Not all predators are equal: a continent‐scale analysis of the effects of predator control on Australian mammals

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Strength of a Trophic Cascade Between an Apex Predator,Mammalian Herbivore and Grasses in a Desert Ecosystem Does Not Vary with Temporal Fluctuations in Primary Productivity

There has long been debate regarding the primacy of bottom-up and top-down effects as factors shaping ecosystems. The exploitation ecosystems hypothesis (EEH) predicts that predators indirectly benefit plants because their top-down effects limit herbivores’ consumption of plants, and that the strength of trophic cascade increases with increasing primary productivity. However, in arid environments, pulses of primary productivity produced by irregular rainfall events could decouple herbivore–plant and predator–prey dynamics if high conversion efficiency from seed biomass to consumers allows the rapid build-up of consumer populations. Here, we test predictions of the EEH in an arid environment. We measured activity/abundances of dingoes, red kangaroos and grasses, and diet of dingoes, in landscapes where dingoes were culled or not culled over 3 years. Dingo activity was correlated with rainfall, and their tracks were less frequent at culled sites. Kangaroo abundance was greater at sites where dingoes were culled and increased with rainfall in the previous 6 months. Grass cover was greater at sites where dingoes were not culled and increased with rainfall in the previous 3 months. During a period of average rainfall, dingoes primarily consumed rodents and increased their consumption of kangaroos during a period of drier conditions. Our results are consistent with the hypothesis that suppression of an apex predator triggers a trophic cascade, but are at odds with the EEH’s prediction that the magnitude of trophic cascades should increase with primary productivity. Our study demonstrates that temporal fluctuations in primary productivity can have effects on biomasses of plants and consumers which are in many ways analogous to those observed along spatial gradients of primary productivity.     

The virtuous circle: predator‐friendly farming and ecological restoration in Australia

In Australia, dingoes are widely regarded as enemies of livestock, and accordingly livestock producers commonly attempt to reduce or eradicate them by lethal control. This can have two forms of perverse outcomes: lethal control often does not succeed in reducing dingo populations and can even result in increased attacks on livestock; and the environmental benefits provided by dingoes, some of which are valuable to livestock production, are lost. We describe these outcomes and suggest mechanisms by which tolerance of dingoes could provide benefits to livestock enterprises, at the same time widening the scope of ecological restoration, and humane treatment of wildlife in Australia.     

Resolving the value of the dingo in ecological restoration

There is global interest in restoring populations of apex predators, both to conserve them and to harness their ecological services. In Australia, reintroduction of dingoes (Canis dingo) has been proposed to help restore degraded rangelands. This proposal is based on theories and the results of studies suggesting that dingoes can suppress populations of prey (especially medium‐ and large‐sized herbivores) and invasive predators such as red foxes (Vulpes vulpes) and feral cats (Felis catus) that prey on threatened native species. However, the idea of dingo reintroduction has met opposition, especially from scientists who query the dingo's positive effects for some species or in some environments. Here, we ask ‘what is a feasible experimental design for assessing the role of dingoes in ecological restoration?’ We outline and propose a dingo reintroduction experiment—one that draws upon the existing dingo‐proof fence—and identify an area suitable for this (Sturt National Park, western New South Wales). Although challenging, this initiative would test whether dingoes can help restore Australia's rangeland biodiversity, and potentially provide proof‐of‐concept for apex predator reintroductions globally.     

Top predators as biodiversity regulators: the dingo Canis lupus dingo as a case study

Top‐order predators often have positive effects on biological diversity owing to their key functional roles in regulating trophic cascades and other ecological processes. Their loss has been identified as a major factor contributing to the decline of biodiversity in both aquatic and terrestrial systems. Consequently, restoring and maintaining the ecological function of top predators is a critical global imperative. Here we review studies of the ecological effects of the dingo Canis lupus dingo, Australia's largest land predator, using this as a case study to explore the influence of a top predator on biodiversity at a continental scale. The dingo was introduced to Australia by people at least 3500 years ago and has an ambiguous status owing to its brief history on the continent, its adverse impacts on livestock production and its role as an ecosystem architect. A large body of research now indicates that dingoes regulate ecological cascades, particularly in arid Australia, and that the removal of dingoes results in an increase in the abundances and impacts of herbivores and invasive mesopredators, most notably the red fox Vulpes vulpes. The loss of dingoes has been linked to widespread losses of small and medium‐sized native mammals, the depletion of plant biomass due to the effects of irrupting herbivore populations and increased predation rates by red foxes. We outline a suite of conceptual models to describe the effects of dingoes on vertebrate populations across different Australian environments. Finally, we discuss key issues that require consideration or warrant research before the ecological effects of dingoes can be incorporated formally into biodiversity conservation programs.     

