Philip A. Stephens

Consequences of the Allee effect for behaviour, ecology and conservation

Warder C. Allee brought attention to the possibility of a positive relationship between aspects of fitness and population size 50 years ago. Until recently, however, this concept was generally regarded as an intriguing but relatively unimportant aspect of population ecology. Increasing appreciation that Allee effects must be incorporated into models of population dynamics and habitat use, together with recent interest in the implications of sociality for conservation, have shown that for ecology and conservation the consequences of the Allee effect are profound. The Allee effect can be regarded not only as a suite of problems associated with rarity, but also as the basis of animal sociality.

What Is the Allee Effect

W. C. Allee brought attention to the possibility of a positive relationship between aspects of fitness and population size over fifty years ago. This phenomenon, frequently termed the Allee effect, has been the focus of increased interest over the past two decades in the light of concerns over conservation and the problems of rarity. Use of the term suffers from the absence of a clear definition however, with the result that Allee effects are frequently thought to involve only a narrow range of phenomena and are often overlooked altogether. We propose a definition for the effect and attempt to resolve the major issues underlying the confusion surrounding this term.

Model selection and model averaging in behavioural ecology: the utility of the IT-AIC framework

Behavioural ecologists often study complex systems in which multiple hypotheses could be proposed to explain observed phenomena. For some systems, simple controlled experiments can be employed to reveal part of the complexity; often, however, observational studies that incorporate a multitude of causal factors may be the only (or preferred) avenue of study. We assess the value of recently advocated approaches to inference in both contexts. Specifically, we examine the use of information theoretic (IT) model selection using Akaike’s information criterion (AIC). We find that, for simple analyses, the advantages of switching to an IT-AIC approach are likely to be slight, especially given recent emphasis on biological rather than statistical significance. By contrast, the model selection approach embodied by IT approaches offers significant advantages when applied to problems of more complex causality. Model averaging is an intuitively appealing extension to model selection. However, we were unable to demonstrate consistent improvements in prediction accuracy when using model averaging with IT-AIC; our equivocal results suggest that more research is needed on its utility. We illustrate our arguments with worked examples from behavioural experiments.

Evolution of trust and trustworthiness: social awareness favours personality differences

Interest in the evolution and maintenance of personality is burgeoning. Individuals of diverse animal species differ in their aggressiveness, fearfulness, sociability and activity. Strong trade-offs, mutation–selection balance, spatio-temporal fluctuations in selection, frequency dependence and good-genes mate choice are invoked to explain heritable personality variation, yet for continuous behavioural traits, it remains unclear which selective force is likely to maintain distinct polymorphisms. Using a model of trust and cooperation, we show how allowing individuals to monitor each others cooperative tendencies, at a cost, can select for heritable polymorphisms in trustworthiness. This variation, in turn, favours costly ‘social awareness’ in some individuals. Feedback of this sort can explain the individual differences in trust and trustworthiness so often documented by economists in experimental public goods games across a range of cultures. Our work adds to growing evidence that evolutionary game theorists can no longer afford to ignore the importance of real world inter-individual variation in their models.

Minimum viable populations: is there a 'magic number' for conservation practitioners?

Establishing species conservation priorities and recovery goals is often enhanced by extinction risk estimates. The need to set goals, even in data-deficient situations, has prompted researchers to ask whether general guidelines could replace individual estimates of extinction risk. To inform conservation policy, recent studies have revived the concept of the minimum viable population (MVP), the population size required to provide some specified probability of persistence for a given period of time. These studies conclude that long-term persistence requires ≥5000 adult individuals, an MVP threshold that is unaffected by taxonomy, life history or environmental conditions. Here, we re-evaluate this suggestion. We find that neither data nor theory supports its general applicability, raising questions about the utility of MVPs for conservation planning.

Impact of livestock and settlement on the large mammalian wildlife of Bale Mountains National Park, southern Ethiopia

Large mammals, both wild and domestic, were censused in four study areas in Bale Mountains National Park (BMNP), southern Ethiopia, from April until August 1997, using established road counts, horse-back counts and systematic transects, a new approach to censusing large mammals in BMNP. Data collected since 1983 were also examined to indicate trends in mammalian abundance. Civil unrest following the government changeover in 1991 disrupted management of BMNP, leading to an increase in human settlement and livestock densities. The effect of these factors on BMNPs wildlife is assessed and discussed. Most large mammalian wildlife has declined since 1983 but signs of recovery for some species in recent years are positive. The fates of two charismatic endemics are emphasised. BMNPs current population of mountain nyala (Tragelaphus buxtoni, an endangered antelope) was estimated to be between 1100 and 1300. This is lower than estimates for the late 1980s, but shows an increase from very low levels recorded following social turmoil during the government changeover in 1991. Data on the Ethiopian wolf (Canis simensis, a critically endangered canid) also indicate positive trends in abundance, following severe declines between 1989 and 1992 due to rabies epizootics. Action to reduce human utilisation of parts of BMNP is urgently required to prevent further degradation of the environment and to safeguard the future of both the mountain nyala and Ethiopian wolf.

