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Within-species variation in pollinator behavior is widely observed, but its causes have been minimally investigated. Pollinator sex is associated with large differences in behavior that may lead to predictable differences in flower foraging, but this expectation has not been explicitly tested. We investigate sex-associated differences in nectar-foraging behavior of the hawkmoth Hyles lineata, using pollen on the proboscis as a proxy for flower visitation. We tested two predictions emerging from the literature: (1) the sexes differ in the flower species they visit, (2) females are more specialized in flower choice. We also examined potential drivers underlying these predictions by performing field and laboratory experiments to test whether males (3) switch among flower species more frequently, or (4) fly farther and therefore encounter more species than females. Consistent with prediction (1), pollen load composition differed between the sexes, indicative of visitation differences. Contrary to prediction (2), females consistently carried more species-rich pollen loads than males. (3) Both sexes switched between flower species at similar rates, suggesting that differences in floral fidelity are unlikely to explain why females are less specialized than males. (4) Males flew longer distances than females; coupled with larger between-site differences in pollen composition for females, this result suggests that sex differences in mobility influence foraging, and that females may forage more frequently and in smaller areas than males. Together, our results demonstrate that sex-associated foraging differences can be large and consistent over time, and highlight the importance of sex as a driver of variation in pollinator behavior.

Individuals sometimes exhibit striking constancy to a single behaviour even when they are capable of short-term behavioural flexibility. Constancy enables animals to avoid costs such as memory constraints, but can also inflict significant opportunity costs through behaviour–environment mismatch. It is unclear when individuals should exhibit behavioural constancy and which types of costs most strongly influence such behaviour. We use a case in which individuals within a population exhibit more than one handling tactic for a single food type to investigate whether costs associated with switching among tactics constrain expression of intra-individual variation. Using wild bumble bees (Bombus spp.) that feed on nectar through flower openings (legitimate visits) or through holes at the base of flowers (robbing), we asked three questions. (1) Do individual bees exhibit tactic constancy within and across foraging bouts? (2) Are individuals willing to switch their food-handling tactics? (3) Is constancy in food-handling tactics maintained by costs associated with switching tactics? We measured energetic costs in addition to handling times. We found that bees freely foraging in meadows were highly constant to a single food-handling tactic both within and across bouts. However, experiments with individual captive bees showed that these bees were willing to switch tactics and experienced minimal costs in doing so. Thus, switching costs do not drive the observed constancy in food-handling tactics of bumble bees within and across foraging bouts.

Commensalisms, interactions between two species in which one species benefits and the other experiences no net effect, are frequently mentioned in the ecological literature but are surprisingly little studied. Here we review and synthesize our limited understanding of commensalism. We then argue that commensalism is not a single type of interaction; rather, it is a suite of phenomena associated with distinct ecological processes and evolutionary consequences. For each form of commensalism we define, we present evidence for how, where, and why it occurs, including when it is evolutionarily persistent and when it is an occasional outcome of interactions that are usually mutualistic or antagonistic. We argue that commensalism should be of great interest in the study of species interactions due to its location at the center of the continuum between positive and negative outcomes. Finally, we offer a roadmap for future research.

