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Cooperation is widespread both within and between species, but are intraspecific and interspecific cooperation fundamentally similar or qualitatively different phenomena? This review evaluates this question, necessary for a general understanding of the evolution of cooperation. First, we outline three advantages of cooperation relative to noncooperation (acquisition of otherwise inaccessible goods and services, more efficient acquisition of resources, and buffering against variability), and predict when individuals should cooperate with a conspecific versus a heterospecific partner to obtain these advantages. Second, we highlight five axes along which heterospecific and conspecific partners may differ: relatedness and fitness feedbacks, competition and resource use, resource-generation abilities, relative evolutionary rates, and asymmetric strategy sets and outside options. Along all of these axes, certain asymmetries between partners are more common in, but not exclusive to, cooperation between species, especially complementary resource use and production. We conclude that cooperation within and between species share many fundamental qualities, and that differences between the two systems are explained by the various asymmetries between partners. Consideration of the parallels between intra- and interspecific cooperation facilitates application of well-studied topics in one system to the other, such as direct benefits within species and kin-selected cooperation between species, generating promising directions for future research. [ABSTRACT FROM AUTHOR]

Understanding the conditions under which moderately defended prey evolve to resemble better-defended prey and whether this mimicry is parasitic (quasi-Batesian) or mutualistic (Müllerian) is central to our understanding of warning signals. Models of predator learning generally predict quasi-Batesian relationships. However, predators' attack decisions are based not only on learning alone but also on the potential future rewards.We identify the optimal sampling strategy of predators capable of classifying prey into different profitability categories and contrast the implications of these rules formimicry evolution with a classical Pavlovianmodel based on conditioning. In both cases, the presence of moderately unprofitable mimics causes an increase in overall consumption. However, in the case of the optimal sampling strategy, this increase in consumption is typically outweighed by the increase in overall density of prey sharing the model appearance (a dilution effect), causing a decrease in mortality. It suggests that if predators forage efficiently to maximize their long-term payoff, genuine quasi-Batesianmimicry should be rare, whichmay explain the scarcity of evidence for it in nature. Nevertheless, we show that when moderately defended mimics are profitable to attack by hungry predators, then they can be parasitic on their models, just as classical Batesian mimics are. [ABSTRACT FROM AUTHOR]

Mutualisms are only rarely one-to-one interactions: each species generally interacts with multiple mutualists. Exploitation is ubiquitous in mutualisms, and we would therefore expect that each mutualist interacts with multiple exploiters as well. Exploiter species may also interact with one another. For example, the action of one exploiter species might open the opportunity for exploitation by a second species., Exploitation is common in many plant-pollinator mutualisms: 'primary' nectar robbers feed through holes they make in flowers, which can be subsequently used by 'secondary' nectar robbers unable to create holes themselves. The overall effect of nectar robbing on plant fitness is often (although not always) negative. No study has separated the effects of interacting with primary vs. secondary robbers., Here, we examine the effects of primary vs. secondary nectar robbing on pollinator visitation rate and female fitness in Ipomopsis aggregata. Manipulating the type of nectar robbing that flowers experienced, we found that secondary nectar robbing inflicted fitness costs to plants beyond that inflicted by primary robbing alone. Secondary nectar robbing significantly reduced pollen receipt to flowers, as well as fruit and seed production. Although the causes are elusive, the effect may be attributed to changes in pollinator behaviour at these plants., Synthesis. Our findings provide evidence that interacting with multiple exploiters can lead to increased negative effects for mutualists, and highlight the importance of incorporating multiple exploiters into the conceptual framework of mutualism. [ABSTRACT FROM AUTHOR]

Biologists have taken the concept of organism largely for granted. However, advances in the study of chimerism, symbiosis, bacterial-eukaryote associations, and microbial behavior have prompted a redefinition of organisms as biological entities exhibiting low conflict and high cooperation among their parts. This expanded view identifies organisms in evolutionary time. However, the ecological processes, mechanisms, and traits that drive the formation of organisms remain poorly understood. Recognizing that organismality can be context dependent, we advocate elucidating the ecological contexts under which entities do or do not act as organisms. Here we develop a 'contextual organismality' framework and provide examples of entities, such as honey bee colonies, tumors, and bacterial swarms, that can act as organisms under specific life history, resource, or other ecological circumstances. We suggest that context dependence may be a stepping stone to the development of increased organismal unification, as the most integrated biological entities generally show little context dependence. Recognizing that organismality is contextual can identify common patterns and testable hypotheses across different entities. The contextual organismality framework can illuminate timeless as well as pressing issues in biology, including topics as disparate as cancer emergence, genomic conflict, evolution of symbiosis, and the role of the microbiota in impacting host phenotype. [ABSTRACT FROM AUTHOR]

