[show abstract][hide abstract] ABSTRACT: Why humans are prone to cooperate puzzles biologists, psychologists and economists alike. Between-group conflict has been hypothesized to drive within-group cooperation. However, such conflicts did not have lasting effects in laboratory experiments, because they were about luxury goods, not needed for survival ("looting"). Here, we find within-group cooperation to last when between-group conflict is implemented as "all-out war" (eliminating the weakest groups). Human subjects invested in helping group members to avoid having the lowest collective pay-off, whereas they failed to cooperate in control treatments with random group elimination or with no subdivision in groups. When the game was repeated, experience was found to promote helping. Thus, not within-group interactions alone, not random group elimination, but pay-off-dependent group elimination was found to drive within-group cooperation in our experiment. We suggest that some forms of human cooperation are maintained by multi-level selection: reciprocity within groups and lethal competition among groups acting together.
[show abstract][hide abstract] ABSTRACT: Severe intraspecific competition for mates selects for aggressive individuals but may also lead to the evolution of alternative phenotypes that do not act aggressively, yet manage to acquire matings. The two-spotted spider mite, Tetranychus urticae, shows male mate-guarding behaviour and male-male combat for available females. This may provide opportunity for weaker males to avoid fighting by adopting alternative mating behaviour such as sneaker or satellite tactics as observed in other animals. We investigated male precopulatory behaviour in the two-spotted spider mite by means of video-techniques and found three types of male mating behaviour: territorial, sneaker and opportunistic. Territorial and sneaker males associate with female teleiochrysales and spend much time guarding them. Territorial males are easily disturbed by rival males and engage themselves in fights with them. However, sneaker males are not at all disturbed by rival males, never engage in fights and, strikingly, never face attack by territorial males. Opportunistic males wander around in search of females that are in the teleiochrysalis stage but very close to or at emergence. To quickly classify any given mate-guarding male as territorial or sneaker we developed a method based on the instantaneous response of males to disturbance by a live male mounted on top of a brush. We tested this method against the response of the same males to natural disturbance by two or three other males. Because this method proved to be successful, we used it to collect territorial and sneaker males, and subjected them to morphological analysis to assess whether the various behavioural phenotypes are associated with different morphological characters. However, we found no statistical differences between territorial and sneaker males, concerning the length of the first legs, the stylets, the pedipalps or the body.
Experimental and Applied Acarology 02/2013; · 1.39 Impact Factor
[show abstract][hide abstract] ABSTRACT: It is often suggested that any group selection model can be recast in terms of inclusive fitness. A standard reference to support that claim is "'Quantitative genetics, inclusive fitness, and group selection" by Queller (1992) in the American Naturalist 139 (3), 540-558. In that paper the Price equation is used for the derivation of this claim. Instead of a general derivation, we try out a simple model. For this simple example, we find that the result does not hold. The non-equivalence of group selection and kin selection is therefore not only an important finding in itself, but also a case where the use of the Price equation leads to a claim that is not correct. If results that are arrived at with the Price equation are not correct, they can typically be repaired by adding extra assumptions, or explicitly stating implicit ones. We give examples with relatively mild and with less mild extra assumptions. We also discuss why the Price equation is often referred to as dynamically insufficient, and we try to find out what Price's theorem could be.
Journal of Theoretical Biology 08/2011; 299:64-80. · 2.35 Impact Factor
[show abstract][hide abstract] ABSTRACT: This article considers the evolution of behavioral traits that have both a genetic basis and can be modified. The modifications considered concern an adaptive (learnt) response to environmental influences. In theory, the evolution of such traits may lead to two extreme outcomes; one where the trait becomes genetically fixed (and phenotypically invariable) and the other where it is entirely shaped by environmental influences. Between these extremes lies a spectrum of traits containing a genetic component, but also, to differing degrees, a modifiable component. We review theory considering how learning may affect the genetic evolution of a behavioral trait (commonly referred to as the Baldwin effect). We claim behavioral interactions in tritrophic systems, such as the responses of natural enemies to herbivore-induced plant volatiles, provide an excellent model system to study the Baldwin effect and we illustrate this with recent findings on the searching behavior of the predatory mite Phytoseiulus persimilis.
