Fear creates an Allee effect: experimental evidence from seasonal populations.
ABSTRACT: Allee effects driven by predation can play a strong role in the decline of small populations but are conventionally thought to occur when generalist predators target specific prey (i.e. type II functional response). However, aside from direct consumption, fear of predators could also increase vigilance and reduce time spent foraging as population size decreases, as has been observed in wild mammals living in social groups. To investigate the role of fear on fitness in relation to population density in a species with limited sociality, we exposed varying densities of Drosophila melanogaster to mantid predators either during an experimental breeding season or non-breeding season. The presence of mantids in either season decreased the reproductive performance of individuals but only at low breeding densities, providing evidence for an Allee effect. We then used our experimental results to parametrize a mathematical model to examine the population consequences of fear at low densities. Fear tended to destabilize population dynamics and increase the risk of extinction up to sevenfold. Our study provides unique experimental evidence that the indirect effects of the presence of predators can cause an Allee effect and has important consequences for our understanding of the dynamics of small populations.
Project description:Praying mantids are predators that consume a wide variety of insects. While the gut microbiome of carnivorous mammals is distinct from that of omnivores and herbivores, the role of the gut microbiome among predatory insects is relatively understudied. Praying mantids are the closest known relatives to termites and cockroaches, which are known for their diverse gut microbiota. However, little is known about the mantid gut microbiota or their importance to host health. In this work, we report the results of a 16S rRNA gene-based study of gut microbiome composition in adults and late-instar larvae of three mantid species. We found that the praying mantis gut microbiome exhibits substantial variation in bacterial diversity and community composition. The hindgut of praying mantids were often dominated by microbes that are present in low abundance or not found in the guts of their insect prey. Future studies will explore the role of these microbes in the digestion of the dietary substrates and/or the degradation of toxins produced by their insect prey.
Project description:Some eyespots are thought to deflect attack away from the vulnerable body, yet there is limited empirical evidence for this function and its adaptive advantage. Here, we demonstrate the conspicuous ventral hindwing eyespots found on Bicyclus anynana butterflies protect against invertebrate predators, specifically praying mantids. Wet season (WS) butterflies with larger, brighter eyespots were easier for mantids to detect, but more difficult to capture compared to dry season (DS) butterflies with small, dull eyespots. Mantids attacked the wing eyespots of WS butterflies more frequently resulting in greater butterfly survival and reproductive success. With a reciprocal eyespot transplant, we demonstrated the fitness benefits of eyespots were independent of butterfly behaviour. Regardless of whether the butterfly was WS or DS, large marginal eyespots pasted on the hindwings increased butterfly survival and successful oviposition during predation encounters. In previous studies, DS B. anynana experienced delayed detection by vertebrate predators, but both forms suffered low survival once detected. Our results suggest predator abundance, identity and phenology may all be important selective forces for B. anynana. Thus, reciprocal selection between invertebrate and vertebrate predators across seasons may contribute to the evolution of the B. anynana polyphenism.
Project description:Microtine species in Fennoscandia display a distinct north-south gradient from regular cycles to stable populations. The gradient has often been attributed to changes in the interactions between microtines and their predators. Although the spatial structure of the environment is known to influence predator-prey dynamics of a wide range of species, it has scarcely been considered in relation to the Fennoscandian gradient. Furthermore, the length of microtine breeding season also displays a north-south gradient. However, little consideration has been given to its role in shaping or generating population cycles. Because these factors covary along the gradient it is difficult to distinguish their effects experimentally in the field. The distinction is here attempted using realistic agent-based modelling.By using a spatially explicit computer simulation model based on behavioural and ecological data from the field vole (Microtus agrestis), we generated a number of repeated time series of vole densities whose mean population size and amplitude were measured. Subsequently, these time series were subjected to statistical autoregressive modelling, to investigate the effects on vole population dynamics of making predators more specialised, of altering the breeding season, and increasing the level of habitat fragmentation. We found that fragmentation as well as the presence of specialist predators are necessary for the occurrence of population cycles. Habitat fragmentation and predator assembly jointly determined cycle length and amplitude. Length of vole breeding season had little impact on the oscillations.There is good agreement between our results and the experimental work from Fennoscandia, but our results allow distinction of causation that is hard to unravel in field experiments. We hope our results will help understand the reasons for cycle gradients observed in other areas. Our results clearly demonstrate the importance of landscape fragmentation for population cycling and we recommend that the degree of fragmentation be more fully considered in future analyses of vole dynamics.
