
local species’ diversity (Romero et al. 2015). Ecosystem ground-dwelling birds that build incubation mounds
engineers facilitate the provision of resources (e.g. food, of decomposing leaf-litter and sand scraped from the
water and thermal niches) to other faunal species, and surrounding mallee woodland. Megapode incubation
thus are disproportionately important taxa for conservation. mounds are the largest structure (when compared with
They enhance ecosystem function and may be a valuable body size) created by non-colonial animals (Jones & Göth
tool to assist with ecological restoration, particularly in 2008). Malleefowl mounds are ~4 m wide and 1.5 m high
depauperate, arid systems (Byers et al. 2006; Romero and contain ~3400 kg of material (Frith 1959; Weathers &
et al. 2015; McCullough Hennessy et al. 2016; Catterall Seymour 1998; Jones & Göth 2008). During the breeding
2018). season (September–February), when eggs are incubating
Despite the diversity of ecosystem engineers, 80% in the mound, Malleefowl spend up to 7 hours per day
of the literature focuses on mammals and invertebrates regulating the mound temperature (Weathers & Seymour
(Coggan et al. 2018). The impacts of burrowing mammals 1998; Neilly et al. 2021). Density of active Malleefowl
have been widely studied, with their burrows providing mounds ranges from 1.1 to 5.5 mounds per km2,
habitat for a range of reptiles, birds and invertebrates and increasing with greater annual rainfall (Frith 1962; Booth
other mammals (Read et al. 2008; Davidson et al. 2012; 1987). Post-breeding, inactive Malleefowl mounds persist
Catano & Stout 2015; Hofstede & Dziminski 2017; Coggan in the environment for decades (possibly much longer)
et al. 2018). Other vertebrate ecosystem engineers and can be four times more abundant than active mounds
create or modify habitat in myriad ways: damming rivers (Benshemesh et al. 2020; HN unpubl. data). Malleefowl
(e.g. beavers Castor spp.: Gurnell 1998), creating reefs reproduction necessitates significant soil disturbance,
(e.g. bivalve molluscs: Engel et al. 2017), excavating the movement and accumulation of resources, and
tree-hollows (e.g. woodpeckers of the Picinae: Cockle the creation of a large, raised structure in a landscape
et al. 2011), through foraging on the ground (e.g. lyrebirds devoid of any similar features. Presence of Malleefowl
Menura spp.: Webb & Whiting 2006), modifying vegetation mounds impacts wildfire fuel loads, and can influence
structure via browsing or grazing (e.g. bison Bison spp.: burning patterns at a local scale (Smith et al. 2016), and
Knapp et al. 1999) and by concentrated deposition of it is possible that mounds also influence soil quality and
guano (e.g. seabird colonies: Mosbech et al. 2018). The plant germination. In another megapode, the Australian
majority of faunal species probably act as ecosystem Brush-turkey Alectura lathami, an increase in abundance
engineers to some extent, although engineering effects has been shown to decrease ground-cover and leaf-litter,
range from minor to significant and are dependent and seed and seedling density in the surrounding area
on context (Wright & Jones 2006). Avian ecosystem (Warnken et al. 2004). Additionally, a range of fauna has100 Australian Field Ornithology H. Neilly et al.
been recorded visiting Australian Brush-turkey mounds: Surveillance of Malleefowl mounds
insectivorous birds feeding, mammals foraging, and
reptiles thermoregulating at mounds (Jones 1987). In From 2012 to 2018, annual monitoring of Malleefowl
contrast, very little is known about the impact of Malleefowl mounds was conducted by volunteers at Calperum Station
engineering activities, particularly the use of their mounds (Australian Landscape Trust and the North Calperum
by other vertebrates. Volunteer Group) and Gluepot Reserve (Friends of Gluepot
In this pilot study, we examined how Malleefowl may and BirdLife Australia). At active mounds, motion-sensor
act as ecosystem engineers by creating a novel habitat cameras were installed. The objective of this citizen science
(their incubation mounds) for other vertebrate fauna. project was to examine Malleefowl breeding ecology
We combined camera-trap data from two separate (Neilly et al. 2021) and capture camera footage for use
citizen science projects, to compare the visitation rates in engagement and education. Most camera surveillance
of vertebrates at mounds and away from mounds (non- of mounds was operational during the breeding season
mounds). We hypothesised that Malleefowl mounds would (September–February: see Neilly et al. 2021). This project
be visited at a higher rate and by a more diverse range of was initiated and carried out by volunteers, the equipment
vertebates than non-mound areas. set-up did not follow pre-defined protocols, and the timing
of activity was dictated by availability of volunteers.