Management by proxy? The use of indices in applied ecology

encourages contributions thatcan influence environmental management, policy or both,with evidence based on the most robust science possible.Natural resource management is often contentious, andany perceived weaknesses in the underpinning science areeasily exploited by interest groups to undermine the widerendeavour (see, e.g. the experiences of the Intergovern-mental Panel on Climate Change, Ravindranath 2010).Thus, the robustness of science designed to underpin man-agement and policy is particularly important.Unfortunately, robust and unambiguous results are diffi-cult to obtain in ecology. In particular, causal pathways inecology are seldom linear, but are part of a ‘vast web ofcause and effect’ of which, typically, we can study only asmall part (Peters 1991; p. 134). Meaningful spatial andtemporal scales for ecological processes often defy experi-ments, controlled manipulations and adequate replication;most modern ecological science is reliant on observationaldata and correlation is far easier to demonstrate than cau-sation (

Consistent response of bird populations to climate change on two continents.

Birds populations allied in abundance Changes in climate can cause populations of species to decline, to increase, or to remain steady. Stephens et al. looked across species of common birds in Europe and the United States. Despite many differences between the two regions, expectations about how a species might respond to climate change did predict actual responses. Species predicted to benefit from increasing temperatures, or their associated effects, tended to increase, whereas those predicted to be negatively affected declined. Thus, even across widely varying ecological conditions and communities, climate change can be expected to alter population sizes. Science, this issue p. 84 The impact of climate change on population sizes of birds across continents can be predicted. Global climate change is a major threat to biodiversity. Large-scale analyses have generally focused on the impacts of climate change on the geographic ranges of species and on phenology, the timing of ecological phenomena. We used long-term monitoring of the abundance of breeding birds across Europe and the United States to produce, for both regions, composite population indices for two groups of species: those for which climate suitability has been either improving or declining since 1980. The ratio of these composite indices, the climate impact indicator (CII), reflects the divergent fates of species favored or disadvantaged by climate change. The trend in CII is positive and similar in the two regions. On both continents, interspecific and spatial variation in population abundance trends are well predicted by climate suitability trends.

Warning displays may function as honest signals of toxicity

Many prey species use colourful ‘aposematic’ signalling to advertise the fact that they are toxic. Some recent studies have shown that the brightness of aposematic displays correlates positively with the strength of toxicity, suggesting that aposematic displays are a form of handicap signal, the conspicuousness of which reliably indicates the level of toxicity. The theoretical consensus in the literature is, however, at odds with this finding. It is commonly assumed that the most toxic prey should have less bright advertisements because they have better chances of surviving attacks and can therefore reduce the costs incurred by signalling. Using a novel theoretical model, we show that aposematic signals can indeed function as handicaps. To generate this prediction, we make a key assumption that the expression of bright displays and the storage of anti-predator toxins compete for resources within prey individuals. One shared currency is energy. However, competition for antioxidant molecules, which serve dual roles as pigments and in protecting prey against oxidative stress when they accumulate toxins, provides a specific candidate resource that could explain signal honesty. Thus, contrary to the prevailing theoretical orthodoxy, warning displays may in fact be honest signals of the level of (rather than simply the existence of) toxicity.

The bigger they come, the harder they fall: body size and prey abundance influence predator–prey ratios

Large carnivores are highly threatened, yet the processes underlying their population declines are still poorly understood and widely debated. We explored how body mass and prey abundance influence carnivore density using data on 199 populations obtained across multiple sites for 11 carnivore species. We found that relative decreases in prey abundance resulted in a five- to sixfold greater decrease in the largest carnivores compared with the smallest species. We discuss a number of possible causes for this inherent vulnerability, but also explore a possible mechanistic link between predator size, energetics and population processes. Our results have important implications for carnivore ecology and conservation, demonstrating that larger species are particularly vulnerable to anthropogenic threats to their environment, especially those which have an adverse affect on the abundance of their prey.