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

1Australian Landscape Trust, P.O. Box 955, Renmark SA 5341, 2North Calperum Volunteer Group, 14 Schaefer Drive, Loxton SA 5333, Australia 3BirdLife Australia Gluepot Reserve, via Lunn Road, Waikerie SA 5330, Australia 4Murraylands and Riverland Landscape Board, 110A Mannum Road, Murray Bridge SA 5253, Australia *Corresponding author. Email: [email protected]

Abstract. Ecosystem engineers change the availability of resources for other 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, 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 (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 has 100 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 2746.6 Malleefowl 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 non-mound cameras are indicated on the map. A smaller dataset limited our statistical capabilities but 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 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 was <0.05. Negative binomial distributions were 7.3–18.4 km away from mound cameras (Figure 1). All selected to account for overdispersion and all models were cameras were placed ~600 mm above ground level and validated by examining deviance residuals. Analyses were aimed 15° below horizontal. Camera trigger sensitivity was completed in R (R Core Team 2018). set to normal, a 5-minute delay was imposed following each trigger and only still footage was collected. Results Analysis of camera footage Mound vs non-mound vertebrate visitors: Overall trends The date, time, species, number of individuals, duration of visit and description of activity were recorded for each From 2012 to 2018, 20 active Malleefowl mounds were visit by a vertebrate species. Where species identification monitored via camera surveillance (31,913 hours). We was not possible, visitors were classified as a bird, identified 150 birds from 19 species, 47 reptiles from or mammal. Corvids (Little Crow Corvus bennetti 8 species and 143 mammals from 7 species (Table 2). and Australian Raven C. coronoides) were recorded as Most reptile visits were by bearded dragons (62%) and Corvus sp. and bearded dragons as Pogona sp. as both 79% of mammal visits were by foxes. Where video footage Central Bearded Dragon Pogona vitticeps and Eastern was available, we observed a variety of animal behaviours: Bearded Dragon P. barbarta occur in the survey area but birds foraging and preening (including dust bathing); could not be distinguished on the camera footage. Mound basking (bearded dragons, Mallee Military Dragon cameras were set to record continuously, whereas non- fordi and Painted Dragon C. pictus) or digging mound cameras were set with a 5-minute delay between (Sand Goanna Varanus gouldii); echidnas digging and 102 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 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. Number of visits 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 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- Number of visits 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 and non-mound sites of each vertebrate visitor group from Malleefowl mounds were visited by a range of vertebrate a generalised linear model (GLM) with negative binomial species. The higher overall species richness and visitation distribution (Tukey’s post-hoc test P <0.05). rate at mounds may indicate that mounds provide different types or quantities of resources compared with non- mound areas. Although we did not test these mechanisms, ecosystem, but ecosystem engineering effects in an arid, our observations suggest that mounds may enhance the nutrient-poor context are likely to be more impactful than availability of invertebrate prey for insectivorous birds and mammals, could provide a favourable microclimate for those in mesic environments (Romero et al. 2015; Coggan reptiles to thermoregulate, and the raised structure of the et al. 2018). The density of active Malleefowl mounds in the mound may be ideal for signalling/social communication more arid parts of their range is relatively low (Booth 1987) (Jones 1987). This pilot study is the first examination but, if non-active mounds experience similar visitation of Malleefowl mounds as habitat for other vertebrate to active mounds, this greatly increases the number of species and supports the pursuit of further, robustly mounds providing habitat for other vertebrates per unit designed research in this area. More broadly, it develops area. As we studied only active mounds, we suggest our understanding of Malleefowl as potentially important that further studies should simultaneously assess faunal ecosystem engineers. We were unable to determine the visitors at active mounds, non-active mounds and non- relative importance of mounds as a habitat in the mallee mounds. Malleefowl 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 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 3.11 104 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 discrepancies Malleefowl mounds as habitat for other vertebrates 105