Project description:Allee effects may render exploited animal populations extinction prone, but empirical data are often lacking to describe the circumstances leading to an Allee effect. Arbitrary assumptions regarding Allee effects could lead to erroneous management decisions so that predictive modelling approaches are needed that identify the circumstances leading to an Allee effect before such a scenario occurs. We present a predictive approach of Allee effects for polar bears where low population densities, an unpredictable habitat and harvest-depleted male populations result in infrequent mating encounters. We develop a mechanistic model for the polar bear mating system that predicts the proportion of fertilized females at the end of the mating season given population density and operational sex ratio. The model is parametrized using pairing data from Lancaster Sound, Canada, and describes the observed pairing dynamics well. Female mating success is shown to be a nonlinear function of the operational sex ratio, so that a sudden and rapid reproductive collapse could occur if males are severely depleted. The operational sex ratio where an Allee effect is expected is dependent on population density. We focus on the prediction of Allee effects in polar bears but our approach is also applicable to other species.
Project description:Many populations collapse suddenly when reaching low densities even if they have abundant food conditions, a phenomenon known as an Allee effect. Such collapses can have disastrous consequences, for example, for loss of biodiversity. In this paper, we formulate a stage-structured consumer-resource biomass model in which adults only reproduce at the beginning of each growing season, and investigate the effect of an increasing stage-independent background mortality rate of the consumer. As the main difference with previously studied continuous-time models, seasonal reproduction can result in an Allee effect and consumer population collapses at high consumer mortality rate. However, unlike the mechanisms reported in the literature, in our model the Allee effect results from the time difference between the maturation of juveniles and the reproduction of adults. The timing of maturation plays a crucial role because it not only determines the body size of the individuals at maturation but also influences the duration of the period during which adults can invest in reproductive energy, which together determine the reproductive output at the end of the season. We suggest that there exists an optimal timing of maturation and that consumer persistence is promoted if individuals mature later in the season at a larger body size, rather than maturing early, despite high food availability supporting rapid growth.
Project description:Theoretical investigations of competitive dynamics have noted that numbers of predator and prey influence each other. However, few empirical studies have demonstrated how a life-history trait of the prey (such as fecundity) can be affected simultaneously by its own density and the density of predators. For instance, density dependence can reduce fecundity with increasing number of prey, while inverse density dependence or Allee effects may occur especially when the prey is a social organism. Here we analysed an intraguild predator-prey system of two seabird species at a large spatio-temporal scale. As expected, we found that fecundity of prey was negatively affected by predator density. Nevertheless, fecundity of prey also increased nonlinearly with its own density and strikingly with the prey-predator ratio. Small groups of prey were probably not able to defend their nests especially against large number of predators. At the highest prey densities (i.e. when anti-predator strategies should be most efficient), prey fecundity also lowered, suggesting the appearance of density dependence mediated by food competition. Allee effects and density dependence occurred across a broad range of population sizes of both the prey and the predator at several local populations facing different ecological environments.
Project description:Non-consumptive fear effects are an important determinant of foraging decisions by consumers across a range of ecosystems. However, how fear effects associated with the presence of predators interact with those associated with habitat structure remain unclear. Here, we used predator fish models (<i>Plectropomus leopardus</i>) and experimental patches of the macroalga <i>Sargassum ilicifolium</i> of varying densities to investigate how predator- and habitat-associated fear effects influence herbivory on coral reefs. We found the removal of macroalgal biomass (i.e. herbivory) was shaped by the interaction between predator- and habitat-associated fear effects. Rates of macroalgal removal declined with increasing macroalgal density, likely due to increased visual occlusion by denser macroalgae patches and reduced ability of herbivorous fishes to detect the predators. The presence of the predator model reduced herbivory within low macroalgal density plots, but not within medium- and high-density macroalgal plots. Our results suggest that fear effects due to predator presence were greatest at low macroalgal density, yet these effects were lost at higher densities possibly due to greater predation risk associated with habitat structure and/or the inability of herbivorous fishes to detect the predator model.