Journal of Plant Interactions 01/2011; 6:77-80. · 0.90 Impact Factor
[show abstract][hide abstract] ABSTRACT: Nutrient limitation determines the primary production and species composition of many ecosystems. Here we apply an adaptive dynamics approach to investigate evolution of the ecological stoichiometry of primary producers and its implications for plant-herbivore interactions. The model predicts a trade-off between the competitive ability and grazing susceptibility of primary producers, driven by changes in their nutrient uptake rates. High nutrient uptake rates enhance the competitiveness of primary producers but also increase their nutritional quality for herbivores. This trade-off enables coexistence of nutrient exploiters and grazing avoiders. If herbivores are not selective, evolution favors runaway selection toward high nutrient uptake rates of the primary producers. However, if herbivores select nutritious food, the model predicts an evolutionarily stable strategy with lower nutrient uptake rates. When the model is parameterized for phytoplankton and zooplankton, the evolutionary dynamics result in plant-herbivore oscillations at ecological timescales, especially in environments with high nutrient availability and low selectivity of the herbivores. High herbivore selectivity stabilizes the community dynamics. These model predictions show that evolution permits nonequilibrium dynamics in plant-herbivore communities and shed new light on the evolutionary forces that shape the ecological stoichiometry of primary producers.
The American Naturalist 10/2010; 176(6):E162-76. · 4.55 Impact Factor
[show abstract][hide abstract] ABSTRACT: Despite the directional selection acting on life-history traits, substantial amounts of standing variation for these traits have frequently been found. This variation may result from balancing selection (e.g., through genetic trade-offs) or from mutation-selection balance. These mechanisms affect allele frequencies in different ways: Under balancing selection alleles are maintained at intermediate frequencies, whereas under mutation-selection balance variation is generated by deleterious mutations and removed by directional selection, which leads to asymmetry in the distribution of allele frequencies. To investigate the importance of these two mechanisms in maintaining heritable variation in oviposition rate of the two-spotted spider mite, we analyzed the response to artificial selection. In three replicate experiments, we selected for higher and lower oviposition rate, compared to control lines. A response to selection only occurred in the downward direction. Selection for lower oviposition rate did not lead to an increase in any other component of fitness, but led to a decline in female juvenile survival. The results suggest standing variation for oviposition rate in this population consists largely of deleterious alleles, as in a mutation-selection balance. Consequently, the standing variation for this trait does not appear to be indicative of its adaptive potential.
[show abstract][hide abstract] ABSTRACT: Plants infested with herbivores release specific volatile compounds that are known to recruit natural enemies. The response of natural enemies to these volatiles may be either learned or genetically determined. We asked whether there is genetic variation in the response of the predatory mite Phytoseiulus persimilis to methyl salicylate (MeSa). MeSa is a volatile compound consistently produced by plants being attacked by the two-spotted spider mite, the prey of P. persimilis. We predicted that predators express genetically determined responses during long-distance migration where previously learned associations may have less value. Additionally, we asked whether these responses depend on odors from uninfested plants as a background to MeSa. To infer a genetic basis, we analyzed the variation in response to MeSa among iso-female lines of P. persimilis by using choice-tests that involved either (1) MeSa presented as a single compound or (2) MeSa with background-odor from uninfested lima bean plants. These tests were conducted for starved and satiated predators, i.e., two physiological states, one that approximates migration and another that mimics local patch exploration. We found variation among iso-female lines in the responses to MeSa, thus showing genetic variation for this behavior. The variation was more pronounced in the starved predators, thus indicating that P. persimilis relies on innate preferences when migrating. Background volatiles of uninfested plants changed the predators' responses to MeSa in a manner that depended on physiological state and iso-female line. Thus, it is possible to select for context-dependent behavioral responses of natural enemies to plant volatiles.