Are dingoes a trophic regulator in arid Australia? A comparison of mammal communities on either side of the dingo fence

The direct and indirect interactions that large mammalian carnivores have with other species can have far‐reaching effects on ecosystems. In recent years there has been growing interest in the role that Australia's largest terrestrial predator, the dingo, may have in structuring ecosystems. In this study we investigate the effect of dingo exclusion on mammal communities, by comparing mammal assemblages where dingoes were present and absent. The study was replicated at three locations spanning 300 km in the Strzelecki Desert. We hypothesized that larger species of mammal subject to direct interactions with dingoes should increase in abundance in the absence of dingoes while smaller species subject to predation by mesopredators should decrease in abundance because of increased mesopredator impact. There were stark differences in mammal assemblages on either side of the dingo fence and the effect of dingoes appeared to scale with body size. Kangaroos and red foxes were more abundant in the absence of dingoes while Rabbits and the Dusky Hopping‐mouse Notomys fuscus were less abundant where dingoes were absent, suggesting that they may benefit from lower red fox numbers in the presence of dingoes. Feral cats and dunnarts (Sminthopsis spp.) did not respond to dingo exclusion. Our study provides evidence that dingoes do structure mammal communities in arid Australia; however, dingo exclusion is also associated with a suite of land use factors, including sheep grazing and kangaroo harvesting that may also be expected to influence kangaroo and red fox populations. Maintaining or restoring populations of dingoes may be useful strategies to mitigate the impacts of mesopredators and overgrazing by herbivores.     

Prey use by dingoes in a contested landscape: Ecosystem service provider or biodiversity threat?

In Australia, dingoes (Canis lupus dingo) have been implicated in the decline and extinction of a number of vertebrate species. The lowland Wet Tropics of Queensland, Australia is a biologically rich area with many species of rainforest‐restricted vertebrates that could be threatened by dingoes; however, the ecological impacts of dingoes in this region are poorly understood. We determined the potential threat posed by dingoes to native vertebrates in the lowland Wet Tropics using dingo scat/stomach content and stable isotope analyses of hair from dingoes and potential prey species. Common mammals dominated dingo diets. We found no evidence of predation on threatened taxa or rainforest specialists within our study areas. The most significant prey species were northern brown bandicoots (Isoodon macrourus), canefield rats (Rattus sordidus), and agile wallabies (Macropus agilis). All are common species associated with relatively open grass/woodland habitats. Stable isotope analysis suggested that prey species sourced their nutrients primarily from open habitats and that prey choice, as identified by scat/stomach analysis alone, was a poor indicator of primary foraging habitats. In general, we find that prey use by dingoes in the lowland Wet Tropics does not pose a major threat to native and/or threatened fauna, including rainforest specialists. In fact, our results suggest that dingo predation on “pest” species may represent an important ecological service that outweighs potential biodiversity threats. A more targeted approach to managing wild canids is needed if the ecosystem services they provide in these contested landscapes are to be maintained, while simultaneously avoiding negative conservation or economic impacts.     

