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Supply and demand affect the values of goods exchanged in cooperative trades. Studies of humans and other species typically describe the standard scenario that an increase in demand leads to a higher price. Here, we challenge the generality of that logic with empirical data and a theoretical model. In our study system, "client" fishes visit cleaner wrasse (

Glochidion (Phyllanthaceae; leafflower trees) is a genus of trees which is widely reported to be pollinated by leafflower moths (Gracillariidae: Epicephala) in temperate and subtropical Asia, Australia, and the Pacific islands. However, the pollination ecology of Glochidion is not well described from tropical Asia, the region where it is most species-rich at both local (≤9 spp.) and regional (~200 spp.) scales. Here we report investigations of pollination biology and species-specificity of five Glochidion species in tropical Southeast Asia (Cambodia). Through nocturnal observations and fruit dissections, we find that at least three and likely five Glochidion species in Cambodia are pollinated by seed-parasitic leafflower moths. We find no evidence that any of these leafflower moths are non-mutualistic parasites, despite known examples of such parasites of this mutualism elsewhere in Asia. While the presence of a single larva in a fruit results in only a fraction of seeds being consumed, the presence of more than one larva per fruit—a frequent occurrence in some species—can result in almost all seeds within the fruit being infested. Multilocus phylogenetic analysis indicates that there are five different minimally monophyletic leafflower moth clades, each of which pollinates a unique Glochidion host species. Our results indicate that in its center of diversity in tropical Asia this system is an obligate pollination mutualism as previously described at the global margins of its distribution. These findings provide insights into the processes that generate and maintain biodiversity and maintain mutualism stability in plant–insect interactions in this biodiversity hotspot.

Premise of research. Root allocation and morphology can be influenced by nutritional symbiosis with arbuscular mycorrhizal (AM) fungi. In the presence of AM fungi, plants could alter root allocatio...

Species interactions shape the evolution of traits, life histories, and the pattern of speciation. What is less clear is whether certain types of species interaction are more or less likely to lead to phenotypic divergence among species. We used the brood pollination mutualism between yuccas and yucca moths to test how mutualistic (pollination) and antagonistic (oviposition) traits differ in the propensity to increase phenotypic divergence among pollinator moths. We measured traits of the tentacular mouthparts, structures used by females to actively pollinate flowers, as well as ovipositor traits to examine differences in the rate of evolution of these two suites of traits among pollinator species. Morphological analyses revealed two distinct groups of moths based on ovipositor morphology, but no such groupings were identified for tentacle morphology, even for moths that pollinated distantly related yuccas. In addition, ovipositor traits evolved at significantly faster rates than tentacular traits. These results support theoretical work suggesting that antagonism is more likely than mutualism to lead to phenotypic divergence.

The information on biological activity and species interaction in an ecosystem is essential to understand the mutualism and resource acquisition. A study was taken up to understand the interspecific association among whitefly, its predator and the associated ant species in Capsicum ecosystem. The whitefly was identified as invasive solanum whitefly, Aleurothrixus trachoides (Back), the coccinellid, which was predating on the whitefly as Axinoscymnus puttarudriahi Kapur and Munshi and the associated ant as Tapinoma melanocephalum (Fabricius). Studies on spatial distribution of the pest, predator and ant revealed that the pest and the predator followed aggregate distribution in upper and middle leaves of the plant. A highly significant positive correlation was exhibited between the predator grub and the immature stages of the whitefly as confirmed by Chi-square test, which was evident throughout the plant. Tapinoma melanocephalum exhibited positive association with the pest only in the lower leaves of Capsicum. The non-significant association of T. melanocephalum with the coccinellid predator, indicated that the ant did not interfere with the performance of the predator. The significant positive association of the coccinellid predator larvae and adults with the immature stages of the pest throughout the plant indicated the predatory efficiency of A. puttarudriahi, which needs to be conserved.

Generalization of pollination systems is widely accepted by ecologists in the studies of plant–pollinator interaction networks at the community level, but the degree of generalization of pollination networks remains largely unknown at the individual pollinator level. Using potential legitimate pollinators that were constantly visiting flowers in two alpine meadow communities, we analyzed the differences in the pollination network structure between the pollinator individual level and species level. The results showed that compared to the pollinator species‐based networks, the linkage density, interaction diversity, interaction evenness, the average plant linkage level, and interaction diversity increased, but connectance, degree of nestedness, the average of pollinator linkage level, and interaction diversity decreased in the pollinator individual‐based networks, indicating that pollinator individuals had a narrower food niche than their counterpart species. Pollination networks at the pollinator individual level were more specialized at the network level (H′2) and the plant species node level (d′) than at the pollinator species‐level networks, reducing the chance of underestimating levels of specialization in pollination systems. The results emphasize that research into pollinator individual‐based pollination networks will improve our understanding of the pollination networks at the pollinator species level and the coevolution of flowering plants and pollinators., Our finding revealed that the pollinator individual‐level pollination networks were specialized at network and plant node level. The pollinator individual‐based pollination networks reduce the chance of underestimating levels of specialization and improve our understanding of coevolution of flowering plants and pollinators in pollination systems.