Exploitation in cooperative interactions both within and between species is widespread. Although it is assumed to be costly to be exploited, mechanisms to control exploitation are surprisingly rare, making the persistence of cooperation a fundamental paradox in evolutionary biology and ecology. Focusing on between-species cooperation (mutualism), we hypothesize that the temporal sequence in which exploitation occurs relative to cooperation affects its net costs and argue that this can help explain when and where control mechanisms are observed in nature. Our principal prediction is that when exploitation occurs late relative to cooperation, there should be little selection to limit its effects (analogous to “tolerated theft” in human cooperative groups). Although we focus on cases in which mutualists and exploiters are different individuals (of the same or different species), our inferences can readily be extended to cases in which individuals exhibit mixed cooperative-exploitative strategies. We demonstrate that temporal structure should be considered alongside spatial structure as an important process affecting the evolution of cooperation. We also provide testable predictions to guide future empirical research on interspecific as well as intraspecific cooperation. [ABSTRACT FROM AUTHOR]

Symbiosis generally causes an expansion of the niche of each partner along the axis for which a service is mutually provided. However, for other axes, the niche can be restricted to the intersection of each partner's niche and can thus be constrained rather than expanded by mutualism.We explore this phenomenon using Artemia as a model system. This crustacean is able to survive at very high salinities but not at low salinities, although its hemolymph's salinity is close to freshwater. We hypothesized that this low-salinity paradox results from poor performance of its associated microbiota at low salinity. We showed that, in sterile conditions, Artemia had low survival at all salinities when algae were the only source of carbon. In contrast, survival was high at all salinities when fed with yeast. We also demonstrated that bacteria isolated from Artemia's gut reached higher densities at high salinities than at low salinities, including when grown on algae. Taken together, our results show that Artemia can survive at low salinities, but their gut microbiota, which are required for algae digestion, have reduced fitness. Widespread facultative symbiosis may thus be an important determinant of niche limits along axes not specific to the mutualistic interaction. [ABSTRACT FROM AUTHOR]

Core principles of ecological theory predict that, in the absence of other factors, mutualisms should experience destabilizing positive feedback and should be vulnerable to extinction through competitive exclusion by exploiter species. Many effective stabilizing mechanisms address one issue or the other, and many turn upon additional features. Using an explicitly demographic approach, I show that indirect, demography-mediated interactions between mutualists and exploiters can enable mutualist-exploiter coexistence, which in turn can stabilize the abundances of mutualists, exploiters, and their shared resources. This occurs because of the distinct resource demographic responses that are inherent to interaction with mutualistic and exploitative partners and can occur in long-lasting, exclusive interactions, such as protection mutualisms, as well as in apparently very different, short-lived mutualistic interactions, such as pollination. The key necessary factor-demographic response to interspecific interaction-is common in nature. Some demographic structure is also necessary and is generated through interspecific interaction in long-lasting associations; it is also very common in natural populations. Thus, the explicitly demographic and multispecies approach taken here constitutes a potentially promising single explanation for the apparent stability ofmutualismin a wide range of natural systems. [ABSTRACT FROM AUTHOR]

Climate change is altering the timing of life history events in a wide array of species, many of which are involved in mutualistic interactions. Because many mutualisms can form only if partner species are able to locate each other in time, differential phenological shifts are likely to influence their strength, duration and outcome. At the extreme, climate change-driven shifts in phenology may result in phenological mismatch: the partial or complete loss of temporal overlap of mutualistic species. We have a growing understanding of how, when, and why phenological change can alter one type of mutualism - pollination. However, as we show here, there has been a surprising lack of attention to other types of mutualism. We generate a set of predictions about the characteristics that may predispose mutualisms in general to phenological mismatches. We focus not on the consequences of such mismatches but rather on the likelihood that mismatches will develop. We explore the influence of three key characteristics of mutualism: 1) intimacy, 2) seasonality and duration, and 3) obligacy and specificity. We predict that the following characteristics of mutualism may increase the likelihood of phenological mismatch: 1) a non-symbiotic life history in which co-dispersal is absent; 2) brief, seasonal interactions; and 3) facultative, generalized interactions. We then review the limited available data in light of our a priori predictions and point to mutualisms that are more and less likely to be at risk of becoming phenologically mismatched, emphasizing the need for research on mutualisms other than plant - pollinator interactions. Future studies should explicitly focus on mutualism characteristics to determine whether and how changing phenologies will affect mutualistic interactions. [ABSTRACT FROM AUTHOR]