Project description:The fruit fly, Drosophila melanogaster, has proven to be an excellent model organism for genetic, genomic and neurobiological studies. However, relatively little is known about the natural history of D. melanogaster. In particular, neither the natural predators faced by wild populations of D. melanogaster, nor the anti-predatory behaviors they may employ to escape and avoid their enemies have been documented. Here we observe and describe the influence of two predators that differ in their mode of hunting: zebra jumping spiders, Salticus scenicus (active hunters) and Chinese praying mantids, Tenodera sinensis (ambush predators) on the behavioral repertoire of Drosophila melanogaster. We documented three particularly interesting behaviors: abdominal lifting, stopping, and retreat-which were performed at higher frequency by D. melanogaster in the presence of predators. While mantids had only a modest influence on the locomotory activity of D. melanogaster, we observed a significant increase in the overall activity of D. melanogaster in the presence of jumping spiders. Finally, we observed considerable among-individual behavioral variation in response to both predators.
Project description:Commercial fisheries may impact marine ecosystems and affect populations of predators like seabirds. In the Southern Ocean, there is an extensive fishery for Antarctic krill Euphausia superba that is projected to increase further. Comparing distribution and prey selection of fishing operations versus predators is needed to predict fishery-related impacts on krill-dependent predators. In this context, it is important to consider not only predators breeding near the fishing grounds but also the ones breeding far away and that disperse during the non-breeding season where they may interact with fisheries. In this study, we first quantified the overlap between the distribution of the Antarctic krill fisheries and the distribution of a krill dependent seabird, the Antarctic petrel Thalassoica antarctica, during both the breeding and non-breeding season. We tracked birds from the world biggest Antarctic petrel colony (Svarthamaren, Dronning Maud Land), located >1000 km from the main fishing areas, during three consecutive seasons. The overall spatial overlap between krill fisheries and Antarctic petrels was limited but varied greatly among and within years, and was high in some periods during the non-breeding season. In a second step, we described the length frequency distribution of Antarctic krill consumed by Antarctic petrels, and compared this with results from fisheries, as well as from diet studies in other krill predators. Krill taken by Antarctic petrels did not differ in size from that taken by trawls or from krill taken by most Antarctic krill predators. Selectivity for specific Antarctic krill stages seems generally low in Antarctic predators. Overall, our results show that competition between Antarctic petrels and krill fisheries is currently likely negligible. However, if krill fisheries are to increase in the future, competition with the Antarctic petrel may occur, even with birds breeding thousands of kilometers away.
Project description:Polar bears (Ursus maritimus) from East Greenland and Svalbard exhibited very high concentrations of polychlorinated biphenyls (PCBs) in the 1980s and 1990s. In Svalbard, slow population growth during that period was suspected to be linked to PCB contamination. In this case study, we explored how PCBs could have impacted polar bear population growth and/or male reproductive success in Svalbard during the mid-1990s by reducing the fertility of contaminated males. A dose-response relationship linking the effects of PCBs to male polar bear fertility was extrapolated from studies of the effects of PCBs on sperm quality in rodents. Based on this relationship, an individual-based model of bear interactions during the breeding season predicted fertilization success under alternative assumptions regarding male-male competition for females. Contamination reduced pregnancy rates by decreasing the availability of fertile males, thus triggering a mate-finding Allee effect, particularly when male-male competition for females was limited or when infertile males were able to compete with fertile males for females. Comparisons of our model predictions on age-dependent reproductive success of males with published empirical observations revealed that the low representation of 10-14-year-old males among breeding males documented in Svalbard in mid-1990s could have resulted from PCB contamination. We conclude that contamination-related male infertility may lead to a reduction in population growth via an Allee effect. The magnitude of the effect is largely dependent on the population-specific mating system. In eco-toxicological risk assessments, appropriate consideration should therefore be given to negative effects of contaminants on male fertility and male mating behaviour.