Interactions between Sphagnum (peat moss) and cyanobacteria play critical roles in terrestrial carbon and nitrogen cycling processes. Knowledge of the metabolites exchanged, the physiological processes involved, and the environmental conditions allowing the formation of symbiosis is important for a better understanding of the mechanisms underlying these interactions. In this study, we used a cross-feeding approach with spatially resolved metabolite profiling and metatranscriptomics to characterize the symbiosis between Sphagnum and Nostoc cyanobacteria. A pH gradient study revealed that the Sphagnum–Nostoc symbiosis was driven by pH, with mutualism occurring only at low pH. Metabolic cross-feeding studies along with spatially resolved matrix-assisted laser desorption/ionization mass spectrometry imaging (MALDI-MSI) identified trehalose as the main carbohydrate source released by Sphagnum, which were depleted by Nostoc along with sulfur-containing choline-O-sulfate, taurine and sulfoacetate. In exchange, Nostoc increased exudation of purines and amino acids. Metatranscriptome analysis indicated that Sphagnum host defense was downregulated when in direct contact with the Nostoc symbiont, but not as a result of chemical contact alone. The observations in this study elucidated environmental, metabolic, and physiological underpinnings of the widespread plant–cyanobacterial symbioses with important implications for predicting carbon and nitrogen cycling in peatland ecosystems as well as the basis of general host-microbe interactions.

In this review, we highlight the main insights that have been gathered from recent developments using large-scale genomics of fungal saprotrophs and symbiotrophs (including ectomycorrhizal and orchid and ericoid mycorrhizal fungi) inhabiting forest ecosystems. After assessing the goals and motivations underlying our approach, we explore our current understanding of the limits and future potential of using genomics to understand the ecological roles of these forest fungi. Comparative genomics unraveled the molecular machineries involved in lignocellulose decomposition in wood decayers, soil and litter saprotrophs, and mycorrhizal symbionts. They also showed that transitions from saprotrophy to mutualism entailed widespread losses of lignocellulose-degrading enzymes; diversification of novel, lineage-specific symbiosis-induced genes; and convergent evolution of genetic innovations that facilitate the accommodationof mutualistic symbionts within their plant hosts. We also identify the major questions that remain unanswered and propose new avenues of genome-based research to understand the role of soil fungi in sustainable forest ecosystems.

Mutually beneficial interspecific interactions are abundant throughout the natural world, including between microbes. Mutualisms between microbes are critical for everything from human health to global nutrient cycling. Studying model microbial mutualisms in the laboratory enables highly controlled experiments for developing and testing evolutionary and ecological hypotheses. In this review, we begin by describing the tools available for studying model microbial mutualisms. We then outline recent insights that laboratory systems have shed on the evolutionary origins, evolutionary dynamics, and ecological features of microbial mutualism. We touch on gaps in our current understanding of microbial mutualisms, note connections to mutualism in nonmicrobial systems, and call attention to open questions ripe for future study.

Symbiotic interactions can shift along a mutualism-parasitism continuum. While there are many studies examining dynamics typically considered to be mutualistic that sometimes shift toward parasitism, little is known about conditions underlying shifts from parasitism toward mutualism. In lake populations, we observed that infection by a microsporidian gut symbiont sometimes conferred a reproductive advantage and other times a disadvantage to its Daphnia host. We hypothesized that the microsporidian might benefit its host by reducing infection by more virulent parasites, which attack via the gut. In a laboratory study using field-collected animals, we found that spores of a virulent fungal parasite were much less capable of penetrating the guts of Daphnia harboring the microsporidian gut symbiont. We predicted that this altered gut penetrability could cause differential impacts on host fitness depending on ecological context. Field survey data revealed that microsporidian-infected Daphnia hosts experienced a reproductive advantage when virulent parasites were common while resource scarcity led to a reproductive disadvantage, but only in lakes where virulent parasites were relatively rare. Our findings highlight the importance of considering multiparasite community context and resource availability in host-parasite studies and open the door for future research into conditions driving shifts along parasitism to mutualism gradients.