Plants allocate defences in order to decrease costs and maximize benefits against herbivores. The Optimal Defense Theory (ODT) predicts that continuously expressed (i.e. constitutive) defences are expected in structures of high value, whereas defences that are expressed or that increase their expression only after damage or upon risk of damage (i.e. induced defences) are expected in structures of low value. Although there are several studies evaluating ODT predictions, few studies have successfully tested them as a way of measuring ecological investment in extrafloral nectary (EFN)‐mediated ant–plant interactions. Here we compared extrafloral nectar production and ant attractiveness to EFNs located on vegetative versus reproductive plant structures on Qualea multiflora plants subjected to different levels of simulated herbivory. We then addressed the following predictions emerging from the ODT: (a) extrafloral nectar produced in inflorescence EFNs will have higher volumes and calories and will attract more ants than extrafloral nectar produced in leaf EFNs; (b) extrafloral nectar production (volume and calories) and ant attendance will increase after simulated herbivory in leaf EFNs but not in inflorescence EFNs; (c) higher simulated leaf herbivory will induce higher extrafloral nectar production in EFNs on leaves and (d) more attractive extrafloral nectar (higher volume and calories) will attract more ants. Extrafloral nectar volume and calorie content, as well as ant abundance, were higher in EFNs of inflorescences compared to EFNs of leaves both before and after simulated herbivory, consistent with one of our predictions. However, EFNs on both leaves and inflorescences, not on leaves only, were induced by simulated herbivory, a pattern opposite to our prediction. Plants subjected to higher levels of leaf damage produced more and higher calorie extrafloral nectar, but showed similar ant abundance. Finally, more attractive extrafloral nectar attracted more ants. Synthesis. Our results show that extrafloral nectar production before and after simulated herbivory, as well as ant recruitment, varies according to the plant structure on which EFNs are located. Our study is the first to show that ant recruitment via extrafloral nectar follows predictions from Optimal Defense Theory, and that the ant foraging patterns may be shaped by the plant part attacked and the level of damage it receives.

Many adult Lepidoptera nectar at flowers of plant species on which they also deposit their eggs. As a consequence, the same partner may act as both a pollinator and herbivore. How high are the costs associated with such herbivorous pollinators, relative to the potential benefits they confer? We addressed this question in the association between Datura wrightii (Solanaceae) and Manduca sexta (Sphingidae), which in southern Arizona, USA is both the plant’s most important pollinator (as adults) and most significant herbivore (as larvae). We manually inflicted two biologically relevant levels of artificial folivory (33% and 66% leaf removal) at two points during the growing season, to nearly 200 D. wrightii plants established in a common-garden outdoor plot. We recorded plant survival, growth, and several components of reproduction (bud, flower, and fruit production). We found no detectable effect of damage on plant growth rate, fruit set in the same or the following growing season, or plant volume or survival in the same or the following three seasons. These results suggest that D. wrightii may be highly tolerant of leaf consumption by the herbivorous offspring of its specialized, nectar-feeding pollinators. We argue that this mechanism would likely foster persistence of the pollination mutualism more effectively than would resistance to, or deterrence of, herbivores. Tolerance to the costs of mutualism is an underexplored phenomenon that could contribute to the stability of these interactions.

Plant–herbivore and plant–pollinator interactions are both well-studied, but largely independent of each other. It has become increasingly recognized, however, that pollination and herbivory interact extensively in nature, with consequences for plant fitness. Here, we explore the idea that trade-offs in investment in insect flight and reproduction may be a mechanistic link between pollination and herbivory. We first provide a general background on trade-offs between flight and fecundity in insects. We then focus on Lepidoptera; larvae are generally herbivores while most adults are pollinators, making them ideal to study these links. Increased allocation of resources to flight, we argue, potentially increases a Lepidopteran insect pollinator’s efficiency, resulting in higher plant fitness. In contrast, allocation of resources to reproduction in the same insect species reduces plant fitness, because it leads to an increase in herbivore population size. We examine the sequence of resource pools available to herbivorous Lepidopteran larvae (maternally provided nutrients to the eggs, as well as leaf tissue), and to adults (nectar and nuptial gifts provided by the males to the females), which potentially are pollinators. Last, we discuss how subsequent acquisition and allocation of resources from these pools may alter flight–fecundity trade-offs, with concomitant effects both on pollinator performance and the performance of larval herbivores in the next generation. Allocation decisions at different times during ontogeny translate into costs of herbivory and/or benefits of pollination for plants, mechanistically linking herbivory and pollination.