Eye contact and sociability data suggests that Australian dingoes were never domesticated

Dogs were the first animal to become domesticated by humans, and they represent a classic model system for unraveling the processes of domestication. We compare Australian dingo eye contact and socialization with Basenji and German Shepherd dog (GSD) breeds. Australian dingoes arrived in Australia 5,000–8,000 BP, and there is debate whether they were domesticated before their arrival. The Basenji represents a primitive breed that diverged from the remaining breeds early in the domestication process, while GSDs are a breed dog selected from existing domestic dogs in the late 1800s. We conducted a 4-phase study with unfamiliar and familiar investigators either sitting passively or actively calling each canid. We found 75% of dingoes made eye contact in each phase. In contrast, 86% of Basenjis and 96% of GSDs made eye contact. Dingoes also exhibited shorter eye-gaze duration than breed dogs and did not respond to their name being called actively. Sociability, quantified as a canid coming within 1 m of the experimenter, was lowest for dingoes and highest for GSDs. For sociability duration, dingoes spent less time within 1 m of the experimenter than either breed dog. When compared with previous studies, these data show that the dingo is behaviorally intermediate between wild wolves and Basenji dogs and suggest that it was not domesticated before it arrived in Australia. However, it remains possible that the accumulation of mutations since colonization has obscured historical behaviors, and dingoes now exist in a feralized retamed cycle. Additional morphological and genetic data are required to resolve this conundrum.     

Death by sex in an Australian icon: a continent‐wide survey reveals extensive hybridization between dingoes and domestic dogs

Hybridization between domesticated animals and their wild counterparts can disrupt adaptive gene combinations, reduce genetic diversity, extinguish wild populations and change ecosystem function. The dingo is a free‐ranging dog that is an iconic apex predator and distributed throughout most of mainland Australia. Dingoes readily hybridize with domestic dogs, and in many Australian jurisdictions, distinct management strategies are dictated by hybrid status. Yet, the magnitude and spatial extent of domestic dog–dingo hybridization is poorly characterized. To address this, we performed a continent‐wide analysis of hybridization throughout Australia based on 24 locus microsatellite DNA genotypes from 3637 free‐ranging dogs. Although 46% of all free‐ranging dogs were classified as pure dingoes, all regions exhibited some hybridization, and the magnitude varied substantially. The southeast of Australia was highly admixed, with 99% of animals being hybrids or feral domestic dogs, whereas only 13% of the animals from remote central Australia were hybrids. Almost all free‐ranging dogs had some dingo ancestry, indicating that domestic dogs could have poor survivorship in nonurban Australian environments. Overall, wild pure dingoes remain the dominant predator over most of Australia, but the speed and extent to which hybridization has occurred in the approximately 220 years since the first introduction of domestic dogs indicate that the process may soon threaten the persistence of pure dingoes.     

Exploitation ecosystems and trophic cascades in non‐equilibrium systems: pasture – red kangaroo – dingo interactions in arid Australia

The exploitation ecosystems hypothesis (EEH) proposes that 1) plant biomass reflects the primary productivity of an ecosystem modified by the regulating effect of herbivory, and 2) herbivore abundance reflects the productivity of plants modified by the regulating effect of predation. Primary productivity thus determines the number of trophic levels in an ecosystem and the extent to which bottom–up and top–down regulation influence the biomass ratios of adjacent and non‐adjacent trophic levels (i.e. trophic cascading). We constructed an interactive model of plant (pasture), herbivore (red kangaroo Macropus rufus) and predator (dingo Canis lupus dingo), a system in which trophic cascades have been suggested to occur, and used it to test the effects of increasing stochastic variation in primary productivity and dingo culling on predictions of the EEH. The model contained four feedback loops: the predator–herbivore and herbivore–plant feedback loops, and the predator and plant density‐dependent feedback loops. The equilibrium conditions along the primary productivity gradient reproduced the three zones of trophic dynamics predicted by the EEH, plus an additional zone at productivities above which the maximum density of a predator is achieved due to social regulation: that zone is characterized by increasing herbivore density and decreasing plant biomass. Culling dingoes produced trophic cascades that were strongly attenuated at primary productivities below which the maximum density of dingoes was attained. Results were robust to uncertainty in kangaroo off‐take by dingoes and to the efficacy of dingo culling, but prey switching by dingoes from red kangaroos to reptiles would weaken trophic cascades. We conclude that social regulation of carnivores has important implications for expression of the EEH and trophic cascades, and that attenuation of trophic cascades increases with increasing stochasticity in primary productivity. Our model also provides a framework for understanding the conditions in which dingo‐mediated trophic cascades might be expected to occur, and generates testable predictions about the effects of higher dingo densities (e.g. by stopping culling or reintroduction to former range) on kangaroo and pasture dynamics.     