PREMISE Animal pollination is critical to plant reproduction and may cause convergent evolution of pollination syndromes. Pollination syndromes in Castilleja have been distinguished based on floral traits and historical observations of floral visitors. Here we addressed these questions: (1) Can pollination syndromes be distinguished using floral morphological traits or volatile organic compound emissions? (2) Is there significant variation in floral traits within a pollination syndrome at the level of populations or species? (3) Do pollination syndromes predict the most frequent floral visitor to Castilleja? METHODS Floral traits and visitation were measured for five co-occurring Castilleja species (C. applegatei, C. linariifolia, C. miniata, C. nana, and C. peirsonii), representing three pollination syndromes (bee, fly, and hummingbird), at four sites in the Sierra Nevada Mountains. We used nonmetric multidimensional scaling (NMDS) and multiple linear regressions to address key questions in the differentiation of Castilleja and floral visitors. RESULTS Our analyses revealed that both morphological traits and floral VOCs can be used to distinguish between some pollination syndromes and Castilleja species. Morphological traits defined pollination syndromes reliably, but within the hummingbird syndrome, there was also significant variation among populations and species. Pollination syndrome was a poor predictor of visitors to Castilleja. CONCLUSIONS Floral trait differentiation among Castilleja individuals reflects both taxonomy and pollination syndromes. Differentiation was generally more evident in morphological traits compared to VOCs. Furthermore, a priori notions of pollination syndromes in this system are overly simplistic and fail to predict which animals most frequently visit Castilleja in natural populations.

Heat-induced mass mortalities involving ecosystem engineers may have long-lasting detrimental effects at the community level, eliminating the ecosystem services they provide. Intertidal mussels are ecologically and economically valuable with some populations facing unprecedented heat-induced mass mortalities. Critically, mussels are also frequently infested by endolithic parasites that modify shell albedo, hence reducing overheating and mortality rates under heat stress. Using a biophysical model, we explored the topographical and meteorological conditions under which endolithically driven thermal buffering becomes critical to survival. Based on meteorological data from a global climate analysis, we modelled body temperatures of infested and non-infested mussels over the last decade (2010-2020) at nine sites spread across c. 20 degrees of latitude. We show that thermal buffering is enhanced where and when heat stress is greatest, that is, on sun-exposed surfaces under high solar radiation and high air temperature. These results suggest that new co-evolutionary pathways are likely to open for these symbiotic organisms as climate continues to change, potentially tipping the balance of the relationship from a parasitic to a more mutualistic one. However, endolithically driven reductions in body temperatures can also occur at or below optimal temperatures, thereby reducing the host's metabolic rates and making the interplay of positive and negative effects complex. In parallel, we hindcasted body temperatures using empirical data from nearby weather stations and found that predictions were very similar with those obtained from two global climate reanalyses (i.e. NCEP-DOE Reanalysis 2 and ECMWF Reanalysis v5). This result holds great promise for modelling the distribution of terrestrial ectotherms at ecologically relevant spatiotemporal scales, as it suggests we can reasonably bypass the practical issues associated with weather stations. For intertidal ectotherms, however, the challenge will be incorporating body temperatures over the full tidal cycle. info:eu-repo/semantics/publishedVersion

Nutritional mutualism between insects and symbiotic bacteria is widespread. The various sap-feeding whitefly species within the Bemisia tabaci complex associate with the same obligate symbiont (Portiera) and multiple secondary symbionts. It is often assumed that some of the symbionts residing in the whiteflies play crucial roles in the nutritional physiology of their insect hosts. Although effort has been made to understand the functions of the whitefly symbionts, the metabolic complementarity offered by these symbionts to the hosts is not yet well understood. We examined two secondary symbionts, Arsenophonus and Wolbachia, in two species of the B. tabaci whitefly complex, provisionally named as Asia II 3 and China 1. Genomic sequence analyses revealed that Arsenophonus and Wolbachia retained genes responsible for the biosynthesis of B vitamins. We then conducted transcriptomic surveys of the bacteriomes in these two species of whiteflies together with that in another species named MED of this whitefly complex previously reported. The analyses indicated that several key genes in B vitamin syntheses from the three whitefly species were identical. Our findings suggest that, similar to another secondary symbiont Hamiltonella, Arsenophonus and Wolbachia function in the nutrient provision of host whiteflies. Although phylogenetically distant species of symbionts are associated with their respective hosts, they have evolved and retained similar functions in biosynthesis of some B vitamins. Such metabolic complementarity between whiteflies and symbionts represents an important feature of their coevolution.