Resource dynamics influence the contemporary ecology of consumer-resource mutualisms. Suites of resource traits, such as floral nectar components, also evolve in response to different selective pressures, changing the ecological dynamics of the interacting species at the evolutionary equilibrium. Here we explore the evolution of resource-provisioning traits in a biotically pollinated plant that produces nectar as a resource for beneficial consumers. We develop a mathematical model describing natural selection on two quantitative nectar traits: maximum nectar production rate and maximum nectar reservoir volume. We use this model to examine how nectar production dynamics evolve under different ecological conditions that impose varying cost-benefit regimes on resource provisioning. The model results predict that natural selection favors higher nectar production when ecological factors limit the plant or pollinator’s abundance (e.g., a lower productivity environment or a higher pollinator conversion efficiency). We also find that nectar traits evolve as a suite in which higher costs of producing one trait select for a compensatory increase in investment in the other trait. This empirically explicit approach to studying the evolution of consumer-resource mutualisms illustrates how natural selection acting via direct and indirect pathways of species interactions generates patterns of resource provisioning seen in natural systems.

Datura wrightii (Solanaceae) , a common shrub in the southwestern United States, bears massive, white, night-blooming flowers that attract and reward hawkmoth pollinators. However, Apis mellifera (honeybee) foragers are often observed on its flowers, especially at dusk and dawn hours. Their foraging activities are focused on the anthers, suggesting they could be pollen thieves. We used a series of observations and manipulative experiments to determine if honeybees are detrimental or beneficial to D. wrightii. We found that honeybees were the most frequent visitors to D. wrightii flowers at both dusk and dawn, and that they removed and carried large amounts of D. wrightii pollen. Flowers were capable of being pollinated at dusk and dawn and a single visit by a honeybee was sufficient to pollinate the flowers and produce fruit. There was no evidence that restricting visitation to diurnal hours yielded fruit set that was different from when we restricted visitation (likely by hawkmoths) to evening hours. These results suggest that honeybees are capable of effective pollination of D. wrightii. Although honeybees might interfere with pollen transmission mediated by their highly specialized hawkmoth pollinators , they may also increase plant fitness by pollinating D. wrightii when hawkmoths are not present.

There is now good evidence that many mutualisms evolved from antagonism; why or how, however, remains unclear. We advance the Co-Opted Antagonist (COA) Hypothesis as a general mechanism explaining evolutionary transitions from antagonism to mutualism. COA involves an eco-coevolutionary process whereby natural selection favors co-option of an antagonist to perform a beneficial function and the interacting species coevolve a suite of phenotypic traits that drive the interaction from antagonism to mutualism. To evaluate the COA hypothesis, we present a generalized eco-coevolutionary framework of evolutionary transitions from antagonism to mutualism and develop a data-based, fully ecologically-parameterized model of a small community in which a lepidopteran insect pollinates some of its larval host plant species. More generally, our theory helps to reconcile several major challenges concerning the mechanisms of mutualism evolution, such as how mutualisms evolve without extremely tight host fidelity (vertical transmission) and how ecological context influences evolutionary outcomes, and vice-versa., Nature Communications, 12 (1), ISSN:2041-1723

It is increasingly apparent that mutualisms are central components of ecological communities, generating linkages among species in ways that lead to rich and persistent assemblages. I propose here an underlying rationale for why biological diversity tends to accumulate around mutualisms. I first highlight four key features of mutualism: most involve one-way or two-way exchange of resources; these resources are commonly used by many different mutualists; as they are difficult or impossible to modulate to mutualists’ needs, there would appear to be an advantage to overproduction; and many species other than mutualists subsist on these resources. As a consequence, I argue, (a) interaction networks form around mutualisms that extend well beyond the mutualists themselves; (b) non-mutualists taking advantage of one species’ resources may often be mutualists of other species in the community; and (c) different mutualistic networks are also linked, because successful mutualisms often generate resources that form the base of yet other mutualisms. I present three examples of plant/pollinator interactions that illustrate how mutualisms contribute to the accumulation of biological diversity in their local habitats. I conclude by offering a set of predictions to move these ideas further.

The application of ecological concepts and principles to the study of organization design and system development is now commonplace. Population ecology is one biological perspective that has been especially influential within the organization design literature. In the current study, we illustrate for the first time how this framework can be used to reveal when and in what ways intervention is warranted within a system that, like a biological population in nature, has emerged with little to no purposeful design. Specifically, we use 20 years of data on 1,074 Arizona charter schools to illustrate how population analysis can be used to uncover the characteristics of individual organizations that have the highest ability to survive over time within systems that have been allowed to emerge with little to no purposeful design. Our findings lead us to argue how pre-intervention population analysis at the system level can influence organization-level design choices in a way that enables dynamic fit and thereby enhances the likelihood of organization survivorship and innovation over time.

