Predator interactions, mesopredator release and biodiversity conservation

There is growing recognition of the important roles played by predators in regulating ecosystems and sustaining biodiversity. Much attention has focused on the consequences of predator-regulation of herbivore populations, and associated trophic cascades. However apex predators may also control smaller ‘mesopredators’ through intraguild interactions. Removal of apex predators can result in changes to intraguild interactions and outbreaks of mesopredators (‘mesopredator release’), leading in turn to increased predation on smaller prey. Here we provide a review and synthesis of studies of predator interactions, mesopredator release and their impacts on biodiversity. Mesopredator suppression by apex predators is widespread geographically and taxonomically. Apex predators suppress mesopredators both by killing them, or instilling fear, which motivates changes in behaviour and habitat use that limit mesopredator distribution and abundance. Changes in the abundance of apex predators may have disproportionate (up to fourfold) effects on mesopredator abundance. Outcomes of interactions between predators may however vary with resource availability, habitat complexity and the complexity of predator communities. There is potential for the restoration of apex predators to have benefits for biodiversity conservation through moderation of the impacts of mesopredators on their prey, but this requires a whole-ecosystem view to avoid unforeseen negative effects.

‘Nothing has changed since I began.

My eye has permitted no change.

I am going to keep things like this.’

From ‘Hawk Roosting’, by Ted Hughes.

Predator control promotes invasive dominated ecological states

Invasive species are regarded as one of the top five drivers of the global extinction crisis. In response, extreme measures have been applied in an attempt to control or eradicate invasives, with little success overall. We tested the idea that state shifts to invasive dominance are symptomatic of losses in ecosystem resilience, due to the suppression of apex predators. This concept was investigated in Australia where the high rate of mammalian extinctions is largely attributed to the destructive influence of invasive species. Intensive pest control is widely applied across the continent, simultaneously eliminating Australia’s apex predator, the dingo (Canis lupus dingo). We show that predator management accounts for shifts between two main ecosystem states. Lethal control fractures dingo social structure and leads to bottom-up driven increases in invasive mesopredators and herbivores. Where control is relaxed, dingoes re-establish top–down regulation of ecosystems, allowing for the recovery of biodiversity and productivity.

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.

Effects of predator control on behaviour of an apex predator and indirect consequences for mesopredator suppression

1. Apex predators can benefit ecosystems through top–down control of mesopredators and herbivores. However, apex predators are often subject to lethal control aimed at minimizing attacks on livestock. Lethal control can affect both the abundance and behaviour of apex predators. These changes could in turn influence the abundance and behaviour of mesopredators.

2. We used remote camera surveys at nine pairs of large Australian rangeland properties, comparing properties that controlled dingoes Canis lupus dingo with properties that did not, to test the effects of predator control on dingo activity and to evaluate the responses of a mesopredator, the feral cat Felis catus.

3. Indices of dingo abundance were generally reduced on properties that practiced dingo control, in comparison with paired properties that did not, although the effect size of control was variable. Dingoes in uncontrolled populations were crepuscular, similar to major prey. In populations subject to control, dingoes became less active around dusk, and activity was concentrated in the period shortly before dawn.

4. Shifts in feral cat abundance indices between properties with and without dingo control were inversely related to corresponding shifts in indices of dingo abundance. There was also a negative relationship between predator visitation rates at individual camera stations, suggesting cats avoided areas where dingoes were locally common. Reduced activity by dingoes at dusk was associated with higher activity of cats at dusk.

5. Our results suggest that effective dingo control not only leads to higher abundance of feral cats, but allows them to optimize hunting behaviour when dingoes are less active. This double effect could amplify the impacts of dingo control on prey species selected by cats. In areas managed for conservation, stable dingo populations may thus contribute to management objectives by restricting feral cat access to prey populations.

6. Synthesis and applications. Predator control not only reduces indices of apex predator abundance but can also modify their behaviour. Hence, indicators other than abundance, such as behavioural patterns, should be considered when estimating a predator's capacity to effectively interact with lower trophic guilds. Changes to apex predator behaviour may relax limitations on the behaviour of mesopredators, providing enhanced access to resources and prey.

