Coevolution by different functional mechanisms modulates the structure and dynamics of antagonistic and mutualistic networks

A central problem in the study of species interactions is to understand the underlying ecological and evolutionary mechanisms that shape and are shaped by trait evolution in interacting assemblages. The patterns of interaction among species (i.e. network structure) provide the pathways for evolution and coevolution, which are modulated by how traits affect individual fitness (i.e. functional mechanisms). Functional mechanisms, in turn, also affect the likelihood of an ecological interaction, shaping the structure of interaction networks. Here, we build adaptive network models to explore the potential role of coevolution by two functional mechanisms, trait matching and exploitation barrier, in driving trait evolution and the structure of interaction networks. We use these models to explore how different scenarios of coevolution and functional mechanisms reproduce the empirical network patterns observed in antagonistic and mutualistic interactions and affect trait evolution. Scenarios assuming coevolutionary feedback with a strong effect of functional mechanism better reproduce the empirical structure of networks. Antagonistic and mutualistic networks, however, are better explained by different functional mechanisms and the structure of antagonisms is better reproduced than that of mutualisms. Scenarios assuming coevolution by strong trait matching between interacting partners better explain the structure of antagonistic networks, whereas those assuming strong barrier effects better reproduce the structure of mutualistic networks. The dynamics resulting from the feedback between strong functional mechanisms and coevolution favor the stability of antagonisms and mutualisms. Selection favoring trait matching reduces temporal trait fluctuation and the magnitude of arms races in antagonisms, whereas selection due to exploitation barriers reduces temporal trait fluctuations in mutualisms. Our results indicate that coevolutionary models better reproduce the network structure of antagonisms than those of mutualisms and that different functional mechanisms may favor the persistence of antagonistic and mutualistic interacting assemblages.     

Costs and constraints in aphid-ant mutualism

While many studies have demonstrated that ants provide beneficial services to aphids, Bristow (Ant-plant interactions, Oxford University Press, Oxford, 104–119, 1991) first questioned why so few aphid species are ant-attended. Phylogenetic trees have demonstrated multiple gains and loss of ant-attendance in the course of aphid-ant interactions, implying that mutualisms easily form and dissolve. Several studies have reported the factors that influence the formation and maintenance of aphid-ant interactions. Examples include the physiological costs of ant attendance, competition for mutualistic ants, ant predation on aphids, the influence of host plants, and parasitoid wasps. Recent physiological techniques have also revealed the chemical component of aphid-ant mutualisms. The honeydew of ant-attended aphids contains melezitose (a trisaccharide), which has an important role in aphid-ant interactions. Studies of cuticular hydrocarbons on aphids and ants have clarified the underlying mechanisms of ant predation on aphids. Attending ants also reduce aphid dispersal ability, causing the formation of fragmented aphid populations with low genetic diversity in each population. The reduced aphid dispersal could be partly explained by higher wing loading and reduction of flight apparatus due to ant attendance. Whether ant attendance is associated with the range of host plants of aphids or genetic variation in microorganism in aphids remain to be explored.     

On the evolution of non-specific mutualism

It has been argued that mutualisms are non-specific when mutualistic interactions are weak and transient, and become more specific as interactions increase in strength. However, this runs counter to the observation that there exist tightly linked mutualisms of great antiquity that are highly nonspecific. Here we argue that mutualism generates positive, interspecific, frequency-dependent selection, which acts as a cohesive evolutionary force, discouraging evolution of specificity. A simple mathematical model is constructed to analyse the evolution of a community consisting of two guilds of species with mutualistic between-guild interactions, two competing species in each guild and two genetically distinct phenotypes within each species. With some simplifying assumptions, the trajectories in the neighbourhood of the only interior equilibrium point are determined analytically in terms of interactions between individuals. These show that the equilibrium is locally stable (no evolution) when there is little differentiation between phenotypes in mutualistic and interspecific, competitive interactions. On the other hand, when there is strong differentiation between phenotypes in their mutualistic interactions, the equilibrium is unstable and the community starts to evolve towards non-specificity. There are, however, two forces counteracting this tendency which, if sufficiently potent, cause evolution towards specificity. The first is generated by strong differentiation between phenotypes in interspecific competition; the second is caused by specificity which already exists between species in their mutualistic interactions. Thus, the tendency for non-specificity or specificity to evolve depends on the interplay between antagonistic and mutualistic interactions in the community. We illustrate these results with some numerical examples and, finally, survey some data on specificity of mutualisms in the light of the analysis.     