Many structural patterns have been found to be important for the stability and robustness of mutualistic plant–pollinator networks. These structural patterns are impacted by a suite of variables, including species traits, species abundances, their spatial configuration, and their phylogenetic history. Here, we consider a specific trait: phenology, or the timing of life history events. We expect that timing and duration of activity of pollinators, or of flowering in plants, could greatly affect the species' roles within networks in which they are embedded. Using plant–pollinator networks from 33 sites in southern British Columbia, Canada, we asked (a) how phenological species traits, specifically timing of first appearance in the network and duration of activity in a network, were related to species' roles within a network, and (b) how those traits affected network robustness to phenologically biased species loss. We found that long duration of activity increased connection within modules for both pollinators and plants and among modules for plants. We also found that date of first appearance was positively related to interaction strength asymmetry in plants but negatively related to pollinators. Networks were generally more robust to the loss of pollinators than plants, and robustness increased if the models allow new interactions to form when old ones are lost, constrained by overlapping phenology of plants and pollinators. Robustness declined with the loss of late‐flowering plants, which tended to have higher interaction strength asymmetry. In addition, robustness declined with loss of early‐flying or long‐duration pollinators. These pollinators tended to be among‐module connectors. Our results point to networks being limited by early‐flying pollinators. If plants flower earlier due to climate change, plant fitness may decline as they will depend on early emerging pollinators, unless pollinators also emerge earlier., In plant–pollinator networks of southern British Columbia, we found that networks were generally more robust to the loss of pollinators than plants. Robustness declined with the loss of late‐flowering plants, and with the loss of early‐flying or long‐duration pollinators.

Insect mutualisms are essential for reproduction of many plants, protection of plants and other insects, and provisioning of nutrients for insects. Disruption of these mutualisms by global change can have important implications for ecosystem processes. Here, we assess the general effects of global change on insect mutualisms, including the possible impacts on mutualistic networks. We find that the effects of global change on mutualisms are extremely variable, making broad patterns difficult to detect. We require studies focusing on changes in cost-benefit ratios, effects of partner dependency, and degree of specialization to further understand how global change will influence insect mutualism dynamics. We propose that rapid coevolution is one avenue by which mutualists can ameliorate the effects of global change.

Plants with a small number of specific pollinators may be vulnerable to fluctuations in the availability of those pollinators, which could limit plant reproductive success and even result in extinction. Plants can develop mechanisms to mitigate this risk, such as apomixis. Reproductive assurance mechanisms have been largely ignored in obligate pollination mutualisms (OPMs), that are some of the most specialised of plant-pollinator interactions. Furthermore, although OPMs are often referred to as obligate, this is rarely tested. We performed a flower-bagging experiment to test if the unisexual flowers of Breynia oblongifolia could set fruit in the absence of its highly specialised seed-eating moth pollinators. Surprisingly, many bagged female flowers developed fruits, suggesting apomixis. We therefore conducted a second series of experiments in which we 1) added or excluded pollinators from caged plants; and 2) surveyed a wild population for apomictic reproduction using mother-offspring genotyping. In the absence of pollinators, no fruits developed. In addition, we detected no genetic evidence for apomixis when comparing between mothers and their offspring or between adults in a wild population. We explain the production of fruits in bagged branches by our discovery that B. oblongifolia can retain pollinated female flowers over the winter period. These flowers develop to fruits in the spring in the absence of male flowers or pollinators. Our study thus shows that B. oblongifolia is unable to produce fruit in the absence of its specialist moth pollinators. Thus, the highly specific interaction between plant and pollinators appears to be truly obligate.

Farming of fungi by ants, termites, or beetles has led to ecologically successful societies fueled by industrial‐scale food production. Another type of obligate insect agriculture in Fiji involves the symbiosis between the ant Philidris nagasau and epiphytes in the genus Squamellaria (Rubiaceae) that the ants fertilize, defend, harvest, and depend on for nesting. All farmed Squamellaria form tubers (domatia) with preformed entrance holes and complex cavity networks occupied by P. nagasau. The inner surface of the domatia consists of smooth‐surfaced walls where the ants nest and rear their brood, and warty‐surfaced walls where they fertilize their crop by defecation. Here, we use RNA sequencing to identify gene expression patterns associated with the smooth versus warty wall types. Since wall differentiation occurred in the most recent common ancestor of all farmed species of Squamellaria, our study also identifies genetic pathways co‐opted following the emergence of agriculture. Warty‐surfaced walls show many upregulated genes linked to auxin transport, root development, and nitrogen transport consistent with their root‐like function; their defense‐related genes are also upregulated, probably to protect these permeable areas from pathogen entry. In smooth‐surfaced walls, genes functioning in suberin and wax biosynthesis are upregulated, contributing to the formation of an impermeable ant‐nesting area in the domatium. This study throws light on a number of functional characteristics of plant farming by ants and illustrates the power of genomic studies of symbiosis., We performed the first transcriptomic analysis of the newly discovered obligate farming symbiosis involving ants farming plants. Our analyses reveal a number of candidate genes associated with the differentiation of the plant tissue for ant nesting and fertilization.