Journal of Chemical Ecology 07/2010; 36(7):680-8. · 2.46 Impact Factor
[show abstract][hide abstract] ABSTRACT: Although phenotypic plasticity can be advantageous in fluctuating environments, it may come too late if the environment changes fast. Complementary chromatic adaptation is a colorful form of phenotypic plasticity, where cyanobacteria tune their pigmentation to the prevailing light spectrum. Here, we study the timescale of chromatic adaptation and its impact on competition among phytoplankton species exposed to fluctuating light colors. We parameterized a resource competition model using monoculture experiments with green and red picocyanobacteria and the cyanobacterium Pseudanabaena, which can change its color within approximately 7 days by chromatic adaptation. The model predictions were tested in competition experiments, where the incident light color switched between red and green at different frequencies (slow, intermediate, and fast). Pseudanabaena (the flexible phenotype) competitively excluded the green and red picocyanobacteria in all competition experiments. Strikingly, the rate of competitive exclusion was much faster when the flexible phenotype had sufficient time to fully adjust its pigmentation. Thus, the flexible phenotype benefited from its phenotypic plasticity if fluctuations in light color were relatively slow, corresponding to slow mixing processes or infrequent storms in their natural habitat. This shows that the timescale of phenotypic plasticity plays a key role during species interactions in fluctuating environments.
The American Naturalist 11/2008; 172(5):169-85. · 4.55 Impact Factor
[show abstract][hide abstract] ABSTRACT: Diapause in arthropods is a physiological state of dormancy that is generally thought to promote survival during harsh seasons and dispersal, but it may also serve to avoid predation in space and time. Here, we show that predation-related odours induce diapause in female adult spider mites. We argue that this response allows them to move into an area where they are free of enemies, yet forced to survive without food. Spider mites are specialised leaf feeders, but--in late summer--they experience severe predation on leaves. Hence, they face a dilemma: to stay on the leaf and risk being eaten or to move away from the leaf and risk death from starvation and thirst. Female two-spotted spider mites solve this dilemma by dramatically changing their physiology when exposed to predation-associated cues. This allows them to disperse away from leaves and to survive in winter refuges in the bark of trees or in the soil. We conclude that the mere presence of predation-associated cues causes some herbivorous mites to seek refuge, thereby retarding the growth rate of the population as a whole: a trait-mediated indirect effect that may have consequences for the stability of predator-prey systems and for ecosystem structure.
[show abstract][hide abstract] ABSTRACT: Explaining the evolution and maintenance of cooperation among unrelated individuals is one of the fundamental problems in biology and the social sciences. Recent findings suggest that altruistic punishment is an important mechanism maintaining cooperation among humans. We experimentally explore the boundaries of altruistic punishment to maintain cooperation by varying both the cost and the impact of punishment, using an exceptionally extensive subject pool. Our results show that cooperation is only maintained if conditions for altruistic punishment are relatively favourable: low cost for the punisher and high impact on the punished. Our results indicate that punishment is strongly governed by its cost-to-impact ratio and that its effect on cooperation can be pinned down to one single variable: the threshold level of free-riding that goes unpunished. Additionally, actual pay-offs are the lowest when altruistic punishment maintains cooperation, because the pay-off destroyed through punishment exceeds the gains from increased cooperation. Our results are consistent with the interpretation that punishment decisions come from an amalgam of emotional response and cognitive cost-impact analysis and suggest that altruistic punishment alone can hardly maintain cooperation under multi-level natural selection. Uncovering the workings of altruistic punishment as has been done here is important because it helps predicting under which conditions altruistic punishment is expected to maintain cooperation.