PREMISE In addition to its role as the male gamete, pollen is often used as a food reward for pollinators. Roughly 20,000 species of angiosperms are strictly pollen-rewarding, providing no other rewards to their pollinators. However, the influence of this strategy on pollinator behavior and plant reproduction is poorly understood, especially relative to the nectar-reward strategy. We performed a field experiment using the strictly pollen-rewarding Lupinus argenteus to explore how the absence of nectar influences pollinator behavior and plant reproduction. METHODS We added artificial nectar to Lupinus argenteus individuals to simulate a phenotype that would reward pollinators with both nectar and pollen. We compared bee pollinator behavior, via direct observation, and female reproduction between nectar-added and nectarless control plants. RESULTS Bees exhibited behavioral responses to the novel reward, collecting nectar as well as pollen and spending 27% longer per flower. Pollen transfer increased with flower visit duration. However, plants in the study population were not pollen-limited; consequently, the observed changes in pollinator behavior did not result in changes in female components of plant reproduction. CONCLUSIONS The addition of nectar to pollen-rewarding plants resulted in modest increases in per-flower pollinator visit duration and pollen transfer, but had no effect on reproduction because, at the place and time the experiment was conducted, plants were not pollen-limited. These results suggest that a pollen-only reward strategy may allow plants that are visited by pollen foragers to minimize some costs of reproduction by eliminating investment in other rewards, such as nectar, without compromising female plant fitness.

Mutualisms are critically important in maintaining the biodiversity and functioning of ecosystems. Mutualisms include a diverse array of interactions that result in reciprocal positive effects for both partners, including plant–pollinator, plant–seed disperser, and plant–rhizobia interactions. There is growing recognition that global environmental change can affect the ecological outcomes of mutualisms, but less attention has been paid to how urbanization in particular affects their evolution. This chapter builds from an ecological perspective and considers how urban landscapes may affect the evolutionary ecology of mutualism. It reviews the adaptive evolutionary processes that could affect mutualism in urban landscapes. It then surveys transportation, protection, and nutritional mutualisms to assess how urbanization may affect these mutualistic interactions in an evolutionary framework. The survey described in the chapter highlights a dearth of empirical and theoretical investigations on urban mutualisms from an evolutionary perspective despite potentially strong changes in selection pressures in urban areas. The chapter ends by outlining research directions to further the study of the evolutionary ecology of mutualisms in urban landscapes.

Although dispersal is generally viewed as a crucial determinant for the fitness of any organism, our understanding of its role in the persistence and spread of plant populations remains incomplete. Generalizing and predicting dispersal processes are challenging due to context dependence of seed dispersal, environmental heterogeneity and interdependent processes occurring over multiple spatial and temporal scales. Current population models often use simple phenomenological descriptions of dispersal processes, limiting their ability to examine the role of population persistence and spread, especially under global change. To move seed dispersal ecology forward, we need to evaluate the impact of any single seed dispersal event within the full spatial and temporal context of a plant’s life history and environmental variability that ultimately influences a population’s ability to persist and spread. In this perspective, we provide guidance on integrating empirical and theoretical approaches that account for the context dependency of seed dispersal to improve our ability to generalize and predict the consequences of dispersal, and its anthropogenic alteration, across systems. We synthesize suitable theoretical frameworks for this work and discuss concepts, approaches and available data from diverse subdisciplines to help operationalize concepts, highlight recent breakthroughs across research areas and discuss ongoing challenges and open questions. We address knowledge gaps in the movement ecology of seeds and the integration of dispersal and demography that could benefit from such a synthesis. With an interdisciplinary perspective, we will be able to better understand how global change will impact seed dispersal processes, and potential cascading effects on plant population persistence, spread and biodiversity., Although dispersal is generally viewed as a crucial determinant for the fitness of any organism, our understanding of its role in the persistence and spread of plant populations remains incomplete. In this perspective, we provide guidance on integrating empirical and theoretical approaches that account for the context dependency of seed dispersal to improve our ability to generalize and predict the consequences of dispersal, and its anthropogenic alteration, across systems. With an interdisciplinary perspective, we will be able to better understand how global change will impact seed dispersal processes, and potential cascading effects on plant population persistence, spread and biodiversity.