in camera-trap models and settings between the mound Booth, D. (1987). Home range and hatching success of Malleefowl, and non-mound locations. This may affect the detection Leipoa-ocellata Gould (Megapodiidae), in Murray Mallee Near of small animals (Randler & Kalb 2018) although the Renmark, SA. Wildlife Research 14, 95–104. smallest vertebrates observed (Mallee Military Dragons) Byers, J.E., Cuddington, K., Jones, C.G., Talley, T.S., Hastings, A., Lambrinos, J.G., Crooks, J.A. & Wilson, W.G. (2006). Using were detected at both non-mound and mound sites. At ecosystem engineers to restore ecological systems. Trends in non-mounds, the field of view was greater because the Ecology & Evolution 21, 493–500. camera was not obstructed by a mound, so we expected Catano, C.P. & Stout, I.J. (2015). Functional relationships reveal that detectability would be higher at non-mounds. keystone effects of the gopher tortoise on vertebrate diversity However, despite the possible detection disadvantage, in a longleaf pine savanna. Biodiversity and Conservation 24, higher vertebrate visitation rates were still recorded at 1957–1974. mounds. There was substantially greater surveillance time Catterall, C.P. (2018). Fauna as passengers and drivers in vegetation restoration: A synthesis of processes and evidence. at non-mounds compared with mounds. When considering Ecological Management & Restoration 19, 54–62. species richness, it is possible that the lower surveillance Cockle, K.L., Martin, K. & Wesołowski, T. (2011). Woodpeckers, time at mounds has inflated the value of species decay, and the future of cavity-nesting vertebrate communities richness per 1000 hours. Species richness is not directly worldwide. Frontiers in Ecology and the Environment 9, proportional to effort; once richness plateaus, any increase 377–382. in surveillance time will artificially push down value of Coggan, N.V., Hayward, M.W. & Gibb, H. (2018). A global richness per unit time. Additionally, we needed to create a database and “state of the field” review of research into subset of the data, to minimise the confounding influence ecosystem engineering by land animals. Journal of Animal Ecology 87, 974–994. of camera location and time, and this limited the ability to Davidson, A.D., Detling, J.K. & Brown, J.H. (2012). Ecological perform certain statistical analyses. Despite these issues, roles and conservation challenges of social, burrowing, we have confidence that the trends observed support our herbivorous mammals in the world’s grasslands. Frontiers in hypothesis and justify further investigation. In the future, Ecology and the Environment 10, 477–486. it would be beneficial to consider the potential for Engel, F.G., Alegria, J., Andriana, R., Donadi, S., Gusmao, J.B., crossovers and multi-use of related citizen science projects van Leeuwe, M.A., Matthiessen, B. & Eriksson, B.K. (2017). in the design phase, to allow for more seamless integration Mussel beds are biological power stations on intertidal flats. Estuarine, Coastal and Shelf Science 191, 21–27. of data. Frith, H.J. (1959). Breeding of the mallee fowl, Leipoa ocellata Malleefowl are listed internationally and in Australia Gould (Megapodiidae). CSIRO Wildlife Research 4, 31–60. as Vulnerable (IUCN Red List of Threatened Species, Frith, H.J. (1962). Conservation of the mallee fowl, Leipoa ocellata Gould (Megapodiidae). CSIRO Wildlife Research , 33–51. Environment Protection and Biodiversity Conservation Act 7 Gurnell, A.M. (1998). The hydrogeomorphological effects of 1999) and are Endangered in some parts of their range beaver dam-building activity. Progress in Physical Geography: (New South Wales Threatened Species Conservation Act Earth and Environment 22, 167–189 1998, Victorian Advisory List of Threatened Vertebrate Hauser, C.E., Southwell, D., Lahoz-Monfort, J.J., Fauna in Victoria 2013). Most research on Malleefowl Rumpff, L., Benshemesh, J., Burnard, T., van Hespen, J., has focused on identifying threats and opportunities for Wright, J., Wintle, B. & Bode, M. (2019). Adaptive management conservation. Our data suggest that the conservation of informs conservation and monitoring of Australia’s threatened Malleefowl is likely to positively impact a suite of mallee malleefowl. Biological Conservation 233, 31–40. Hofstede, L. & Dziminski, M.A. (2017). Greater bilby burrows: birds and reptiles. Although not tested here, we also predict Important structures for a range of species in an arid that Malleefowl activities will have much broader, long-term environment. Australian Mammalogy 39, 227–237. impacts, influencing invertebrates, soil function and the Jones, C.G., Lawton, J.H. & Shachak, M. (1994). Organisms as germination and establishment of vegetation. Malleefowl ecosystem engineers. Oikos 69, 373–386. may be an important tool to help improve ecosystem Jones, D. (1987). Animals using the incubation mounds of the function in mallee woodlands (Byers et al. 2006; Catterall Australian brush-turkey. Sunbird 17 (2), 32–35. 