Proceedings of the Royal Society B: Biological Sciences 05/2008; 275(1637):871-8. · 5.68 Impact Factor
[show abstract][hide abstract] ABSTRACT: Recent theoretical studies have analyzed the evolution of habitat specialization using either the logistic or the Ricker equation. These studies have implemented evolutionary change directly in population-level parameters such as habitat-specific intrinsic growth rates r or carrying capacities K. This approach is a shortcut to a more detailed analysis where evolutionary change is studied in underlying morphological, physiological, or behavioral traits at the level of the individual that contribute to r or K. Here we describe two pitfalls that can occur when such a shortcut is employed. First, population-level parameters that appear as independent variables in a population dynamical model might not be independent when derived from processes at the individual level. Second, patterns of covariation between individual-level traits are usually not conserved when mapped to the level of demographic parameters. Nonlinear mappings constrain the curvature of trade-offs that can sensibly be assumed at the population level. To illustrate these results, we derive a two-habitat version of the logistic and Ricker equations from individual-level processes and compare the evolutionary dynamics of habitat-specific carrying capacities with those of underlying individual-level traits contributing to the carrying capacities. Finally, we sketch how our viewpoint affects the results of earlier studies.
The American Naturalist 12/2006; 168(5):E148-62. · 4.55 Impact Factor
[show abstract][hide abstract] ABSTRACT: The Western flower thrips Frankliniella occidentalis produces conspicuous anal droplets that function as a direct defense against various predators. These droplets also function in pheromonal communication in that they contain a mixture of decyl acetate and dodecyl acetate, which acts as an alarm. Exposure of thrips to synthetic pheromone is known to promote takeoff or refuge seeking, but the effect of the natural pheromone has not yet been studied. Here, we not only studied the response to natural pheromone, but also tested the new hypothesis that the alarm pheromone primes a defensive response in thrips. This test was carried out by measuring the reaction time to a simulated predator attack after exposure to synthetic or natural alarm pheromone (against a control with no pheromone at all). The reaction was quantified in terms of the time it takes a thrips larva to produce a droplet after attack. We found that thrips larvae produce droplets of alarm pheromone faster when cues associated with danger are present. There were no significant differences in reaction times of responses to synthetic pheromone, natural pheromone, or odors from a patch with a predator attacking a thrips larva. This implies that the synthetic pheromone mimics the natural pheromone, and that other cues emanating from the predator play a minor role. We conclude that the alarm pheromone increases the vigilance of the thrips, and this may promote its survival.
Journal of Chemical Ecology 08/2006; 32(7):1599-603. · 2.46 Impact Factor
[show abstract][hide abstract] ABSTRACT: Explaining the evolution and maintenance of cooperation among unrelated individuals is one of the fundamental problems in biology and the social sciences. Recent experimental evidence suggests that altruistic punishment is an important mechanism to maintain cooperation among humans. In this paper we explore the boundary conditions for altruistic punishment to maintain cooperation by systematically varying the cost and impact of punishment, using a subject pool which extends beyond the standard student population. We find that the economics of altruistic punishment lead to the demise of cooperation when punishment is relatively expensive and/or has low impact. Our results indicate that the 'decision to punish' comes from an amalgam of emotional response and cognitive cost-benefit analysis. Additionally, earnings are lowest when punishment promotes cooperation, suggesting that the scope for altruistic punishment as a means to maintain cooperation is limited.
[show abstract][hide abstract] ABSTRACT: We study the combined evolutionary dynamics of herbivore specialization and ecological character displacement, taking into account foraging behavior of the herbivores, and a quality gradient of plant types. Herbivores can adapt by changing two adaptive traits: their level of specialization in feeding efficiency and their point of maximum feeding efficiency along the plant gradient. The number of herbivore phenotypes, their levels of specialization, and the amount of character displacement among them are the result of the evolutionary dynamics, which is driven by the underlying population dynamics, which in turn is driven by the underlying foraging behavior. Our analysis demonstrates broad conditions for the diversification of a herbivore population into many specialized phenotypes, for basically any foraging behavior focusing use on highest gains while also including errors. Our model predicts two characteristic phases in the adaptation of herbivore phenotypes: a fast character-displacement phase and a slow coevolutionary niche-shift phase. This two-phase pattern is expected to be of wide relevance in various consumer-resource systems. Bringing together ecological character displacement and the evolution of specialization in a single model, our study suggests that the foraging behavior of herbivorous arthropods is a key factor promoting specialist radiation.