Adaptive speciation can occur when a population undergoes assortative mating and disruptive selection caused by frequency- dependent intraspecific competition. However, other interactions, such as mutualisms based on trait matching, may generate conflicting selective pressures that constrain species diversification. We used individual-based simulations to explore how different types of mutualism affect adaptive diversification. A magic trait was assumed to simultaneously mediate mate choice, intraspecific competition, and mutualisms. In scenarios of intimate, specialized mutualisms, individuals interact with one or few individual mutualistic partners, and diversification is constrained only if the mutualism is obligate. In other scenarios, increasing numbers of different partners per individual limit diversification by generating stabilizing selection. Stabilizing selection emerges from the greater likelihood of trait mismatches for rare, extreme phenotypes than for common intermediate phenotypes. Constraints on diversification imposed by increased numbers of partners decrease if the trait matching degree has smaller positive effects on fitness. These results hold after the relaxation of various assumptions. When trait matching matters, mutualism-generated stabilizing selection would thus often constrain diversification in obligate mutualisms, such as ant-myrmecophyte associations, and in low-intimacy mutualisms, including plant-seed disperser systems. Hence, different processes, such as trait convergence favoring the incorporation of nonrelated species, are needed to explain the higher richness of low-intimacy assemblages—shown here to be up to 1 order of magnitude richer than high-intimacy systems. [ABSTRACT FROM AUTHOR]

Coevolution of plants and pollinators has been suggested as a mechanism driving diversification of plant-pollinator mutualisms. There is increasing recognition that predators or competitors can influence the abundance and behavior of pollinators and indirectly affect the fitness of plants. However, existing theories on plantpollinator diversification focus exclusively on mutualistic interactions between plants and pollinators. Here we used simulations to evaluate whether predation on pollinators promotes coevolutionary diversification of plant-pollinator mutualisms.We developed an individualbased simulation model in which the blooming season of plants and the active seasons of pollinators and predators can evolve. In simulations without predators, plant-pollinator coevolution caused diversification in blooming/active seasons for both plants and pollinators, but this diversification resulted in polymorphisms, not speciation. The introduction of predators promoted a split of plant and pollinator populations into reproductively isolated subpopulations with corresponding blooming and active seasons or a directional shift of blooming and active seasons, increasing the possibility of plant-pollinator cospeciation. This result suggests that predation on pollinators can promote sympatric and allopatric divergence of plant-pollinator mutualisms. Joint action of antagonistic and mutualistic interactions may be fundamentally important for diversification in coevolutionary interactions. [ABSTRACT FROM AUTHOR]

Both the abiotic environment and abiotic interactions among species contribute to shaping species assemblages. While the roles of habitat filtering and competitive interactions are clearly established, less is known about how positive interactions, whereby species benefit from the presence of one another, affect community structure. Here we assess the importance of positive interactions by studying Andean communities of butterflies that interact mutualistically via Müllerian mimicry. We show that communities at similar altitudes have a similar phylogenetic composition, confirming that filtering by altitude is an important process. We also provide evidence that species that interact mutualistically (i.e., species that share the same mimicry wing pattern) coexist at large scales more often than expected by chance. Furthermore, we detect an association between mimicry structure and altitude that is stronger than expected even when phylogeny is corrected for, indicating adaptive convergence for wing pattern and/or altitudinal range driven by mutualistic interactions. Positive interactions extend far beyond Müllerian mimicry, with many examples in plants and animals, and their role in the evolution and assembly of communities may be more pervasive than is currently appreciated. Our findings have strong implications for the evolution and resilience of community structure in a changing world. [ABSTRACT FROM AUTHOR]

