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Martijn Egas

Publications and source records attributed to Martijn Egas.

10 recordsLinked to original sources

Evolutionary predictions should be based on individual-level traits.

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.

Animals↗

Pheromone-induced priming of a defensive response in Western flower thrips.

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.

Animals↗

Diapause incidence in the two-spotted spider mite increases due to predator presence, not due to selective predation.

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.

Adaptation, Physiological↗

Evolution of specialization and ecological character displacement of herbivores along a gradient of plant quality.

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.

Adaptation, Biological↗

Evolution restricts the coexistence of specialists and generalists: the role of trade-off structure.

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.

Adaptation, Physiological↗

Predation risk affects diapause induction in the spider mite Tetranychus urticae.

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.

Adaptation, Physiological↗

Dispersal and the evolution of specialisation in a two-habitat type metapopulation.

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.

Adaptation, Physiological↗

Intraspecific variation in induction of feeding preference and performance in a herbivorous mite.

Induction of food preference has often been observed in herbivorous insects. The term is used to indicate preference of individuals for the host plant they have already experienced over one they have not experienced. A typical set-up is one where individuals first feed on host X or Y, and are then offered a choice between host X and Y. This set-up--and hence the body of empirical data--has been criticised for lack of a control treatment to untangle the effects of the separate hosts. In this study, we use a design with a third, unrelated host as control to investigate induction of preference in the herbivorous arthropod, Tetranychus urticae. We provide evidence of induced preference, as well as induced performance, and show that there is considerable variation in these two traits among strains. We suggest induced resistance to toxic secondary plant chemicals as one potential explanation for induced performance. This in itself suggests associative learning as the most likely candidate learning mechanism for induction of preference in this species. Phenotypically plastic effects underlying induced performance may be a general aspect of induction of preference in herbivorous arthropods, which warrants closer attention to these phenomena.

Animals↗

Adaptive learning in arthropods: spider mites learn to distinguish food quality.

Many herbivorous arthropods have been shown to possess learning capabilities, yet fitness effects of learning are largely unknown. In this paper, we test whether two-spotted spider mites (Tetranychus urticae) learn to distinguish food quality in choice tests, and whether this results in fitness benefits. Food consisted of cucumber plants with one of three degrees of feeding damage: undamaged (no mites), mildly damaged (infested by a mite strain adapted to tomato) and heavily damaged (infested by a mite strain adapted to cucumber). Mites were subjected to one choice test in a greenhouse and three sequential choice tests on leaf disks. Thereafter, individual mite performance was measured as oviposition rate over four days. In the course of the three small-scale choice tests, preference shifted towards less damaged food. The performance tests showed that learning was adaptive: mites learned to prefer the food type that yielded the higher oviposition rate. Interestingly, innate preferences in the greenhouse tests were close to those shown after learning in the small-scale tests. Given that both strains of mites had not experienced cucumber for several years, we hypothesize that the preference in the greenhouse was due to avoidance of mite odours rather than odours of damaged plants. Through its effect on foraging behaviour, adaptive learning may promote the evolution of host plant specialization in herbivorous arthropods.

Adaptation, Physiological↗

On the evolution of cytoplasmic incompatibility in haplodiploid species.

The most enigmatic sexual manipulation by Wolbachia endosymbionts is cytoplasmic incompatibility (CI): infected males are reproductively incompatible with uninfected females. In this paper, we extend the theory on population dynamics and evolution of CI, with emphasis on haplodiploid species. First, we focus on the problem of the threshold to invasion of the Wolbachia infection in a population. Simulations of the dynamics of infection in small populations show that it does not suffice to assume invasion by drift alone (or demographic "accident"). We propose several promising alternatives that may facilitate invasion of Wolbachia in uninfected populations: sex-ratio effects, meta population structure, and other fitness-compensating effects. Including sex-ratio effects of Wolbachia allows invasion whenever infected females produce more infected daughters than uninfected females produce uninfected daughters. Several studies on haplodiploid species suggest the presence of such sex-ratio effects. The simple metapopulation model we analyzed predicts that, given that infecteds are better "invaders," uninfecteds must be better "colonizers" to maintain coexistence of infected and uninfected patches. This condition seems more feasible for species that suffer local extinction due to predation (or parasitization) than for species that suffer local extinction due to overexploiting their resource(s). Finally, we analyze the evolution of CI in haplodiploids once a population has been infected. Evolution does not depend on the type of CI (female mortality or male production), but hinges solely on decreasing the fitness cost and/or increasing the transmission efficiency. Our models offer new perspectives for increasing our understanding of the population and evolutionary dynamics of CI.

Bacterial Infections↗