Biodiversity friend or foe: land use by a top predator,the dingo in contested landscapes of the Australian Wet Tropics

Dingoes (Canis dingo) in the coastal lowlands of Australia's Wet Tropics are perceived as a major threat to biodiversity and subjected to broad‐spectrum lethal control. However, evidence of their impacts is equivocal, and control programmes generally ignore the ecological benefits that dingoes might provide. Previous diet analysis has shown that dingoes in the Wet Tropics primarily prey on common, terrestrial mammals. However, little is known of dingo habitat use or prey acquisition in the region despite these activities having major implications for biodiversity conservation. We investigated land use by dingoes in the lowland Wet Tropics to enable predictions of potential prey types, relative prey use and modes of prey acquisition. Nine dingoes were tracked for 3–6 months. Home ranges and resting areas were estimated using multiple estimators, and habitat use was analysed using compositional analysis of habitat use and generalized additive models. Dingo ranging behaviour suggested that anthropogenic food subsidies were infrequently used. Each territory comprised several sclerophyll forest rest areas with adjacent sugarcane‐grassland high activity areas. Individuals used each rest‐activity area for extended durations before moving on to another. Sclerophyll and rainforests, which contain the fauna species of primary conservation concern, were generally used for rest/sleep, or movement between rest‐activity areas. Activity patterns were consistent with dingoes hunting in open sugarcane‐grassland habitats during daylight hours. Dingo activity was low in areas where fauna species of conservation concern occur, which suggests that dingoes do not pose a threat to their survival. Consequently, current broad‐spectrum lethal control may have minimal benefits or even incur costs for biodiversity. Maximizing the ecosystem services provided by dingoes while simultaneously minimizing their negative impacts requires a more targeted location‐specific management approach, one that assesses and mitigates impacts specifically where background circumstances suggest particular packs may be either a conservation or economic threat.     

Habitat selection by feral cats and dingoes in a semi‐arid woodland environment in central Australia

Habitat use by feral cats and dingoes was examined within a heterogeneous semi‐arid woodland site in central Australia over 2 years. Density estimates of feral cats based on tracks were higher in mulga habitat than in open habitat. Isodar analysis implied that this pattern of habitat use by feral cats was consistent with the consumer‐resource model of density‐dependent habitat selection, which is an ideal free solution. The reason why mulga supported higher densities of feral cats was unclear. Foraging success of feral cats may be higher in the mulga because the stalk and ambush hunting tactics typically employed by felids are well suited to dense cover. Mulga may also have offered feral cats more protection from dingo predation. Dingo activity was distributed uniformly across habitats. The dingo isodar was statistically non‐significant, suggesting that habitat selection by dingoes was independent of density.     

Home range,activity and sociality of a top predator,the dingo: a test of the Resource Dispersion Hypothesis

The idea that groups of individuals may develop around resource patches led to the formulation of the Resource Dispersion Hypothesis (RDH). We tested the predictions of the RDH, within a quasi‐experimental framework, using Australia’s largest terrestrial predator, the dingo Canis lupus dingo. Average dingo group sizes were higher in areas with abundant focal food sources around two mine sites compared with those in more distant areas. This supports the notion that resource richness favours larger group size, consistent with the RDH. Irrespective of season or sex, average home range estimates and daily activity for dingoes around the mine sites were significantly less than for dingoes that lived well away. Assuming that a territory is the defended part of the home range and that territory size is correlated with home range size, consistent with the RDH, the spatial dispersion of food patches therefore determined territory size for dingoes in our study. However, although sample size was small, some dingoes that accessed the supplementary food resource at the mines also spent a large proportion of their time away, suggesting a breakdown of territorial defence around the focal food resource. This, in combination with the large variation in home range size among dingoes that accessed the same supplementary food resource, limits the predictive capabilities of the RDH for this species. We hypothesize that constraints on exclusive home range occupancy will arise if a surfeit of food resources (in excess of requirements for homeostasis) is available in a small area, and that this will have further effects on access to mates and social structure. We present a conceptual model of facultative territorial defence where focal resources are available to demonstrate our findings.     