Chemical defence,offence and alliance in ants–aphids–ladybirds relationships

Chemicals, which mediate the interactions between aphids, ladybirds and ants, are reviewed. Special emphasis is laid on autogenous and plant-derived chemical defence in aphids and ladybirds. Evidences for chemical cues used during foraging and oviposition in ladybirds are assessed. Possible mutualistic interaction between plants and the third trophic level is illustrated by the as yet few reports of indirect plant-defence volatiles induced by aphids or coccids attracting parasitoids or ladybirds. The use of chemical signals allowing aphid parasitoids or ladybirds to squeeze into ant–aphid mutualistic association is briefly described. Questions are raised and hypotheses suggested which could stimulate further research on aphid host-plant influence on ladybird foraging behaviour and fitness, and on the cues used by aphid-web partners for their mutual recognition.     

Ants indirectly reduce the reproductive performance of a leafless shrub by benefiting aphids through predator deterrence

Ant–aphid mutualisms can generate cascade effects on the host plants, but these impacts depend on the ecological context. We studied the consequences of ant–aphid interactions on the reproductive performance of a Mediterranean leafless shrub (Retama sphaerocarpa), through direct and indirect effects on the arthropod community. By manipulating the presence of ants and aphids in the field, we found that ants increased aphid abundance and their persistence on the plant and reduced aphid predators by nearly half. However, the presence of ants did not affect the abundance of other plant herbivores, which were relatively scarce in the studied plants. Aphids, and particularly those tended by ants, had a negative impact on the plant reproductive performance by significantly reducing the number of fruits produced. However, fruit and seed traits were not changed by the presence of aphids or those tended by ants. We show that ants favoured aphids by protecting them from their natural enemies but did not indirectly benefit plants through herbivory suppression, resulting in a net negative impact on the plant reproductive performance. Our study suggests that the benefits obtained by plants from hosting ant–aphid mutualisms are dependent on the arthropod community and plant traits.

     

Insights into a novel three-partner interaction between ants,coreids (Hemiptera: Coreidae) and extrafloral nectaries: implications for the study of protective mutualisms

Extrafloral nectar of plants and honeydew of hemipterans are the common mediators of facultative interactions that involve ants as a mobile strategy of defence. The outcome of these interactions can vary from mutualistic to commensalistic or even antagonistic, depending on the ecological context and the interacting species. Here, we explore a novel, three-partner interaction involving ants, the coreid Dersagrena subfoveolata (Hemiptera) and the extrafloral nectaries (EFNs) bearing plant Senna aphylla (Fabaceae) in semi-arid Northwest Argentina. We surveyed natural areas and conducted ant exclusion experiments, to understand how each pairwise interaction influences the overall outcome among the three interacting parts. The outcome of the interactions was assessed for experimental plants as the reproductive output and herbivore abundances and for coreids as predator abundances. We found that the coreids occurred exclusively on S. aphylla plants and that at least nine ant species interacted with the EFNs as well as with the coreids. Coreid occurrence and abundance depended on ant densities, which in turn, was determined by the presence of actively secreting EFNs. Coreid and ant presence did not influence plant reproductive success, and ants provided to coreids some biotic defence, mainly against vespid wasp predators, but had no effect on non-coreid herbivores. We conclude that the interaction outcome is commensalistic between ants and plants (assuming that EF nectar is not costly for the plant), antagonistic between coreids and plants, and mutualistic between coreids and ants. The sum of all outcomes is net positive effect for ants and coreids, and net slightly negative to neutral for plants.     

Mutualism, hybrid inviability and speciation in a tropical ant-plant

Although biotic interactions are particularly intricate in the tropics, few studies have examined whether divergent adaptations to biotic interactions lead to speciation in tropical organisms. Ant-plant mutualisms are widespread in the tropics. Within Leonardoxa africana, two subspecies present contrasting defences against herbivores. Young leaves of subsp. africana are defended by mutualistic ants, whereas subsp. gracilicaulis satiates herbivores by synchronized leaf production. Subsp. africana possesses hollow internodes and many large foliar nectaries, housing and feeding ants. We detected no genetic introgression between the two subspecies in the contact zone between them. F1 hybrids were present. They were intermediate in phenotype, expressing reduced, nonfunctional but costly myrmecophilic traits. However, they suffered more herbivory than their parents. Hybrids remained small, failing to reach reproductive size, probably due to their maladapted defence phenotype. Hence, there could be a direct link between adaptation to mutualism and reproductive isolation: biotic interactions could be a driver of tropical diversity.     