Throughout the study, 20 active mounds were monitored:
Material and methods 14 from Calperum Station and six from Gluepot Reserve
(Table 1). Each camera (Little Acorn LTL-6210 or LTL-
Site location 6310) was positioned on a 1.5-m-high tripod ~2 m from
the edge of a mound and set to continuous recording
Calperum Station (238,638 ha, owned by Australian (0 second delay), but whether still or video footage was
Landscape Trust) and Gluepot Reserve (54,390 ha, being recorded varied between mounds.
owned by BirdLife Australia) are in the Riverland region of
South Australia (Figure 1). Both are ex-pastoral properties
destocked in the mid 1990s, and now managed for Non-mound surveillance
conservation. The average annual rainfall in this area is
256 mm but is highly variable (90–517 mm per annum) and Non-mound cameras (eight at Gluepot Reserve and
rainfall events can be unpredictable and irregular. Eucalypt eight at Calperum Station, ScoutGuard DTC-560K and
mallee communities (Red Mallee Eucalyptus socialis, Giant SG560K: Table 1) were set up in September 2016 as
Mallee E. oleosa, White Mallee E. dumosa and Yorrell part of the Australia-wide Adaptive Management Predator
E. gracilis) dominate the sand-dune system landscape. Experiment project (Hauser et al. 2019). This project
Table 1. The surveillance duration of motion-sensor cameras at 20 Malleefowl mounds and 16 non-mound sites at
Calperum Station and Gluepot Reserve, South Australia, 2012–2018.
Camera name Location Year Total Camera name Location Year Total
surveillance (h) surveillance (h)
Mounds sites Non-mound sites
CAL_2012a Calperum 2012–2013 287.8 CA Calperum 2016–2018 12,624
CAL_2012b Calperum 2012–2013 814.6 CB Calperum 2016–2018 16,224
CAL_2012c Calperum 2012–2013 646.7 CC Calperum 2016–2018 12,624
GLU_2013a Gluepot 2013–2014 4176.0 CD Calperum 2016–2018 15,096
GLU_2013b Gluepot 2013–2014 2452.6 CE Calperum 2016–2018 13,176
CAL_2013a Calperum 2013–2014 578.3 CF Calperum 2016–2018 14,280
CAL_2013b Calperum 2013–2014 1121.1 CG Calperum 2016–2018 12,432
GLU_2014 Gluepot 2014–2015 1824.0 CH Calperum 2016–2018 16,200
CAL_2014a Calperum 2014–2015 702.7 GA Gluepot 2016–2018 15,912
CAL_2014b Calperum 2014–2015 288.5 GB Gluepot 2016–2018 14,376
CAL_2014c Calperum 2014–2015 584.1 GC Gluepot 2016–2018 14,568
CAL_2015a Calperum 2015–2016 388.6 GD Gluepot 2016–2018 15,096
CAL_2015b Calperum 2015–2016 354.5 GE Gluepot 2016–2018 14,568
CAL_2015c Calperum 2015–2016 91.4 GF Gluepot 2016–2018 7656
GLU_2016 Gluepot 2016–2017 9312.0 GG Gluepot 2016–2018 13,320
CAL_2016a Calperum 2016–2017 341.6 GH Gluepot 2016–2018 16,992
CAL_2016b Calperum 2016–2017 1074.5
GLU_2017a Gluepot 2017–2018 1272.0
GLU_2017b Gluepot 2017–2018 2856.0
CAL_2017 Calperum 2017–2018 2746.6Malleefowl mounds as habitat for other vertebrates 101
photographs. To account for this difference, visits (and
their duration) were treated as continuous if consecutive
footage had a break of ≤10 minutes but were recorded
separately if >10 minutes.