Could a rabies incursion spread in the northern Australian dingo population? Development of a spatial stochastic simulation model

Australia, home to the iconic dingo, is currently free from canine rabies. However northern Australia, including Indigenous communities with large free-roaming domestic dog populations, is at increased risk of rabies incursion from nearby Indonesia. We developed a novel agent-based stochastic spatial rabies spread model to evaluate the potential spread of rabies within the dingo population of the Northern Peninsula Area (NPA) region of northern Australia. The model incorporated spatio-temporal features specific to this host-environment system, including landscape heterogeneity, demographic fluctuations, dispersal movements and dingo ecological parameters—such as home range size and density—derived from NPA field studies. Rabies spread between dingo packs in nearly 60% of simulations. In such situations rabies would affect a median of 22 dingoes (approximately 14% of the population; 2.5–97.5 percentiles: 2–101 dingoes) within the study area which covered 1,131 km², and spread 0.52 km/week for 191 days. Larger outbreaks occurred in scenarios in which an incursion was introduced during the dry season (vs. wet season), and close to communities (vs. areas with high risk of interaction between dingoes and hunting community dogs). Sensitivity analyses revealed that home range size and duration of infectious clinical period contributed most to the variance of outputs. Although conditions in the NPA would most likely not support a sustained propagation of the disease in the dingo population, due to the predicted number of infected dingoes following a rabies incursion and the proximity of Indigenous communities to dingo habitat, we conclude that the risk for human transmission could be substantial.     

Could direct killing by larger dingoes have caused the extinction of the thylacine from mainland Australia?

Invasive predators can impose strong selection pressure on species that evolved in their absence and drive species to extinction. Interactions between coexisting predators may be particularly strong, as larger predators frequently kill smaller predators and suppress their abundances. Until 3500 years ago the marsupial thylacine was Australia's largest predator. It became extinct from the mainland soon after the arrival of a morphologically convergent placental predator, the dingo, but persisted in the absence of dingoes on the island of Tasmania until the 20th century. As Tasmanian thylacines were larger than dingoes, it has been argued that dingoes were unlikely to have caused the extinction of mainland thylacines because larger predators are rarely killed by smaller predators. By comparing Holocene specimens from the same regions of mainland Australia, we show that dingoes were similarly sized to male thylacines but considerably larger than female thylacines. Female thylacines would have been vulnerable to killing by dingoes. Such killing could have depressed the reproductive output of thylacine populations. Our results support the hypothesis that direct killing by larger dingoes drove thylacines to extinction on mainland Australia. However, attributing the extinction of the thylacine to just one cause is problematic because the arrival of dingoes coincided with another the potential extinction driver, the intensification of the human economy.     

Phenotypic variation and promiscuity in a wild population of pure dingoes (Canis dingo)

Phenotypic diversity occurs in natural populations as a result of the interaction between an individual's genotype and the environment. Nevertheless, individual variation in phenotypic traits such as coat colour and body size is routinely used to differentiate between “pure” dingoes Canis dingo and dingo‐dog hybrids. Extensive anthropogenic impacts and widespread hybridization with domestic dogs has hindered our ability to study intact dingo populations and, therefore, most of our basic understanding of dingo biology (e.g., phenotypic variation, mating systems, genetic diversity) stems from observational studies on perturbed populations. We sampled a relatively undisturbed population of dingoes, from arid Australia, to determine their purity and genetic diversity. We explored their mating strategy using a pedigree built from genetic data and examined how phenotypic variation was influenced by age, sex, heterozygosity, and relatedness. Coat colour was our measure of phenotype and our population displayed four types (sandy, black & tan, white, and sable). All dingoes (n = 83) possessed a high level of dingo ancestry (mean purity > 90%) and were closely related to each other; with all but one individual related as full‐sibling or parent–offspring. Our pedigree shows both monogamous and promiscuous mating strategies exist within an undisturbed population. Variation in coat colour or body size cannot be used to infer a dingo's level of purity because the phenotype of pure dingoes is intrinsically variable. The breeding system of dingoes was long thought to be monogamous, but we provide genetic evidence for numerous mating strategies including both long‐term monogamy and extreme promiscuity.     