In the light of rapid losses of biodiversity worldwide, it has become more important than ever to study the factors that ensure the continued existence of diverse ecological communities. Whereas the diversity-enhancing effects of antagonistic interactions are relatively well understood, much less is known about the contribution of mutualistic interactions to biodiversity maintenance. This study assesses the influence of mutualistic interactions with pollinators on the diversity of plant communities with alternative means of reproduction besides animal pollination. In contrast to a recent more general model of plant-animal mutualisms, the results of our simulations suggest that interactions with pollinators do not generally promote plant diversity, irrespective of the structure of the interaction network. Despite a potential for increased plant species richness through the positive effect of pollinators on plant birth rates, species richness was mostly negatively affected by the presence of pollinators because existing abundance asymmetries were amplified by animal pollination. Our results imply that for plant communities with alternative means of reproduction, the loss of pollinators will usually not lead to decreased diversity. However, whereas the immediate effects of pollinator loss on plant community composition may be negligible, the long-term population genetic consequences are likely to be severe. [ABSTRACT FROM AUTHOR]

Background Plants in over one hundred families in habitats worldwide bear extrafloral nectaries (EFNs). EFNs display a remarkable diversity of evolutionary origins, as well as diverse morphology and location on the plant. They secrete extrafloral nectar, a carbohydrate-rich food that attracts ants and other arthropods, many of which protect the plant in return. By fostering ecologically important protective mutualisms, EFNs play a significant role in structuring both plant and animal communities. And yet researchers are only now beginning to appreciate their importance and the range of ecological, evolutionary and morphological diversity that EFNs exhibit. Scope This Highlight features a series of papers that illustrate some of the newest directions in the study of EFNs. Here, we introduce this set of papers by providing an overview of current understanding and new insights on EFN diversity, ecology and evolution. We highlight major gaps in our current knowledge, and outline future research directions. Conclusions Our understanding of the roles EFNs play in plant biology is being revolutionized with the use of new tools from developmental biology and genomics, new modes of analysis allowing hypothesis-testing in large-scale phylogenetic frameworks, and new levels of inquiry extending to community-scale interaction networks. But many central questions remain unanswered; indeed, many have not yet been asked. Thus, the EFN puzzle remains an intriguing challenge for the future. [ABSTRACT FROM PUBLISHER]

Recent years have seen an explosion in the diversity of partner control mechanisms hypothesised to stabilise cooperative behaviour among unrelated individuals. Game theory suggests numerous strategies, each with specific decision rules that allow cooperators to control a non-contributing partner. This diversity of hypothetical strategies seems likely to reflect diversity in the types of intraspecific cooperation and interspecific mutualism that exist in nature. It is therefore important to provide a framework that explains similarities and differences between the various hypothetical strategies and that predicts how key parameters that describe the natural history of natural systems favour different control mechanisms. We develop a novel unifying framework for pairwise interactions between unrelated individuals, in which we link specific control mechanisms to specific game structures. The latter are defined by unique combinations of the states of five parameters that describe investment, aspects of the payoff matrix, the number of interactions and partner choice. We find that specific control mechanisms potentially have utility in a limited number of game structures; conversely, each game structure may typically offer a few competing control mechanisms. Our framework offers theoreticians specific problems that await mathematical exploration, while at the same time offering empiricists guidelines for evaluating the game structure and corresponding control mechanisms in their systems. [ABSTRACT FROM AUTHOR]

With the growing recognition of exploiters as a prominent and enduring feature of many mutualisms, there is a need to understand the ecological and evolutionary dynamics of mutualisms in the context of exploitation. Here, we model coevolution between mutualist and exploiter birth rates, using an obligate pollinating seed parasite mutualism associated with a nonpollinating exploiter as a reference system. In this system, mutualist and exploiter larvae parasitize the host plant, competing for and consuming seeds. Evolution of the mutualist determines which exploiters can invade successfully. Subsequent coevolution with an exploiter has a strong, predictable influence on mutualist-exploiter coexistence, mutualist and exploiter phenotypes, and species abundances. Weak mutualist competition promotes "evolutionary purging" of the exploiter, while weak exploiter competition leads to "evolutionary suicide" of the system. When stable, long-term coexistence occurs, we identify two main "trait-abundance syndromes" that have three novel implications. (1) Persistent, highly parasitic exploiters can be favored by coevolution. (2) Even then, the density of coevolved mutualists can be high. (3) Low plant density results primarily from the evolution of mutualist, not exploiter, birth rate and density. To evaluate these predictions, studies are needed that identify and compare populations with and without exploiters and compare life-history traits of mutualists and exploiters. [ABSTRACT FROM AUTHOR]