Analysis of data
Visit data were divided by the number of surveillance
hours for each camera and multiplied by 1000, to give
the number of visits per 1000 hours of footage. Species
were grouped as reptiles; birds; Red Fox Vulpes vulpes;
Goat Capra hircus; Cat Felis catus; and other mammals
(European Hare Lepus europaeus, Short-beaked Echidna
Tachyglossus aculeatus, Common Brushtail Possum
Trichosurus vulpecula and Sheep Ovies aries).
For statistical analysis, a subset of the data was created
that included only observations from Calperum Station
cameras (to minimise the issue of spatial separation of
mound and non-mound sites, which was greater at Gluepot
Reserve: Figure 1) and within years 2016–2018 (when
non-mound and mound camera sites were simultaneously
in operation) to avoid the confounding influence of differing
rainfall, vegetation condition or management decisions
that might have impacted the number of vertebrates across
different years. Subset data were from three Malleefowl
Figure 1. Calperum Station and Gluepot Reserve mounds―CAL_2106a, CAL_2016b and CAL_2017―and
are adjacent properties in South Australia. The from the eight Calperum Station non-mound cameras.
locations of Malleefowl mound cameras and
A smaller dataset limited our statistical capabilities but
non-mound cameras are indicated on the map.
NSW = New South Wales, SA = South Australia. allowed us to assess whether the trends suggested by
the full dataset were an artefact of location or time. In the
analysis of subset data, values were calculated per camera
monitors Malleefowl productivity in areas under different site, and means with standard errors are presented.
feral-predator management regimes. The cameras
were arranged in an evenly spaced grid, predetermined To examine the difference in (1) total abundance and
using GPS, within a 2 km × 5 km area within annual richness by camera site and (2) the number of visitations
Malleefowl mound monitoring grids (i.e. in homogenous by animal type and camera site, we used the raw subset
mallee woodland that is known Malleefowl habitat). At count data in a generalised linear model (GLM) with a
Calperum Station, the non-mound cameras were in the negative binomial distribution and an offset for number of
same Malleefowl monitoring grid as four of the active surveillance hours in lme4 (Bates et al. 2015). Pairwise
mounds; the other active mounds were located in the comparisons were made of the significant terms in the
adjacent monitoring grids, ~7 km and 12 km away. At model using the Tukey test in lsmeans (Lenth 2016).
Gluepot Reserve, the non-mound cameras were located Differences were considered significant if probability
P was102 Australian Field Ornithology H. Neilly et al.
presumably feeding; foxes digging, predating a Malleefowl
egg (one instance observed), urinating and defaecating
on the mound; and a cat was observed pouncing into the
centre of a mound. Malleefowl were not present at the
same time as other vertebrate visitors.
From 2016 to 2018, 16 non-mound sites were monitored
Number of visits
by camera surveillance (225,144 h). We identified
215 birds from 18 species, 34 reptiles from 7 species
and 1135 mammals from 6 species, including 141 foxes
(Table 3). Of the total mammals observed, 81% were
Western Grey Kangaroos Macropus fuliginosus.
Using the full dataset, mean total number of vertebrate
visits was more than 1.5 times higher at mounds
(10.64 ± standard error 0.69 visits/1000 h footage) than
non-mounds (6.33 ± 1.24 visits/1000 h footage). Mean
species richness of vertebrate visitors was more than Vertebrate visitor group
six times higher at mounds (1.44 ± 0.21 species/1000 h
footage) than non-mounds (0.22 ± 0.15 species/1000 h Figure 2. The mean number of visits/1000 h of camera
footage ± standard error of vertebrates at Malleefowl
footage). The number of visits by birds, reptiles and foxes
mounds (2012–2018) and non-mounds (2016–2018) at
was higher at mounds, whereas kangaroo numbers were Calperum Station and Gluepot Reserve, South Australia.
higher at non-mounds, and there was very little difference
in the number of visits by cats, goats and other mammals
(Figure 2).