[show abstract][hide abstract] ABSTRACT: We recently reported evidence for increased diapause incidence in the spider mite Tetranychus urticae in presence of the predatory mite Typhlodromus pyri. This effect may arise from (1) selective predation on non-diapause spider mites, (2) predator-induced diapause in spider mites, or (3) both. Using a different strain of T. urticae, we first recovered increased diapause incidence in association with predators. Then, we tested for selective feeding in two-choice experiments with equal numbers of non-diapause and diapause spider mites. We found that the predatory mite had a significant preference for the latter. This indicates that increased diapause incidence in association with predatory mites is not due to selective predation. Therefore, predator-mediated physiological induction of diapause seems a more likely explanation. The cues leading to induction appear to relate to the predators, not their effects, since predation simulated by spider-mite removal or puncturing did not significantly affect diapause incidence. Why spider mites benefit from this response, remains an open question.
Enperimental and Applied Acarology 02/2005; 35(1-2):73-81. · 1.85 Impact Factor
[show abstract][hide abstract] ABSTRACT: Metapopulation theory for the evolution of specialisation is virtually absent. In this article, therefore, we study a metapopulation model for consumers with a fitness trade-off between two habitats. We focus on effects of habitat abundance, dispersal rate and trade-off strength on the evolution of specialisation under two types of trade-off. Adaptation affects either the intrinsic growth rates r or the carrying capacities K. Depending on dispersal rate and trade-off strength, evolution can result in one generalist, one specialist or two specialist types. Higher dispersal rate and a weaker trade-off favour the evolution of a generalist, for both trade-off structures. However, we also find differences between the two trade-off structures. Our results are qualitatively similar to analyses of two-patch models, suggesting that insights from such simpler models can be extrapolated to metapopulation models. Additional effects, however, occur because in classical metapopulations patch lifetime depends on extinction rate. Counterintuitively, this favours the evolution of specialisation when the trade-off affects r.
Theoretical Population Biology 12/2004; 66(3):233-48. · 1.24 Impact Factor
[show abstract][hide abstract] ABSTRACT: Environmental variability and adaptive foraging behavior have been shown to favor coexistence of specialists and generalists on an ecological timescale. This leaves unaddressed the question of whether such coexistence can also be expected on an evolutionary timescale. In this article, we study the attainability, through gradual evolution, of specialist-generalist coexistence, as well as the evolutionary stability of such communities when allowing for immigration. Our analysis shows that the potential for specialist-generalist coexistence is much more restricted than originally thought and strongly depends on the trade-off structure assumed. We establish that ecological coexistence is less likely for species facing a trade-off between per capita reproduction in different habitats than when the trade-off acts on carrying capacities alone. We also demonstrate that coexistence is evolutionarily stable whenever it is ecologically stable but that in most cases, such coexistence cannot be reached through gradual evolution. We conclude that an evolutionarily stable community of specialists and generalists may be created only through immigration from elsewhere or through mutations of large effect. Our results highlight that trade-offs in fitness-determining traits can have counterintuitive effects on the evolution of specialization.
The American Naturalist 05/2004; 163(4):518-31. · 4.55 Impact Factor
[show abstract][hide abstract] ABSTRACT: Whenever diapause induction triggers movement into another microhabitat or the development of protective morphological structures, this may also alter predation risk. If the risk of being eaten is lower in the diapause phase, then there may be selection favouring diapause induction in response to predators or their cues. In this article, we studied the effect of the predatory mite Typhlodromus pyri on diapause induction in the spider mite Tetranychus urticae. We used a Greek strain because under long-night photoperiods and low temperature only part of the population enters diapause, thereby leaving room for the impact of another factor. In spider mite groups under predation, the percentage diapause induction increased whenever night-lengths were such that diapause was induced (13-16 h of night). Given this diapause induction in response to predation risk, the question arises whether entering diapause helps spider mites to escape from predation and contribute more offspring to the spring generation next year.
Enperimental and Applied Acarology 02/2004; 34(3-4):307-14. · 1.85 Impact Factor