1. While the relationship between facilitation and competition has been explored extensively in recent years, there is also a natural link between facilitation and mutualism, as both are interspecific interactions that confer benefits. Yet, the relationship between these two interactions has been minimally explored. 2. Here, I explore parallels and differences between mutualism and facilitation. Five focal areas organize current research on mutualism evolution: trait evolution; the continuum from specialization to generalization; the evolutionary origins and maintenance of the interaction; co-evolution of partners; and the prevalence and implications of cheating. These foci are also helpful for investigating how facilitation evolves, a much less explored issue. 3. Testable hypotheses regarding the evolution of facilitation include the following: selection should be stronger on traits of facilitated species than on traits of facilitators; facilitative interactions with mutualistic (++) and commensal (+0) outcomes should exhibit greater evolutionary stability than those with antagonistic (+−) outcomes; co-evolution should be possible in mutualistic and antagonistic facilitation only; when co-evolution occurs, it should produce a geographic mosaic of interaction outcomes; and antagonistic facilitation could lead to selection on facilitators to either escape or to tolerate the neighbours that benefit from them. 4. Synthesis. Three gaps in our knowledge currently impede progress on evolutionary questions surrounding facilitation. First, reciprocal effects are rarely investigated; facilitation might evolve like mutualism, commensalism or antagonism, depending on effects on the facilitator species. Secondly, the genetics of relevant traits are not yet well explored; the traits themselves are better known for facilitator species than for the facilitated, which are more likely to evolve in the context of the interaction. Finally, the fitness costs and benefits associated with facilitation have rarely been measured. Filling these gaps should permit rapid progress in understanding how facilitation arises, persists and evolves. [ABSTRACT FROM AUTHOR]

Background and Aims: A deeper understanding of mutualism can be reached by studying systems with measurable costs and benefits. Most studies of this type focus on an unusual class of obligate, species-specific pollination mutualisms. The interaction between Datura wrightii (Solanaceae) and the hawkmoth Manduca sexta offers similar advantages but greater generality. Adult moths both nectar at and deposit eggs on the same plant; larvae are herbivorous. The antagonistic component of this interaction has been well studied. Here the role of M. sexta as a pollinator of D. wrightii, particularly in the context of this moth's frequent nectaring visits to the bat-pollinated plant Agave palmeri, is documented. [ABSTRACT FROM PUBLISHER]

Mutualisms are ubiquitous in nature, as is their exploitation by both conspecific and heterospecific cheaters. Yet, evolutionary theory predicts that cheating should be favoured by natural selection. Here, we show theoretically that asymmetrical competition for partners generally determines the evolutionary fate of obligate mutualisms facing exploitation by third-species invaders. When asymmetry in partner competition is relatively weak, mutualists may either exclude exploiters or coexist with them, in which case their co-evolutionary response to exploitation is usually benign. When asymmetry is strong, the mutualists evolve towards evolutionary attractors where they become extremely vulnerable to exploiter invasion. However, exploiter invasion at an early stage of the mutualism's history can deflect mutualists’ co-evolutionary trajectories towards slightly different attractors that confer long-term stability against further exploitation. Thus, coexistence of mutualists and exploiters may often involve an historical effect whereby exploiters are co-opted early in mutualism history and provide lasting ‘evolutionary immunization’ against further invasion. [ABSTRACT FROM AUTHOR]

Contents Summary 412 I. Introduction 413 II A historical perspective on mutualism 413 III. Insect pollination 414 IV. Protection of plants by ants 417 V. Ant-mediated seed dispersal 419 VI. Discussion 420 Acknowledgements 423 References 424 Mutualisms (cooperative interactions between species) have had a central role in the generation and maintenance of life on earth. Insects and plants are involved in diverse forms of mutualism. Here we review evolutionary features of three prominent insect–plant mutualisms: pollination, protection and seed dispersal. We focus on addressing five central phenomena: evolutionary origins and maintenance of mutualism; the evolution of mutualistic traits; the evolution of specialization and generalization; coevolutionary processes; and the existence of cheating. Several features uniting very diverse insect–plant mutualisms are identified and their evolutionary implications are discussed: the involvement of one mobile and one sedentary partner; natural selection on plant rewards; the existence of a continuum from specialization to generalization; and the ubiquity of cheating, particularly on the part of insects. Plant–insect mutualisms have apparently both arisen and been lost repeatedly. Many adaptive hypotheses have been proposed to explain these transitions, and it is unlikely that any one of them dominates across interactions differing so widely in natural history. Evolutionary theory has a potentially important, but as yet largely unfilled, role to play in explaining the origins, maintenance, breakdown and evolution of insect–plant mutualisms. [ABSTRACT FROM AUTHOR]