Interactions between seed predators/pollinators and their host plants: a first step towards mutualism?

The mutualisms between fig trees and their pollinator fig wasps and between yucca plants and yucca moths are spectacular examples of coevolution. The characteristics of these independently evolved mutualisms have resulted from long‐term processes, the first stages of which are unknown. A fundamental question in the study of mutualism is how these interactions evolve. Seed predator/pollinator and host plant interactions, which may initially be considered as mainly antagonistic, have the potential to provide good model systems for the study of the first stages of evolution towards mutualism. We present here theoretical models assessing the consequences of interactions between specialized seed predator insects and their host plants. These models describe the parameters that affect the fitness of an individual female seed predator and her influence on the fitness of the host plant. In an optimal strategy for the seed predator, the number of eggs laid in each flower depends on the interaction between the adult and larva survival. Along with a growing predation pressure on adults and larvae several eggs must be laid in each flower by the female seed predator to enhance her fitness. However, in a situation where the host plant selectively aborts flowers with a high number of eggs the fitness of the seed predator will seriously decrease. If the cost of selective abortion is less than the cost of seed predation the host plant will maintain fitness. In a mutualistic relationship a balance between the cost and the benefit of the parameters in the fitness models of the seed predator and the host plant has to occur so that the net seed output is larger than zero (0). Any unselfish behaviour or quality of the seed predator that would benefit the host plant in such a way that the net seed output increases might be a first stage in an interaction becoming mutualistic. The models presented here will not only provide a platform for empirical studies on interactions that may swing from parasitism to mutualism, but also for seed predator/pollinator and host plant interactions in general.     

The indirect consequences of a mutualism: comparing positive and negative components of the net interaction between honeydew-tending ants and host plants

1. In ecological webs, net indirect interactions between species are composed of interactions that vary in sign and magnitude. Most studies have focused on negative component interactions (e.g. predation, herbivory) without considering the relative importance of positive interactions (e.g. mutualism, facilitation) for determining net indirect effects. 2. In plant/arthropod communities, ants have multiple top-down effects via mutualisms with honeydew-producing herbivores and harassment of and predation on other herbivores; these ant effects provide opportunities for testing the relative importance of positive and negative interspecific interactions. We manipulated the presence of ants, honeydew-producing membracids and leaf-chewing beetles on perennial host plants in field experiments in Colorado to quantify the relative strength of these different types of interactions and their impact on the ant's net indirect effect on plants. 3. In 2007, we demonstrated that ants simultaneously had a positive effect on membracids and a negative effect on beetles, resulting in less beetle damage on plants hosting the mutualism. 4. In 2008, we used structural equation modelling to describe interaction strengths through the entire insect herbivore community on plants with and without ants. The ant's mutualism with membracids was the sole strong interaction contributing to the net indirect effect of ants on plants. Predation, herbivory and facilitation were weak, and the net effect of ants reduced plant reproduction. This net indirect effect was also partially because of behavioural changes of herbivores in the presence of ants. An additional membracid manipulation showed that the membracid's effect on ant activity was largely responsible for the ant's net effect on plants; ant workers were nearly ten times as abundant on plants with mutualists, and effects on other herbivores were similar to those in the ant manipulation experiment. 5. These results demonstrate that mutualisms can be strong relative to negative direct interspecific interactions and that positive interactions deserve attention as important components of ecological webs.     