Mound vs non-mound vertebrate visitors:
Subset data
The subset data from 2016 to 2018 at Calperum Station
Number of visits
showed similar trends to the full dataset (Figure 3). Mean
total number of vertebrate visits was significantly higher
at mounds (14.03 ± 5.32 visits/1000 h footage) than non-
mounds (5.84 ± 0.93 visits/1000 h footage). Mean species
richness of vertebrate visitors was significantly higher at
mounds (5.06 ± 0.65 species/1000 h footage) than non-
mounds (1.44 ± 0.16 species/1000 h footage).
The number of visits by birds and reptiles was
significantly higher at mounds compared with non-mound
sites, whereas the inverse was true for kangaroo visits.
Differences among cat, goat and other mammal visits
between mounds and non-mound sites were not detected. Vertebrate visitor group
Likewise, mean fox visits were not significantly different at
mounds compared with non-mound sites. Figure 3. The mean number of visits/1000 h of camera
footage ± standard error of vertebrates at Malleefowl
mounds (n = 3) and non-mounds (n = 8), from subset data
Discussion from 2016–2018, Calperum Station, South Australia. An
asterisk indicates a significant difference between mound
Malleefowl mounds were visited by a range of vertebrate and non-mound sites of each vertebrate visitor group from
a generalised linear model (GLM) with negative binomial
species. The higher overall species richness and visitation distribution (Tukey’s post-hoc test PMalleefowl mounds as habitat for other vertebrates 103
Table 2. Visits (total number and number/1000 h camera footage) by birds, reptiles and mammals recorded at Malleefowl
mounds, 2012–2018, at Calperum Station and Gluepot Reserve, South Australia.
Species Calperum Gluepot
Total No./1000 h Total No./1000 h
Birds 43 5.02 107 4.38
Common Bronzewing Phaps chalcoptera 0 0.00 5 0.20
Australian Owlet-nightjar Aegotheles cristatus 0 0.00 2 0.08
White-eared Honeyeater Nesoptilotis leucotis 0 0.00 1 0.04
Spiny-cheeked Honeyeater Acanthagenys rufogularis 11 1.29 0 0.00
Singing Honeyeater Gavicalis virescens 1 0.12 0 0.00
Yellow-throated Miner Manorina flavigula 1 0.12 9 0.37
White-browed Babbler Pomatostomus superciliosus 0 0.00 2 0.08
Crested Bellbird Oreoica gutturalis 0 0.00 3 0.12
Chestnut Quail-thrush Cinclosoma castanotum 1 0.12 9 0.37
Grey Shrike-thrush Colluricincla harmonica 1 0.12 4 0.16
Pied Currawong Strepera graculina 2 0.23 0 0.00
Grey Currawong Strepera versicolor 0 0.00 2 0.08
Australian Magpie Gymnorhina tibicen 6 0.70 0 0.00
Grey Butcherbird Cracticus torquatus 12 1.40 8 0.33
Masked Woodswallow Artamus personatus 0 0.00 24 0.98
Corvus sp. 1 0.12 14 0.57
White-winged Chough Corcorax melanorhamphos 0 0.00 13 0.53
Red-capped Robin Petroica goodenovii 0 0.00 5 0.20
Southern Scrub-robin Drymodes brunneopygia 0 0.00 3 0.12
Unidentified bird 7 0.82 3 0.12
Reptiles 8 0.93 39 1.60
Mallee Tree Dragon Amphibolurus norrisi 0 0.00 1 0.04
Mallee Military Dragon Ctenophorus fordi 0 0.00 2 0.08
Painted Dragon Ctenophorus pictus 0 0.00 2 0.08
Nobbi Dragon Diporiphora nobbi 0 0.00 6 0.25
Bearded dragon Pogona sp. 7 0.82 22 0.90
Mulga Snake Pseudechis australis 0 0.00 1 0.04
Sleepy Lizard Tiliqua rugosa 1 0.12 0 0.00
Sand Goanna Varanus gouldii 0 0.00 5 0.20
Mammals 46 5.37 97 3.97
Goat Capra hircus 0 0.00 1 0.04
Cat Felis catus 5 0.58 0 0.00
Western Grey Kangaroo Macropus fuliginosus 5 0.58 8 0.33
Sheep Ovis aries 0 0.00 5 0.20
Short-beaked Echidna Tachyglossus aculeatus 0 0.00 6 0.25
Common Brushtail Possum Trichosurus vulpecula 0 0.00 1 0.04
Red Fox Vulpes vulpes 36 4.21 76 3.11104 Australian Field Ornithology H. Neilly et al.