Many plants that bear hidden or recessed floral nectar experience nectar robbing, the removal of nectar by a floral visitor through holes pierced in the corolla. Although robbing can reduce plant reproductive success, many studies fail to find such effects. We outline three mechanistic hypotheses that can explain when interactions between plants and nectar‐robbers should be commensal rather than antagonistic: the non‐discrimination (pollinators do not avoid robbed flowers), visitor prevalence (robber visitation is rare relative to pollinator visitation), and pollen saturation (stigmas receive sufficient pollen to fertilize all ovules with one or very few pollinator visits) hypotheses. We then explore these mechanisms in the North American subalpine, bumble bee‐pollinated and nectar‐robbed plant Corydalis caseana (Fumariaceae). We first confirmed that the effects of nectar robbing on female reproductive success were neutral in C. caseana. We then tested the three mechanisms underlying these neutral effects using a combination of observational studies and experiments. We found evidence for all three mechanisms. First, consistent with the non‐discrimination hypothesis, pollinators failed to discriminate against experimentally robbed flowers or inflorescences even though naturally robbed flowers offered significantly lower nectar rewards than unrobbed flowers. Second, C. caseana was more commonly visited by pollinators than by nectar robbers, in accordance with the visitor prevalence hypothesis. Third, stigmas of unvisited flowers as well as those visited once by pollinators were saturated with pollen, with all stigmas bearing pollen loads several orders of magnitude higher than the number of ovules per fruit, consistent with the pollen saturation hypothesis. Our investigation of the mechanisms driving the commensal outcome of nectar robbing in this system deepens our understanding of the ecology of nectar robbing and contributes to a more general understanding of the variation in the outcomes of interactions between species.

Premise of the study Organisms engage in multiple species interactions simultaneously. While pollination studies generally focus on plants and pollinators exclusively, secondary robbing, a behavior that requires other species (primary robbers) to first create access holes in corollas, is common. It has been shown that secondary robbing can reduce plants' female fitness; however, we lack knowledge about its impact on male plant fitness. Methods We experimentally simulated primary and secondary robbing in the monocarpic perennial Ipomopsis aggregata (Polemoniaceae), quantifying indirect effects on pollinator-mediated pollen (dye) donation. We also assessed whether continual nectar removal via the floral opening has similar effects on hummingbird-pollinator behavior as continual secondary robbing through robber holes. Key results We found no significant indirect effects of secondary robbing on a component of Ipomopsis male fitness. Although robbing did reduce pollen (dye) donation due to avoidance of robbed plants by pollinating hummingbirds, pollen donation did not differ between the two robbing treatments. The effects of secondary robbing on hummingbird behavior resembled effects of chronic nectar removal by pollinators. Our results indicate that hummingbird pollinators may use a combination of cues, including cues given by the presence or absence of nectar, to make foraging decisions. Conclusions Combined with prior research, this study suggests that secondary robbing is less costly to a component of male fitness than to female fitness in Ipomopsis, broadening our knowledge of the overall costs of mutualism exploitation to total plant fitness.

Comparing traits of invasive species within and beyond their ancestral range may improve our understanding of processes that promote aggressive spread. Solanum elaeagnifolium (silverleaf nightshade) is a noxious weed in its ancestral range in North America and is invasive on other continents. We compared investment in flowers and ovules, pollination success, and fruit and seed set in populations from Arizona, USA (“AZ”) and Greece (“GR”). In both countries, the populations we sampled varied in size and types of present-day disturbance. Stature of plants increased with population size in AZ samples whereas GR plants were uniformly tall. Taller plants produced more flowers, and GR plants produced more flowers for a given stature and allocated more ovules per flower. Similar functional groups of native bees pollinated in AZ and GR populations, but visits to flowers decreased with population size and we observed no visits in the largest GR populations. As a result, plants in large GR populations were pollen-limited, and estimates of fecundity were lower on average in GR populations despite the larger allocation to flowers and ovules. These differences between plants in our AZ and GR populations suggest promising directions for further study. It would be useful to sample S. elaeagnifolium in Mediterranean climates within the ancestral range (e.g., in California, USA), to study asexual spread via rhizomes, and to use common gardens and genetic studies to explore the basis of variation in allocation patterns and of relationships between visitation and fruit set.