Status and ecological effects of the world's largest carnivores

Large carnivores face serious threats and are experiencing massive declines in their populations and geographic ranges around the world. We highlight how these threats have affected the conservation status and ecological functioning of the 31 largest mammalian carnivores on Earth. Consistent with theory, empirical studies increasingly show that large carnivores have substantial effects on the structure and function of diverse ecosystems. Significant cascading trophic interactions, mediated by their prey or sympatric mesopredators, arise when some of these carnivores are extirpated from or repatriated to ecosystems. Unexpected effects of trophic cascades on various taxa and processes include changes to bird, mammal, invertebrate, and herpetofauna abundance or richness; subsidies to scavengers; altered disease dynamics; carbon sequestration; modified stream morphology; and crop damage. Promoting tolerance and coexistence with large carnivores is a crucial societal challenge that will ultimately determine the fate of Earth’s largest carnivores and all that depends upon them, including humans.

Dingoes can help conserve wildlife and our methods can tell

Management of apex predators is among the most controversial wildlife management issues globally. In Australia, some ecologists have advocated using the dingo, Canis dingo, as a tool for conservation management, due to evidence that they suppress invasive mesopredators. Hayward & Marlow (Journal of Applied Ecology, 51, 2014 and 835) questioned the capacity of dingoes to provide benefits to native biodiversity due to their inability to eradicate foxes and cats. They also argued that indices of abundance commonly used in studies of mesopredator release by dingoes (namely, track-based indices) invalidate the conclusions of the studies. Hayward & Marlow caution conservation practitioners against incorporating dingoes into conservation programmes. Counter to their claims, we summarise research showing that the suppression of invasive mesopredators (cf. eradication) can enhance populations of native species and is therefore a meaningful conservation objective. We highlight literature supporting the hypothesis that dingoes suppress mesopredator abundance and activity, which in turn benefits native biodiversity. We show that Hayward & Marlow overlook many studies of carnivores that show track indices capture a large amount of the variation in the density of medium- and large-sized carnivores. Synthesis and applications. Practitioners cannot afford to wait to act given the perilous state of Australia's mammal species, and we argue that the evidence is sufficiently strong to justify managing dingoes for biodiversity conservation.

Incorporating anthropogenic effects into trophic ecology: predator-prey interactions in a human-dominated landscape

Apex predators perform important functions that regulate ecosystems worldwide. However, little is known about how ecosystem regulation by predators is influenced by human activities. In particular, how important are top-down effects of predators relative to direct and indirect human-mediated bottom-up and top-down processes? Combining data on species' occurrence from camera traps and hunting records, we aimed to quantify the relative effects of top-down and bottom-up processes in shaping predator and prey distributions in a human-dominated landscape in Transylvania, Romania. By global standards this system is diverse, including apex predators (brown bear and wolf), mesopredators (red fox) and large herbivores (roe and red deer). Humans and free-ranging dogs represent additional predators in the system. Using structural equation modelling, we found that apex predators suppress lower trophic levels, especially herbivores. However, direct and indirect top-down effects of humans affected the ecosystem more strongly, influencing species at all trophic levels. Our study highlights the need to explicitly embed humans and their influences within trophic cascade theory. This will greatly expand our understanding of species interactions in human-modified landscapes, which compose the majority of the Earth's terrestrial surface.

How small-scale variation in oyster reef patchiness influences predation on bivalves

As oyster fishing continues to degrade reef habitat along the US Atlantic coast, oyster reefs appear increasingly fragmented on small spatial scales. In outdoor mesocosms, experiments tested how consumption of representatives of 4 different bivalve guilds by each of 3 mesopredators varies between continuous and fine-scale patches of oyster reef habitat. The mesopredator that fed least (stone crab) exhibited no detectable change in consumption on any bivalve (ribbed mussel, bay scallop, hard clam, and 3 size classes of eastern oyster). Consumption of bay scallops by both blue crabs and sheepshead fish was greater in small patches than in continuous oyster reef habitat. Of the bivalve guilds tested, only the scallop possesses swimming motility sufficient to reduce predation, an escape response that would likely leave the bivalve protected within structured habitat in larger continuous oyster reefs. Sheepshead consumed more small oysters in the continuous habitat than in the fine patches, while no other predator-prey interaction exhibited differential feeding as a function of habitat patchiness. Consequently, predation by mesopredators on bivalves can vary with the scale of oyster reef patchiness, but this process may depend upon the bivalve guild. Understanding the role of habitat patchiness on fine scales may be increasingly important in view of the declines in apex predatory sharks leading to mesopredator release, and global climate change directly and indirectly enhancing stone crab abundances, thereby increasing potential predation on bivalves.