Aquatic organisms rely on microbial symbionts for coping with various challenges they encounter during stress and for defending themselves against predators, pathogens and parasites. Microbial symbionts are also often indispensable for the host’s development or life cycle completion. Many aquatic ecosystems are currently under pressure due to diverse human activities that have a profound impact on ecosystem functioning. These human activities are also ex pected to alter interactions between aquatic hosts and their associated microbes. This can directly impact the host’s health and — given the importance and widespread occurrence of microbial symbiosis in aquatic systems — the ecosystem at large. In this review, we provide an overview of the importance of microbial symbionts for aquatic organisms, and we consider how the beneficial services provided by microbial symbionts can be affected by human activities. The scarcity of available studies that assess the functional consequences of human impacts on aquatic microbial symbioses shows that our knowledge on this topic is currently limited, making it difficult to draw general conclusions and predict future changes in microbial symbiont−host relationships in a changing world. To address this important knowledge gap, we provide an overview of ap proaches that can be used to assess the impact of human disturbances on the functioning of aquatic microbial symbioses. ispartof: AME vol:87 issue:Special 7 pages:113-138 status: published

Seed predators have the potential to act as agents of natural selection that influence seed traits and seed fates, which in turn affect the whole plant population dynamic. Accordingly, plants deploy a variety of mechanisms (e.g., resistance and tolerance strategies) to lessen the impact of predation on seed crop or on an individual seed. In this study, we described a novel mechanism, seed cloning strategy, in a tropical plant species in countering animal predation. By conducting field‐ and laboratory‐based germination experiments, we found that both rodent damaged and artificially damaged seed fragments of a large‐seeded tree Garcinia xanthochymus (Clusiaceae) could successfully germinate and establish as seedlings. Tissue culture experiments revealed that G. xanthochymus has no endosperm in seeds, and its seed fragments own strong capacity of differentiation and cloning. Seed damage negatively affected seedling growth and germination, but the seed germination rate was remarkably high. Our study suggests that, seed cloning capacity, adopted by the large‐seeded tree G. xanthochymus may act as a novel strategy counteract for seed predation and would play a significant role in stabilizing the mutualism between plant and animals., We revealed a distinct evolutionary strategy of seed cloning in countering animal predation, as compared to previous ones such as tolerance and resistance.

Several beetle species in the family Coccinellidae have evolved close associations with ants in order to consume ant-tended hemipteran prey. These myrmecophilous lady beetles employ various strategies for avoiding ant aggression, including physical and chemical protection. We asked how the lady beetle Diomus lupusapudoves Vandenberg, Iverson and Liere (Coccinellidae: Diomini) is capable of avoiding ant aggression in a Mexican coffee agroecosystem, given that its larvae are physically unprotected from the aggressive Azteca sericeasur Longino (Formicidae: Leptomyrmecini) ants. Through a classic scale-ant mutualism, these ants protect the primary food source of D. lupusapudoves, the green coffee scale, Coccus viridis Green (Coccidae: Coccini), an agricultural pest of coffee. Through three quantitative behavioral lab experiments, we found that the ants did not show aggression towards either D. lupusapudoves larvae or towards C. viridis. Similarly, the ants were not antagonistic to crickets (Orthoptera), which normally induce aggression, when they were covered in a coating of D. lupusapudoves larvae residues. Through a survival analysis, we found that D. lupusapudoves larvae are not apparently toxic to the ants, as ants that consumed a sugar solution containing the D. lupusapudoves larvae lived longer than those fed only a control sugar solution. From these series of experiments, we deduce that the D. lupusapudoves larvae likely employ chemical mimicry, allowing them to coexist with the aggressive A. sericeasur ants. The myrmecophilous behavior of the D. lupusapudoves larvae assists them in their role as an important biological control agent of the green coffee scale.