In generalized, multispecies mutualisms, competition among members of one guild can influence the net benefits that each species in the other guild receives. Hence seasonal factors that affect the dynamics of competition can also affect net benefits, especially if the benefit or cost of mutualism also varies seasonally. In the Sonoran Desert, two common species of generalist ants compete for access to extrafloral nectaries on the fishhook barrel cactus Ferocactus wislizeni, but their relative abundances vary seasonally; one ant dominates more cacti in winter/spring, and the other in summer/autumn. Recently, a third ant species, which also varies in abundance seasonally, has appeared at our study sites and is competing with resident ants for access to cactus nectaries. This empirical system motivated us to examine a metapopulation model of competition for patches in an open system with periodic forcing. We use the model to: (1) illustrate three ways in which competing species may differ in their sensitivities to environmental conditions that are consistent with seasonally displaced patterns of abundance; (2) ask under what conditions the invasion of a third competitor into a two-species system could alter the sensitivity of the system to environmental forcing at low vs. high frequency; and (3) show how differences among competitors in the pattern of seasonal forcing alone can dramatically alter the outcome of a competitor invasion. We discuss implications of these results for the ways in which seasonal variation among competing mutualists, and the benefits and costs of mutualism, may affect the functioning of generalized, facultative mutualisms. [ABSTRACT FROM AUTHOR]

Studies the conditions for local coexistence of obligate plant/pollinating seed parasite mutualisms and nonpollinating exploiters. Importance of intraspecific and interspecific competition among and between pollinators for achieving three-way coexistence; Strong intraspecific competition facilitating exploiter invasion and coexistence.

abstract: We develop an approach for studying population dy-namics resulting from mutualism by employing functional responses based on density-dependent benefits and costs. These functional re-sponses express how the population growth rate of a mutualist is modified by the density of its partner. We present several possible dependencies of gross benefits and costs, and hence net effects, to a mutualist as functions of the density of its partner. Net effects to mutualists are likely a monotonically saturating or unimodal function of the density of their partner. We show that fundamental differences in the growth, limitation, and dynamics of a population can occur when net effects to that population change linearly, unimodally, or in a saturating fashion. We use the mutualism between senita cactus and its pollinating seed-eating moth as an example to show the influence of different benefit and cost functional responses on pop-ulation dynamics and stability of mutualisms. We investigated two mechanisms that may alter this mutualism's functional responses: distribution of eggs among flowers and fruit abortion. Differences in how benefits and costs vary with density can alter the stability of this mutualism. In particular, fruit abortion may allow for a stable equilibrium where none could otherwise exist. Keywords: benefits and costs, density dependence, functional re-sponse, mutualism, population dynamics, stability. [ABSTRACT FROM AUTHOR]

Presents a quantitative assessment of the frequency of mutualism studies in nine major journals devoted to ecological and evolutionary research. Distribution of topics and viewpoints taken toward mutualism; Factors preventing the integration of data on mutualisms; Eight questions posing broad-based generalizations on evolution and ecology of mutualism.

See also the Commentary by Selosse [ABSTRACT FROM AUTHOR]

Mutualisms, cooperative interactions between species, generally involve an economic exchange: species exchange commodities that are cheap for them to provide, for ones that cannot be obtained affordably or at all. But these associations can only succeed if effective partners can be enticed to interact. In some mutualisms, partners can actively seek one another out. However, plants, which use mutualists for a wide array of essential life history functions, do not have this option. Instead, natural selection has repeatedly favoured the evolution of rewards - nutritional substances (such as sugar-rich nectar and fleshy fruit) with which plants attract certain organisms whose feeding activities can then be co-opted for their own benefit. The trouble with rewards, however, is that they are usually also attractive to organisms that confer no benefits at all. Losing rewards to 'exploiters' makes a plant immediately less attractive to the mutualists it requires; if the reward cannot be renewed quickly (or at all), then mutualistic service is precluded entirely. Thus, it is in plants' interests to either restrict rewards to only the most beneficial partners or somehow punish or deter exploiters. Yet, at least in cases where the rewards are highly nutritious, we can expect counter-selection for exploiter traits that permit them to skirt such control. How, then, can mutualisms persist? In this issue, Orona-Tamayo et al. () describe a remarkable adaptation that safeguards one particularly costly reward from nonmutualists. Their study helps to explain the evolutionary success of an iconic interaction and illuminates one way in which mutualism as a whole can persist in the face of exploitation. [ABSTRACT FROM AUTHOR]