Dingo Howls: The Content and Efficacy of a Long‐Range Vocal Signal

Signals are shaped through selective pressures impacting their information content (i.e. ‘what’ is conveyed) and their efficacy (i.e. ‘how’ is it conveyed). In the case of long‐range communication, a signal's structure should both convey information and allow for detection and discrimination despite the effects of environmental variability during propagation. Canid howls are an ideal example of a long‐range communication signal, although research has mostly focused on the relationships between howls’ acoustic characteristics and their potential information content in wolves, Canis lupus spp., and coyotes, Canis latrans. In this study, we quantified these relationships in another canid, the dingo, Canis familiaris dingo, to determine how their howls compare to that of other wild canids. We found that dingoes’ howls are individually distinctive and may convey gender cues. Playback experiments indicate that dingoes can perform class‐level recognition of familiar and unfamiliar individuals on the basis of vocalizations alone. These results mirror findings for other wild canids such as wolves and coyotes. Lastly, propagation experiments demonstrate that this signal's structure can permit long‐distance detectability and discriminability. We discuss our results in relation to the putative functions of canid howls and propose avenues for future research.     

Temporal and spatial trends in the abundances of an apex predator,introduced mesopredator and ground-nesting bird are consistent with the mesopredator release hypothesis

Apex predator extirpation has been identified as a key driver of biodiversity losses. The mesopredator release hypothesis (MRH) predicts that reduced abundance of apex predators results in an increase in the abundance and predatory impact of mesopredators. Here we test predictions made according to the MRH that an apex predator, the dingo (Canis dingo), benefits a small ground-nesting bird, the little button-quail (Turnix velox), by reducing the abundance of introduced mesopredators, the red fox (Vulpes vulpes) and feral cat (Felis catus). We also examined an alternative hypothesis that herbivore grazing negatively affects little button-quail abundance by reducing ground cover. To test our predictions we compared dingo, mesopredator, quail, herbivore and ground cover abundances and predator diets over a 25 month period and across a 10,000 km² region encompassing areas where dingoes were common and rare, pastoral properties, and conservation reserves. Little button-quails were primarily observed where dingoes were common and foxes rare. Cats were detected at low numbers throughout the sample area irrespective of the index abundance of little button-quails, dingoes or foxes. Birds occurred less frequently in dingo than fox or cat scats. Ground cover and herbivore grazing activity were poor correlates of little button-quail abundance. Our results are consistent with the hypothesis that apex predators’ mesopredator-suppressive effects translate to population-level benefits for a ground-nesting bird. Positive associations between the abundances of dingoes and small-prey species suggests that positive management of dingoes could be incorporated into broad-scale biodiversity conservation programs as a strategy to alleviate the predatory impacts of foxes.     

Testing target‐specific bait delivery for controlling feral pigs in a tropical rainforest

Mitigation of feral pig (Sus scrofa) impacts in Australia’s Wet Tropics World Heritage Area has been impeded by the lack of a target‐specific method for delivering toxic baits in the region. This study evaluated methods to reduce bait‐take by susceptible nontarget species without inhibiting bait‐take by pigs, to enable more effective pig management. We predicted that dingoes would not select an unprocessed corn bait and that other potential nontarget bait consumers would be unable to access the same bait presented under a lightweight cover. Neither of these methods was expected to reduce bait selection or access by pigs. We tested these predictions by monitoring animal interactions with covered and uncovered corn baits, and covered corn and manufactured baits. Use of corn as a bait substrate effectively prevented bait‐take by dingoes. Covering baits substantially reduced bait‐take by other nontarget species and completely prevented nontarget bait‐take when uncovered feed was provided simultaneously. The corn bait preparation was highly acceptable and accessible to feral pigs. We conclude that the methods evaluated here could enable the consideration of poison baiting as a viable method for controlling feral pigs in the World Heritage Area, where it has previously been unavailable.     

Prevalence of Toxoplasma gondii antibodies in dingoes

Serum samples from 62 dingoes (Canis familiaris dingo) trapped in five areas of southeastern New South Wales, Australia were tested for antibodies to Toxoplasma gondii. Six (10%) of the dingoes had direct agglutination test titers for T. gondii of greater than or equal to 1:64, and four of these animals had T. gondii-specific IgM, suggesting recent exposure.