Parasitic plants often attack multiple host species with unique defenses, physiology, and ecology. Reproductive phenology and vectors of parasitic plant genes (pollinators and dispersers) can contribute to or erode reproductive isolation of populations infecting different host species. We asked whether desert mistletoe, Phoradendron californicum (Santalaceae tribe Visceae syn. Viscaceae), differs ecologically across its dominant leguminous hosts in ways affecting reproductive isolation. Parasite flowering phenology on one host species (velvet mesquite, Prosopis velutina) differed significantly from that on four others, and phenology was not predicted by host species phenology or host individual. Comparing mistletoe populations on mesquite and another common host species (catclaw acacia, Senegalia greggii) for which genetically distinct host races are known, we tested for differences in interactions with vectors by quantifying pollinator visitation, reward production, pollen receipt, and fruit consumption. Mistletoes on mesquite produced more pollinator rewards per flower (1.86 times the nectar and 1.92 times the pollen) and received ~ 2 more pollen grains per flower than those on acacia. Mistletoes on the two host species interacted with distinct but overlapping pollinator communities, and pollinator taxa differed in visitation according to host species. Yet, mistletoes of neither host showed uniformly greater reproductive success. Fruit set (0.70) did not differ by host, and the rates of fruit ripening and removal differed in contrasting ways. Altogether, we estimate strong but asymmetric pre-zygotic isolating barriers between mistletoes on the two hosts. These host-associated differences in reproduction have implications for interactions with mutualist vectors and population genetic structure.

Keep Engaging Youth in Science (KEYS) program of the University of Arizona BIO5 Institute; National Institute of Environmental and Health Sciences (NIEHS) [ES006694]; Science Foundation Arizona [BSP 0528-13]; National Science Foundation (NSF) [IOS-1053318]

How do animals forage for variable food resources? For animals foraging at flowers, floral constancy has provided a framework for understanding why organisms visit some flowers while bypassing others. We extend this framework to the flower-handling tactics that visitors employ. Nectar robbers remove nectar through holes bitten in flowers, often without pollinating. Many foragers can switch between robbing and visiting flowers legitimately to gain access to nectar. We document that even though individuals can switch foraging tactics, they often do not. We explore whether individuals exhibit constancy to either robbing or visiting legitimately, which we term tactic constancy. We then extend hypotheses of floral constancy to understand when and why visitors exhibit tactic constancy and raise questions for future research.

Organizational scholars often adopt biological models to explain the emergence and evolution of organizations and human systems. One recent example of such adoption is the organizational ecosystem metaphor. In this article, we contend that taking a rigorous ecological approach over the application of loose ecosystem language has the potential to illuminate patterns in the life span of organizations and human systems. We first define five central properties of biological ecosystems and demonstrate their potential relevance to human-constructed systems (organizational ecosystems). We then argue the value of developing biologically based hypotheses that can be tested in the context of organizational systems. Next, we propose a set of hypotheses specific to organizational stability and disruption, using Arizona charter schools as an example to demonstrate the promise of the rigorous application of the organizational ecosystem metaphor. We close with a discussion of how the insights generated might be applied across other organizational settings and systems.