The article discusses the concept of mutualism and interactions between the species in the occurrence of cooperation in nature. Topics mentioned include the individual unique relationships of the animals, the existence of a mutualism-parasitism continuum and the seminal contribution of mutualism to ecology.

This article is a commentary on Chomicki et al., 211: 1358–1370. [ABSTRACT FROM AUTHOR]

The competitive exclusion principle is fundamental to understanding coexistence. Well‐established theories predict the conditions for coexistence in consumer–resource interactions. Given that species often compete for commodities offered by mutualists, competitive exclusion theory should also be critical to understanding how mutualisms function. We explicitly apply the competitive exclusion principle to mutualism and derive a rule analogous to Tilman's R* rule for exploitative competition. Coexistence is impossible when competitors compete solely for a shared partner‐provided commodity because superior competitors deplete that commodity sufficiently to exclude inferior competitors. We then investigate how competition between two guild members for a partner‐provided commodity and a resource external to the mutualism affects competitor coexistence. There are three key results. First, coexistence is possible via partitioning of a partner‐provided commodity and another resource. Second, unlike in classic R* Theory, competitive outcomes are influenced both by species' abilities to obtain commodities and their mutualisms with the shared commodity‐providing partner, which can indirectly alleviate competitors' commodity limitation. Third, the outcome of competition has important consequences for the commodity‐providing partner, which depend on the type of mutualism and the competitive abilities of competing mutualists. This theory provides a novel framework for investigating how competitors for mutualistic commodities coexist in nature. [ABSTRACT FROM AUTHOR]

• We model two species that interact antagonistically but share a mutualist partner. • The species interact with different life history stages of the shared partner. • Even if the antagonism is not consumptive, persistence of the system is rare. • Strong indirect interactions can lead to complex oscillatory dynamics. • Outcomes critically depend on the degree of specialization of the antagonists. Species often interact with multiple mutualistic partners that provide functionally different benefits and/or that interact with different life-history stages. These functionally different partners, however, may also interact directly with one another in other ways, indirectly altering net outcomes and persistence of the mutualistic system as a whole. We present a population dynamical model of a three-species system involving antagonism between species sharing a mutualist partner species with two explicit life stages. We find that, regardless of whether the antagonism is predatory or non-consumptive, persistence of the shared mutualist is possible only under a restrictive set of conditions. As the rate of antagonism between the species sharing the mutualist increases, indirect rather than direct interactions increasingly determine species' densities and sometimes result in complex, oscillatory dynamics for all species. Surprisingly, persistence of the mutualistic system is particularly dependent upon the degree to which each of the two mutualistic interactions is specialized. Our work investigates a novel mechanism by which changing ecological conditions can lead to extinction of mutualist partners and provides testable predictions regarding the interactive roles of mutualism and antagonism in net outcomes for species' densities. [ABSTRACT FROM AUTHOR]

Extinctions beget further extinctions when species lose obligate mutualists, predators, prey, or hosts. Here, we develop a conceptual model of species and community attributes affecting secondary extinction likelihood, incorporating mechanisms that buffer organisms against partner loss. Specialized interactors, including ‘cryptic specialists’ with diverse but nonredundant partner assemblages, incur elevated risk. Risk is also higher for species that cannot either evolve new traits following partner loss or obtain novel partners in communities reorganizing under changing environmental conditions. Partner loss occurs alongside other anthropogenic impacts; multiple stressors can circumvent ecological buffers, enhancing secondary extinction risk. Stressors can also offset each other, reducing secondary extinction risk, a hitherto unappreciated phenomenon. This synthesis suggests improved conservation planning tactics and critical directions for research on secondary extinctions. [ABSTRACT FROM AUTHOR]