2018). Jones, D. & Göth, A. (2008). Mound-builders. CSIRO Publishing, Melbourne. Knapp, A.K., Blair, J.M., Briggs, J.M., Collins, S.L., Hartnett, D.C., Johnson, L.C. & Towne, E.G. (1999). The keystone role of bison Acknowledgements in North American tallgrass prairie. BioScience 49, 39–50. We thank the many volunteers from the North Calperum Volunteer Lenth, R.V. (2016). Least-squares means: The R package Group, Friends of Gluepot and staff from Australian Landscape lsmeans. Journal of Statistical Software 69, 1–33. Trust and BirdLlife Australia for assisting with data collection. We McCullough Hennessy, S., Deutschman, D.H., Shier, D.M., thank B. Cale for drawing the site map. Special thanks go to Shirly Nordstrom, L.A., Lenihan, C., Montagne, J-P., Wisinski, C.L. & Wells. This study was funded through the Ian Potter Foundation Swaisgood, R.R. (2016). Experimental habitat restoration for Early Career Researcher Fellowship, the Margaret Middleton conserved species using ecosystem engineers and vegetation Fund and Birds SA project fund. We declare no conflicts of management. Animal Conservation 19, 506–514. interest. Mosbech, A., Johansen, K.L., Davidson, T.A., Appelt, M., Grønnow, B., Cuyler, C., Lyngs, P. & Flora, J. (2018). On the crucial importance of a small bird: The ecosystem services of the little auk (Alle alle) population in northwest Greenland in a References long-term perspective. Ambio 47, 226–243. Bates, D., Maechler, M., Bolker, B. & Walker, S. (2015). Fitting Neilly, H., Wells, D., Pascoe, T. & Cale, P. (2019). G’day mate! linear mixed-effects models using lme4. Journal of Statistical Aussie bush birds spectacular greeting. Frontiers in Ecology Software 67, 1–48. and the Environment 17 (8), 428. Benshemesh, J., Southwell, D., Barker, R. & McCarthy, M. (2020). Neilly, H., Wells, D.E., Pascoe, T. & Cale, P. (2021). Malleefowl Citizen scientists reveal nationwide trends and drivers in the Leipoa ocellata breeding behaviour: Insights from citizen breeding activity of a threatened bird, the Malleefowl (Leipoa science camera surveillance. Australian Field Ornithology 38, ocellata). Biological Conservation 246 108573. 87–98. 106 Australian Field Ornithology H. Neilly et al.

Priddel, D. & Wheeler, R. (1997). Efficacy of fox control in Warnken, J., Hodgkison, S., Wild, C. & Jones, D. (2004). The reducing the mortality of released captive-reared Malleefowl, localized environmental degradation of protected areas Leipoa ocellata. Wildlife Research 24, 469–482. adjacent to bird feeding stations: A case study of the Australian Priddel, D., Wheeler, R. & Copley, P. (2007). Does the integrity brush-turkey Alectura lathami. Journal of Environmental or structure of mallee habitat influence the degree of Fox Management 70, 109–118. predation on Malleefowl (Leipoa ocellata)? Emu 107, 100–107. Weathers, W.W. & Seymour, R.S. (1998). Behaviour and time- R Core Team. (2018). R: A language and environment for activity budgets of Malleefowl Leipoa ocellata in South Australia. statistical computing. R Foundation for Statistical Computing, Emu 98, 288–296. Vienna. Available online: https://www.R-project.org. Webb, J.K. & Whiting, M.J. (2006). Does rock disturbance by Randler, C. & Kalb, N. (2018). Distance and size matters: A superb lyrebirds (Menura novaehollandiae) influence habitat comparison of six wildlife camera traps and their usefulness for selection by juvenile snakes? Austral Ecology 31, 58–67. wild birds. Ecology and Evolution 8, 7151–7163. Wright, J.P. & Jones, C.G. (2006). The concept of organisms as Read, J.L., Carter, J., Moseby, K.M. & Greenville, A. (2008). ecosystem engineers ten years on: Progress, limitations, and Ecological roles of rabbit, bettong and bilby warrens in arid challenges. BioScience 56, 203–209. Australia. Journal of Arid Environments 72, 2124–2130. Romero, G.Q., Gonçalves-Souza, T., Vieira, C. & Koricheva, J. (2015). Ecosystem engineering effects on species diversity across ecosystems: A meta-analysis: Ecosystem engineering Received 23 July 2020, accepted 21 April 2020, effects across ecosystems. Biological Reviews 90, 877–890. published online 3 June 2021 Smith, A., Avitabile, S.C. & Leonard, S.W.J. (2016). Less fuel for the fire: Malleefowl Leipoa( ocellata) nesting activity affects fuel loads and fire behaviour.Wildlife Research 43, 640–648.