In mutualistic interactions, the decision whether to cooperate or cheat depends on the relative costs and benefits of each strategy. In pollination mutualisms, secondary nectar robbing is a facultative behavior employed by a diverse array of nectar-feeding organisms, and is thought to be a form of cheating. Primary robbers create holes in floral tissue through which they feed on nectar, whereas secondary robbers, which often lack chewing mouthparts, feed on nectar through existing holes. Because primary robbers make nectar more readily available to secondary robbers, primary robbers facilitate the behaviors of secondary robbers. However, the net effect of facilitation on secondary robber fitness has not been empirically tested: it is unknown whether the benefit secondary robbers receive is strong enough to overcome the cost of competing with primary robbers for a shared resource. We conducted foraging experiments using the bumble bee Bombus bifarius, which can alternatively forage ‘legitimately’ (from the floral opening) or secondary-rob. We measured the relative foraging efficiencies (handling time per flower, flowers visited per minute, proportion of foraging bout spent consuming nectar) of these alternative behaviors, and tested whether the frequency of primary robbing and nectar standing crop in primary-robbed flowers of Linaria vulgaris (Plantaginaceae) affected foraging efficiency. Surprisingly, there was no effect of primary robbing frequency on the foraging efficiency of secondary-robbing B. bifarius. Instead, foraging strategy was a major predictor of foraging efficiency, with legitimate foraging being significantly more efficient than secondary robbing. Legitimate foraging was the more common strategy used by B. bifarius in our study; however, it is rarely used by B. bifarius foraging on L. vulgaris in nature, despite indications that it is more efficient. Our results suggest the need for deeper investigations into why bees adopt secondary robbing as a foraging strategy, specifically, the environmental contexts that promote the behavior.

Whether species interactions are static or change over time has wide-reaching ecological and evolutionary consequences. However, species interaction networks are typically constructed from temporally aggregated interaction data, thereby implicitly assuming that interactions are fixed. This approach has advanced our understanding of communities, but it obscures the timescale at which interactions form (or dissolve) and the drivers and consequences of such dynamics. We address this knowledge gap by quantifying the within-season turnover of plant-pollinator interactions from weekly censuses across 3 years in a subalpine ecosystem. Week-to-week turnover of interactions (1) was high, (2) followed a consistent seasonal progression in all years of study and (3) was dominated by interaction rewiring (the reassembly of interactions among species). Simulation models revealed that species' phenologies and relative abundances constrained both total interaction turnover and rewiring. Our findings reveal the diversity of species interactions that may be missed when the temporal dynamics of networks are ignored.

Indirect defence, the adaptive top-down control of herbivores by plant traits that enhance predation, is a central component of plant-herbivore interactions. However, the scope of interactions that comprise indirect defence and associated ecological and evolutionary processes has not been clearly defined. We argue that the range of plant traits that mediate indirect defence is much greater than previously thought, and we further organise major concepts surrounding their ecological functioning. Despite the wide range of plant traits and interacting organisms involved, indirect defences show commonalities when grouped. These categories are based on whether indirect defences boost natural enemy abundance via food or shelter resources, or, alternatively, increase natural enemy foraging efficiency via information or alteration of habitat complexity. The benefits of indirect defences to natural enemies should be further explored to establish the conditions in which indirect defence generates a plant-natural enemy mutualism. By considering the broader scope of plant-herbivore-natural enemy interactions that comprise indirect defence, we can better understand plant-based food webs, as well as the evolutionary processes that have shaped them.

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.

Minas Gerais Research Foundation; FAPEMIG [APQ-01043-13, PPM-00324-15]; Brazilian Council for Scientific and Technological Development, CNPq [472372/2013-0, 302700/2016-1]; University of Sao Paulo, PRP-USP [18.1.660.41.7]; Alexander von Humboldt-Stiftung, AvH [3.4-8151/15037]; Federal University of Minas Gerais, UFMG/PRPq [01/2013, 14/2013, 02/2014]