Mutualisms are key determinants of community assembly and composition, but urbanization can alter the dynamics of these interactions and associated effects on ecosystem functions. Legume-rhizobia mutualisms are a model interaction to evaluate the ecological and ecosystem-level effects of urbanization, particularly urban-driven eutrophication and nitrogen (N) deposition. Here, we evaluated how urbanization affected the ecology of the mutualism between white clover (Trifolium repens) and its rhizobial symbiont (Rhizobium leguminosarum symbiovar trifolii) along an urbanization gradient. We found that the abundance of rhizobium nodules on white clover decreased with urbanization. White clover acquired N from mixed sources of N fixation and uptake from the soil for the majority of the urbanization gradient, but white clover primarily acquired N from the soil rather than N fixation by rhizobia at the urban and rural limits of the gradient. Importantly, we identified soil N as a critical nexus for urban-driven changes in the white clover-rhizobium mutualism. Taken together, our results demonstrate that urbanization alters the ecological consequences of a legume-rhizobium mutualism, with direct and indirect effects of the urban landscape on an ecologically-important mutualistic interaction.

Cleaning symbiosis is one of the most iconic relationships between fishes. The cleaner picks parasites and diseased tissue from the body of so-called clients. Both parties benefit from this association: the cleaner obtains food and the client has unwanted organisms and material removed. Knowledge about cleaning interactions derives mostly from tropical and temperate reefs, although this relationship also occurs in fresh and brackish water. Cleaners and their clients in brackish water are the least known, mostly due to turbidity that precludes suitable observations. About half of the studies on brackish water cleaners come from aquarium observations, and studies in nature are scarce. I report here on cleaning by silver batfish (Monodactylus argenteus) juveniles in an Australian urban estuary. This cleaner holds stable or temporary cleaning stations and services clients as diverse as mullets, bream, pufferfish and catfish. The clients solicit the cleaner’s services by grouping and posing at the cleaning stations. When not engaged in cleaning the batfish pecks at periphyton, sometimes followed by a queue of clients. In addition to the description and documentation of cleaning interactions, I summarise here the knowledge about cleaners and their clients that dwell in brackish water. Ten cleaner and 16 client species sum up the knowledge about cleaning symbiosis in brackish water to date. I suggest that more instances of cleaning in this environment will come up from potential cleaners with further observational, natural history-oriented studies in estuaries under appropriate visual conditions, or aquarium observations.

Angiosperms have co-evolved with animals over thousands of years leading to an array of mutualistic relationships. Passage of plant seeds through animal intestines leads to an important mutualism providing the animal with food and the plant with seed dispersal and enhanced germination. This phenomenon is well studied in terrestrial angiosperms, but there is less research in aquatic environments. We studied the effect of gut passage in marine mega-herbivores (green sea turtles and dugongs) on seed germination for a common Australian seagrass, Zostera muelleri. We collected fecal samples likely to contain seeds, as well as seagrass seeds from plants at two coastal seagrass meadows in the central Great Barrier Reef World Heritage Area, Australia. Seeds collected from feces and plants were subjected to germination trials across different temperature treatments: low (19°C), medium (26°C), and high (32°C). We found excreted seeds had a significantly greater germination probability (two to four times greater) and germinated significantly faster (18–61% faster) than seeds from the plant. Excreted seeds which had not germinated at the end of the experiment were significantly less likely to be viable compared with seeds taken from the plant. Seeds released from the plant have a slow germination and low germination probability compared with excreted seeds, but retain a high percentage of seed viability. Our study is the first record of marine mega-herbivores enhancing germination of Z. muelleri seeds. By transporting seeds to new locations and enhancing germination, these animals are important in seagrass resilience and connectivity among metapopulations.

The discovery-exploitation tradeoff posits that ant colonies decide between investing in scouts (individual foraging and discovery) or recruits (collective foraging and exploitation) in order to optimize gains. Although this tradeoff has been thoroughly examined through theoretical modelling, research linking model findings to empirical comparative data in ant communities is lacking. We explore the discovery-exploitation tradeoff within an old field ant community in the novel context of a multispecies mutualism with the treehopper Publilia concava. Specifically, we examine the relationship between foraging strategy and ant attentiveness to their treehopper partner, changes in foraging strategy in response to dynamic resources, and how different attendance phenotypes affect partner fecundity. We categorized ants via behavioral assays into “attentive” and “non-attentive” behavioral groups. Foraging strategy differed between ant groups during treehopper oviposition, with “attentive” and “non-attentive” ants favoring collective and individual foraging strategies, respectively. As treehopper nymphs emerged, “non-attentive” Formica subsericea shifted to collective foraging, with a corresponding rapid increase in ant recruitment, while ant recruitment in “attentive” ants remained constant. Treehopper fecundity, however, did not differ when tended by either “attentive” or “non-attentive” ants. We suggest that differences in foraging strategy could be due to residual honeydew left by maternal treehoppers on egg masses, which “attentive” ants exploit. Early recruitment in “attentive” and behaviorally subordinate ants might lead to priority effects, allowing them to persist on host plants and deter invasion from competing ants, which could ultimately lead to species coexistence.