Parasites may help stabilize cooperative relationships

Background  

The persistence of cooperative relationships is an evolutionary paradox; selection should favor those individuals that exploit their partners (cheating), resulting in the breakdown of cooperation over evolutionary time. Our current understanding of the evolutionary stability of mutualisms (cooperation between species) is strongly shaped by the view that they are often maintained by partners having mechanisms to avoid or retaliate against exploitation by cheaters. In contrast, we empirically and theoretically examine how additional symbionts, specifically specialized parasites, potentially influence the stability of bipartite mutualistic associations. In our empirical work we focus on the obligate mutualism between fungus-growing ants and the fungi they cultivate for food. This mutualism is exploited by specialized microfungal parasites (genus Escovopsis) that infect the ant's fungal gardens. Using sub-colonies of fungus-growing ants, we investigate the interactions between the fungus garden parasite and cooperative and experimentally-enforced uncooperative ("cheating") pairs of ants and fungi. To further examine if parasites have the potential to help stabilize some mutualisms we conduct Iterative Prisoner's Dilemma (IPD) simulations, a common framework for predicting the outcomes of cooperative/non-cooperative interactions, which incorporate parasitism as an additional factor.     

Today and tomorrow: impact of climate change on aphid biology and potential consequences on their mutualism with ants

Recent studies about mutualism consider the complexity and versatility of the relationship, in addition to highlighting the importance of the cost/benefit balance between the two protagonists. Because species interactions are highly dependent on the environment, the climate changes foreseen for the coming years are expected to have significant impacts on the evolution of mutualistic interactions. Among mutualisms, the aphid–ant interaction is well documented, partly explained by the pest status of aphids. This literature review focuses on the impact of climate change (particularly atmospheric carbon dioxide concentration and temperature) on aphid biology and the potential consequences with respect to their mutualistic interactions with ants. We provide an overview of the published reports concerned with the effects of temperature and carbon dioxide on aphids, for which a positive, a negative or no effect has been highlighted. We then discuss how climatic changes can alter four major components of aphid biology that are shaping their interaction with ants: (i) aphid population growth; (ii) aphid behaviour and mobility; (iii) honeydew production and composition; and (iv) semiochemistry. Finaly, we discuss the limitations of such studies on aphid–ant mutualism, as well as the information that is still needed to predict how climate change might impact this type of relationship.     

Costs and benefits for phytophagous myrmecophiles: when ants are not always available

Costs and benefits in mutualistic associations between ants and phytophagous myrmecophiles are context dependent. We collected information from the literature on costs and benefits of myrmecophily in aphids, coccids, membracids and lycaenids. A key result of the literature survey is that investment in mutualism with ants entails costs paid not only when ants are present (direct costs) but also when they are absent (indirect costs). We incorporated such a trade-off in a model that investigates the fitness consequences of the decision of a potential myrmecophile whether or not to invest in cooperation with ants. The model shows that whether myrmecophily should be favoured depends on the rate of increase of the population, and, if there are indirect costs, on the frequency of habitats with ants. Both direct and indirect costs can limit or prevent the evolution of myrmecophily even when ants are abundant. To understand the patterns of associations in the field we therefore need to measure the benefits and costs of myrmecophily both in the presence and in the absence of ants.     

Do aphids actively search for ant partners?

The aphid–ant mutualistic relationships are not necessarily obligate for neither partners but evidence is that such interactions provide them strong advantages in terms of global fitness. While it is largely assumed that ants actively search for their mutualistic partners namely using volatile cues; whether winged aphids (i.e., aphids’ most mobile form) are able to select ant‐frequented areas had not been investigated so far. Ant‐frequented sites would indeed offer several advantages for these aphids including a lower predation pressure through ant presence and enhanced chances of establishing mutuaslistic interactions with neighbor ant colonies. In the field, aphid colonies are often observed in higher densities around ant nests, which is probably linked to a better survival ensured by ants’ services. Nevertheless, this could also result from a preferential establishment of winged aphids in ant‐frequented areas. We tested this last hypothesis through different ethological assays and show that the facultative myrmecophilous black bean aphid, Aphis fabae L., does not orientate its search for a host plant preferentially toward ant‐frequented plants. However, our results suggest that ants reduce the number of winged aphids leaving the newly colonized plant. Thus, ants involved in facultative myrmecophilous interactions with aphids appear to contribute to structure aphid populations in the field by ensuring a better establishment and survival of newly established colonies rather than by inducing a deliberate plant selection by aphid partners based on the proximity of ant colonies.     