Intraspecific variation in floral visitor behaviour and pollination efficiency has been much less studied than interspecific variation. Nevertheless, it is clear that large differences in these traits exist within species, and in particular between sexes within species. With the exception of a few well-studied interactions, however, the consequences of these differences in the pollinators and visited plants remain to be investigated. In this review, we document large and consistent differences in the foraging patterns of male and female pollinators that have been demonstrated to directly affect plant reproduction or that have clear potential to do so. Males and females differ in visitation frequency, type of flowers visited, and per-visit pollen transfer. Females gather more and different resources from flowers compared to males, and males generally tend to show more mobile foraging patterns than females. We argue that these sex-associated patterns have broad generality across pollinators, and that sex-associated differences can in some cases be larger than differences between species. We offer predictions about how these patterns will influence pollinator preference, specialization, and fidelity, as well as the cost, quality and quantity of pollination service to plants. In the face of increasing threats to plant-pollinator interactions, understanding their basic functioning and the variation inherent in their component parts is critical. We advocate for more attention to sex-based differences among pollinators in particular, and the consequences of intraspecific variation more broadly.

Summary 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.

Premise of research. Biotic interactions have long been considered to be of less importance in structuring desert systems than other ecosystem types, but biotic interactions often play a critical role in meeting the challenges posed by the extreme conditions of desert environments. The Sonoran Desert, in particular, is home to several textbook examples of mutualisms, such as the interactions between the iconic saguaro cactus and its bat pollinators. But what do we know about the diversity, ecology, and evolution of plant-animal, plant-plant, and plant-microbe interactions and their impacts on individual plants and plant species in the Sonoran Desert?Methodology. To address this question, we review the published research on seven common kinds of plant biotic interactions by revisiting the respective literature, identifying gaps in our knowledge, and outlining future research directions.Pivotal results. Numerous gaps in our knowledge of plant biotic interactions in the Sonoran Desert were identified. Studie...

Interspecific interactions are important structuring forces in ecological communities. Interactions can be disturbed when species are lost from a community. When interactions result in fitness gains for at least one participating organism, that organism may experience reduced fitness as a result of interaction disturbance. However, many species exhibit traits that enable individuals to persist and reproduce in spite of such disruptions, resulting in resilience to interaction disturbance. Such traits can result in interaction generalization, phenotypic and behavioral plasticity, and adaptive capacity. We discuss examples of these traits and use case studies to illustrate how restoration practitioners can use a trait-based approach to examine species of concern, identify traits that are associated with interspecific interactions and are relevant to resilience, and target such traits in restoration. Restoration activities that bolster interaction resilience could include, for example, reintroducing or supporting specific functional groups or managing abiotic conditions to reduce interaction dependence by at-risk species (e.g. providing structural complexity offering shelter and cover). Resilience may also be an important consideration in species selection for restoration. Establishment of resilient species, able to persist after interaction disturbance, may be essential to restoring to a functioning ecological community. Once such species are present, they could help support more specialized species that lack resilience traits, such as many species of concern. Understanding the conditions under which processes linked to resilience may enable species to persist and communities to reform following interaction disturbance is a key application of community ecology to ecological restoration.

Floral communities present complex and shifting resource landscapes for flower-foraging animals. Strong similarities among the floral displays of different plant species, paired with high variability in reward distributions across time and space, can weaken correlations between floral signals and reward status. As a result, it should be difficult for foragers to discriminate between rewarding and rewardless flowers. Building on signal detection theory in behavioural ecology, we use hypothetical probability density functions to examine graphically how plant signals pose challenges to forager decision-making. We argue that foraging costs associated with incorrect acceptance of rewardless flowers and incorrect rejection of rewarding ones interact with community-level reward availability to determine the extent to which rewardless and rewarding species should overlap in flowering time. We discuss the evolutionary consequences of these phenomena from both the forager and the plant perspectives.This article is part of the theme issue ‘Signal detection theory in recognition systems: from evolving models to experimental tests’.

Women have long been underrepresented in the natural sciences, and although great progress has been made in recent decades, many subtle and not-so-subtle barriers persist. In this context, it is easy to get the impression that the early history of ecology and evolutionary biology was exclusively the domain of male researchers. In fact, a number of women made very substantial contributions to The American Naturalist in its first decades. In a follow-up to a series of retrospective essays celebrating 150 years of this journal, we highlight the scientific contributions of the women published in it during its first 50 years (1867-1916). We also discuss the diverse paths that their scientific careers took and the barriers they faced along the way.

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.