Nearly all organisms participate in multiple mutualisms, and complementarity within these complex interactions can result in synergistic fitness effects. However, it remains largely untested how multiple mutualisms impact eco-evolutionary dynamics in interacting species. We tested how multiple microbial mutualists-N-fixing bacteria and mycorrrhizal fungi-affected selection and heritability of traits in their shared host plant (Medicago truncatula), as well as fitness alignment between partners. Our results demonstrate for the first time that multiple mutualisms synergistically affect the selection and heritability of host traits and enhance fitness alignment between mutualists. Specifically, we found interaction with multiple microbial symbionts doubled the strength of natural selection on a plant architectural trait, resulted in 2- to 3-fold higher heritability of plant reproductive success, and more than doubled fitness alignment between N-fixing bacteria and plants. These findings show synergism generated by multiple mutualisms extends to key components of microevolutionary change, emphasising the importance of multiple mutualism effects on evolutionary trajectories.

Ant-hemipteran mutualism is one of the most frequently observed food-for-protection associations in nature, and is recently found to contribute to the invasions of several of the most destructive invasive ants. Chemical communication underlies establishment and maintenance of such associations, in which a multitude of semiochemicals, such as pheromones, cuticular hydrocarbons, honeydew sugars and bacteria-produced honeydew volatiles mediate location, recognition, selection, learning of mutualistic partners. Here, we review what is known about the chemical communication between ants and honeydew-producing hemipterans, and discuss how invasive ants can rapidly recognize and establish a mutualistic relationship with the hemipterans with which they have never coevolved. We also highlight some future directions for a clearer understanding of the chemical communication in ant-hemipteran mutualism and its role in ant invasions.

1. The biotic and abiotic context of individual plants within animal-pollinated plant populations can influence pollinator foraging behaviour. Pollinator movements regulate pollen flow among plant individuals, and ultimately determine individual plant reproductive success. Yet the underlying drivers of this context dependency of interactions at the population level and their functional consequences for individuals remain poorly known. 2. Here we used a well-characterised population of Halimium halimifolium (Cistaceae), a Mediterranean shrub species, in combination with exponential random graph models (ERGMs) to evaluate how the intrapopulation variation in plant attributes configures individual-based plant–pollinator networks and determines their reproductive outcomes. Specifically, we assessed (a) how the intrinsic (i.e. phenotype and phenology) and extrinsic (i.e. microsite) plant attributes influenced the emerging configuration of the bipartite plant–pollinator network and the unipartite plant–plant network derived from pollinator sharing, and (b) how these plant attributes combined with the network topological position of individual plants affect their female fitness, measured as the total seed weight per plant. 3. We found that both intrinsic and extrinsic plant attributes contributed substantially to explain the configuration of both the bipartite and the unipartite pollination network. Besides the effects of plant attributes, the functional group to which pollinator species belonged was also important to determine the variance in plant– pollinator interaction odds, while the probability of plants to share more pollinator species was additionally influenced by the spatial distance between those plants. Furthermore, our results showed that these influences of plant attributes on network structure can be translated into functional outcomes at the plant individual level, with direct consequences for intrapopulation fitness variation. 4. Synthesis. This study builds towards a better understanding of the multiple drivers underlying the context dependency of plant–pollinator interactions and how they mediate the reproductive outputs of individual plants within a population. The application of our analytical framework allows a conceptual shift from descriptive topredictive research on the evolutionary and ecological processes that give rise to complex ecological networks at the population level. Consejo Superior de Investigaciones Científicas de España. JAEINT18_EX_0080 Ministerio de Ciencia e Innovación de España. CGL2017-082847P, CGL2017-92436EXP y FPU19_02552