Sex,competition and mimicry: an eco-evolutionary model reveals unexpected impacts of ecological interactions on the evolution of phenotypes in sympatry

Phenotypic evolution in sympatric species can be strongly impacted by species interactions, either mutualistic or antagonistic. Heterospecific reproductive behaviours between sympatric species have been shown to favour phenotypic divergence of traits used as sexual cues. Those traits may also be involved in local adaptation or in other types of species interactions and, as a result, undergo complex evolutions across sympatric species. Here we focus on mimicry and study how reproductive interference may impair phenotypic convergence between species with various levels of defence. We use a deterministic model assuming two sympatric species where individuals can display two different warning colour patterns. This eco-evolutionary model explores how ecological interactions shape phenotypic evolution within sympatric species. We investigate the effect of 1) the opposing density-dependent selections exerted on colour patterns by predation and reproductive behaviour and 2) the impact of relative species and phenotype abundances on the fitness costs faced by each individual depending on their species and phenotype. Our model shows that reproductive interference may limit the convergent effect of mimetic interactions and may promote phenotypic divergence between Müllerian mimics. The divergent and convergent evolution of traits also strongly depends on the relative species and phenotype abundances and levels of trophic competition, highlighting how the eco-evolutionary feedbacks between phenotypic evolution and species abundances may result in strikingly different evolutionary routes.     

Benefits for plants in ant-plant protective mutualisms: a meta-analysis

Costs and benefits for partners in mutualistic interactions can vary greatly, but surprisingly little is known about the factors that drive this variation across systems. We conducted a meta-analysis of ant-plant protective mutualisms to quantify the effects of ant defenders on plant reproductive output, to evaluate if reproductive effects were predicted from reductions in herbivory and to identify characteristics of the plants, ants and environment that explained variation in ant protection. We also compared our approach with two other recent meta-analyses on ant-plant mutualisms, emphasizing differences in our methodology (using a weighted linear mixed effects model) and our focus on plant reproduction rather than herbivore damage. Based on 59 ant and plant species pairs, ant presence increased plant reproductive output by 49% and reduced herbivory by 62%. The effects on herbivory and reproduction within systems were positively correlated, but the slope of this relationship (0.75) indicated that tolerance to foliar herbivory may be a common plant response to absence of ant guards. Furthermore, the relationship between foliar damage and reproduction varied substantially among systems, suggesting that herbivore damage is not a reliable surrogate for fitness consequences of ant protection. Studies that experimentally excluded ants reported a smaller effect of ant protection on plant reproduction than studies that relied upon natural variation in ant presence, suggesting that study methods can affect results in these systems. Of the ecological variables included in our analysis, only plant life history (i.e., annual or perennial) explained variation in the protective benefit of mutualistic ants: presence of ants benefitted reproduction of perennials significantly more than that of annuals. These results contrast with other quantitative reviews of these relationships that did not include plant life history as an explanatory factor and raise several questions to guide future research on ant-plant protection mutualisms.     

Interactions among mutualism,competition, and predation foster species coexistence in diverse communities

In natural systems, organisms are simultaneously engaged in mutualistic, competitive, and predatory interactions. Theory predicts that species persistence and community stability are feasible when the beneficial effects of mutualisms are balanced by density-dependent negative feedbacks. Enemy-mediated negative feedbacks can foster plant species coexistence in diverse communities, but empirical evidence remains mixed. Disparity between theoretical expectations and empirical results may arise from the effects of mutualistic mycorrhizal fungi. Here, we build a multiprey species/predator model combined with a bidirectional resource exchange system, which simulates mutualistic interactions between plants and fungi. To reach population persistence, (1) the per capita rate of increase of all plant population must exceed the sum of the negative per capita effects of predation, interspecific competition, and costs of mycorrhizal association, and (2) the per capita numerical response of enemies to mycorrhizal plants must exceed the magnitude of the per capita enemy rate of mortality. These conditions reflect the balance between regulation and facilitation in the system. Interactions between plant natural enemies and mycorrhizal fungi lead to shifts in the strength and direction of net mycorrhizal effects on plants over time, with common plant species deriving greater benefits from mycorrhizal associations than rare plant species.     

Phytophagous insect-microbe mutualisms and adaptive evolutionary diversification.