Table 3. Visits (total number and number/1000 h of camera footage) by birds, reptiles and mammals recorded at non-
mound sites, 2016–2018, Calperum Station and Gluepot Reserve, South Australia.
Species Calperum Gluepot
Total No./1000 h Total No./1000 h
Birds 97 0.861 118 1.05
Emu Dromaius novaehollandiae 16 0.142 6 0.05
Malleefowl Leipoa ocellata 9 0.080 4 0.04
Common Bronzewing Phaps chalcoptera 0 0.000 19 0.17
Australian Owlet-nightjar Aegotheles cristatus 1 0.009 1 0.01
Mulga Parrot Psephotellus varius 1 0.009 0 0.00
Red Wattlebird Anthochaera carunculata 0 0.000 1 0.01
Crested Bellbird Oreoica gutturalis 4 0.036 1 0.01
Chestnut Quail-thrush Cinclosoma castanotum 4 0.036 9 0.08
Grey Shrike-thrush Colluricincla harmonica 1 0.009 0 0.00
Australian Magpie Gymnorhina tibicen 9 0.080 12 0.11
Grey Butcherbird Cracticus torquatus 0 0.000 2 0.02
Masked Woodswallow Artamus personatus 7 0.062 0 0.00
Corvus sp. 1 0.009 22 0.20
White-winged Chough Corcorax melanorhamphos 30 0.266 11 0.10
Southern Scrub-robin Drymodes brunneopygia 2 0.018 0 0.00
Unidentified bird 11 0.098 30 0.27
Reptiles 9 0.080 25 0.22
Mallee Tree Dragon Amphibolurus norrisi 0 0.000 1 0.01
Mallee Dragon Ctenophorus fordi 3 0.027 6 0.05
Short-clawed Skink Ctenotus inornatus 0 0.000 3 0.03
Eastern Desert Ctenotus Ctenotus regius 0 0.000 1 0.01
Bearded dragon Pogona sp. 1 0.009 7 0.06
Sleepy Lizard Tiliqua rugosa 0 0.000 4 0.04
Sand Goanna Varanus gouldii 4 0.036 1 0.01
Unidentified reptile 1 0.009 1 0.01
Mammals 447 3.968 688 6.12
Goat Capra hircus 17 0.151 17 0.15
Cat Felis catus 21 0.186 8 0.07
European Hare Lepus europaeus 0 0.000 2 0.02
Western Grey Kangaroo Macropus fuliginosus 329 2.920 585 5.20
Short-beaked Echidna Tachyglossus aculeatus 13 0.115 2 0.02
Red Fox Vulpes vulpes 67 0.595 74 0.66
Kangaroos appeared to avoid active Malleefowl (2010-2011). Considering that only a single predation
mounds, which did not provide vegetative food resources event was observed, foxes might visit mounds because
and perhaps were perceived as an obstacle. Most bird they are inquisitive and tend to visit disturbed areas,
visitors were either exclusively or partially insectivorous rather than visiting mounds to actively pursue predation.
and appeared to be feeding, suggesting that the mound It is, however, suggested that Malleefowl chicks are most
substrate may support a rich invertebrate food source. vulnerable to predation after their emergence from the
The only exclusive granivore recorded, the Common mound (Priddel & Wheeler 1997; Priddel et al. 2007), so
Bronzewing Phaps chalcoptera, was found only at predation events might have occurred near the mound
non-mounds (Table 3). Surprisingly, fox visits were not during chick emergence but were not detected because of
significantly different between mounds and non-mounds the positioning of the mound cameras.
in the subset data, despite this trend appearing in the The interpretation of our results has limitations because
full dataset. Fox numbers were likely higher overall of the study design. The amalgamation of data from two
in 2012–2016 after 2 years of above average rainfall disparate citizen science projects created discrepanciesMalleefowl mounds as habitat for other vertebrates 105
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