Adaptive diversification is a process intrinsically tied to species interactions. Yet, the influence of most types of interspecific interactions on adaptive evolutionary diversification remains poorly understood. In particular, the role of mutualistic interactions in shaping adaptive radiations has been largely unexplored, despite the ubiquity of mutualisms and increasing evidence of their ecological and evolutionary importance. Our aim here is to encourage empirical inquiry into the relationship between mutualism and evolutionary diversification, using herbivorous insects and their microbial mutualists as exemplars. Phytophagous insects have long been used to test theories of evolutionary diversification; moreover, the diversification of a number of phytophagous insect lineages has been linked to mutualisms with microbes. In this perspective, we examine microbial mutualist mediation of ecological opportunity and ecologically based divergent natural selection for their insect hosts. We also explore the conditions and mechanisms by which microbial mutualists may either facilitate or impede adaptive evolutionary diversification. These include effects on the availability of novel host plants or adaptive zones, modifying host-associated fitness trade-offs during host shifts, creating or reducing enemy-free space, and, overall, shaping the evolution of ecological (host plant) specialization. Although the conceptual framework presented here is built on phytophagous insect-microbe mutualisms, many of the processes and predictions are broadly applicable to other mutualisms in which host ecology is altered by mutualistic interactions.     

PHYTOPHAGOUS INSECT–MICROBE MUTUALISMS AND ADAPTIVE EVOLUTIONARY DIVERSIFICATION

Adaptive diversification is a process intrinsically tied to species interactions. Yet, the influence of most types of interspecific interactions on adaptive evolutionary diversification remains poorly understood. In particular, the role of mutualistic interactions in shaping adaptive radiations has been largely unexplored, despite the ubiquity of mutualisms and increasing evidence of their ecological and evolutionary importance. Our aim here is to encourage empirical inquiry into the relationship between mutualism and evolutionary diversification, using herbivorous insects and their microbial mutualists as exemplars. Phytophagous insects have long been used to test theories of evolutionary diversification; moreover, the diversification of a number of phytophagous insect lineages has been linked to mutualisms with microbes. In this perspective, we examine microbial mutualist mediation of ecological opportunity and ecologically based divergent natural selection for their insect hosts. We also explore the conditions and mechanisms by which microbial mutualists may either facilitate or impede adaptive evolutionary diversification. These include effects on the availability of novel host plants or adaptive zones, modifying host-associated fitness trade-offs during host shifts, creating or reducing enemy-free space, and, overall, shaping the evolution of ecological (host plant) specialization. Although the conceptual framework presented here is built on phytophagous insect–microbe mutualisms, many of the processes and predictions are broadly applicable to other mutualisms in which host ecology is altered by mutualistic interactions.     

Bears benefit plants via a cascade with both antagonistic and mutualistic interactions

Predators can influence primary producers by generating cascades of effects in ecological webs. These effects are often non‐intuitive, going undetected because they involve many links and different types of species interactions. Particularly, little is understood about how antagonistic (negative) and mutualistic (positive) interactions combine to create cascades. Here, we show that black bears can benefit plants by consuming ants. The ants are mutualists of herbivores and protect herbivores from other arthropod predators. We found that plants near bear‐damaged ant nests had greater reproduction than those near undamaged nests, due to weaker ant protection for herbivores, which allowed herbivore suppression by arthropod predators. Our results highlight the need to integrate mutualisms into trophic cascade theory, which is based primarily on antagonistic relationships. Predators are often conservation targets, and our results suggest that bears and other predators should be managed with the understanding that they can influence primary producers through many paths.     

Plant killing by Neotropical acacia ants: ecology,decision‐making,and head morphology

Mutualistic species often associate with several partners that vary in the benefits provided. In some protective ant–plant mutualisms, ants vary in the extent at which they kill neighboring vegetation. Particularly, in acacia ants (Pseudomyrmex), the area around the host tree that ants keep free from vegetation (“clearings”) vary depending on the species. This study assessed whether interspecific variation in clearing size corresponds to workers biting on plant tissue of different thickness. As expected, workers from species making the largest clearings bit more often on thicker plant tissues than workers from species making smaller clearings. Because head shape affects mandible force, I also assessed whether pruning on thick tissue in mutualistic ant species or being a predator in non‐mutualistic species correlated with broader heads, which yield stronger mandible force. The species with the broader heads were non‐mutualistic predators or mutualistic pruners of thick tissues, which suggest that pruning neighboring vegetation in non‐predatory species demands force even when the ants do not kill prey with their mandibles. The findings reveal that clearing size variation in mutualistic ant partners of plants can also be observed at the level of individual decision‐making processes among workers, and suggest that head morphology could be a trait under selection in protective ant–plant mutualisms. Abstract in Spanish is available with online material.