Symbioses exert substantial biological influence, with great evolutionary and ecological relevance for disease, major evolutionary transitions, and the structure and function of ecological communities. Yet, much remains unknown about the patterns and processes that characterise symbioses. A major unanswered question is the extent to which symbiont phylogenies mirror those of their hosts and if patterns differ for parasites and mutualists. Addressing this question offers fundamental insights into evolutionary processes, such as whether symbionts typically codiverge with their hosts or if diversity is generated via host switches. Here, we perform a meta-analysis of host-symbiont phylogenetic congruence, encompassing 212 host-symbiont cophylogenetic studies that include ~10,000 species. Our analysis supersedes previous qualitative assessments by utilising a quantitative framework. We show that symbiont phylogeny broadly reflects host phylogeny across biodiversity and life-history, demonstrating a general pattern of phylogenetic congruence in host-symbiont interactions. We reveal two key aspects of symbiont life-history that promote closer ties between hosts and symbionts: vertical transmission and mutualism. Mode of symbiosis and mode of transmission are intimately interlinked, but vertical transmission is the dominant factor. Given the pervasiveness of symbioses, these findings provide important insights into the processes responsible for generating and maintaining the Earth's rich biodiversity.

In recent decades, multipartite mutualisms involving microorganisms such as fungi have been discovered in associations traditionally thought of as bipartite. Ant-plant mutualisms were long thought to be bipartite despite fungi being noticed in an epiphytic ant-plant over 100 years ago. We sequenced fungal DNA from the three distinct domatium chambers of the epiphytic ant-plant Myrmecodia beccarii to establish if fungal communities differ by chamber type across five geographic locations spanning 675 km. The three chamber types serve different ant-associated functions including ‘waste’ chambers, where ant workers deposit waste; ‘nursery’ chambers, where the brood is kept; and ‘ventilation’ chambers, that allow air into the domatium. Overall, fungi from the order Chaetothyriales dominated the chambers in terms of the proportion of operational taxonomic units (OTUs; 13.4%) and sequence abundances of OTUs (28% of the total); however a large portion of OTUs (28%) were unidentified at the order level. Notably, the fungal community in the waste chambers differed consistently from the nursery and ventilation chambers across all five locations. We identified 13 fungal OTUs as ‘common’ in the waste chambers that were rare or in very low sequence abundance in the other two chambers. Fungal communities in the nursery and ventilation chambers overlapped more than either did with the waste chambers but were also distinct from each other. Differences in dominance of the common OTUs drove the observed patterns in the fungal communities for each of the chamber types. This suggests a multipartite mutualism involving fungi exists in this ant-plant and that the role of fungi differs among chamber types.

Specialization in mutualisms is thought to be a major driver of diversification, but few studies have explored how novel specialization evolves, or its relation to the evolution of other niche axes. A fundamental question is whether generalist interactions evolve to become more specialized (i.e., oscillation hypothesis) or if partner switches evolve without any change in niche breadth (i.e., musical chairs hypothesis). We examined alternative models for the evolution of specialization by estimating the mutualistic, climatic, and edaphic niche breadths of sister plant species, combining phylogenetic, environmental, and experimental data on Acmispon strigosus and Acmispon wrangelianus genotypes across their overlapping ranges in California. We found that specialization along all three niche axes was asymmetric across species, such that the species with broader climatic and edaphic niches, Acmispon strigosus, was also able to gain benefit from and invest in associating with a broader set of microbial mutualists. Our data are consistent with the oscillation model of specialization, and a parallel narrowing of the edaphic, climatic, and mutualistic dimensions of the host species niche. Our findings provide novel evidence that the evolution of specialization in mutualism is accompanied by specialization in other niche dimensions.

Plants typically interact with multiple above‐ and below‐ground organisms simultaneously, with their symbiotic relationships spanning a continuum ranging from mutualism, such as with arbuscular mycorrhizal fungi (AMF), to parasitism, including symbioses with plant‐parasitic nematodes (PPN).Although research is revealing the patterns of plant resource allocation to mutualistic AMF partners under different host and environmental constraints, the root ecosystem, with multiple competing symbionts, is often ignored. Such competition is likely to heavily influence resource allocation to symbionts.Here, we outline and discuss the competition between AMF and PPN for the finite supply of host plant resources, highlighting the need for a more holistic understanding of the influence of below‐ground interactions on plant resource allocation. Based on recent developments in our understanding of other symbiotic systems such as legume–rhizobia and AMF‐aphid‐plant, we propose hypotheses for the distribution of plant resources between contrasting below‐ground symbionts and how such competition may affect the host.We identify relevant knowledge gaps at the physiological and molecular scales which, if resolved, will improve our understanding of the true ecological significance and potential future exploitation of AMF‐PPN‐plant interactions in order to optimize plant growth. To resolve these outstanding knowledge gaps, we propose the application of well‐established methods in isotope tracing and nutrient budgeting to monitor the movement of nutrients between symbionts. By combining these approaches with novel time of arrival experiments and experimental systems involving multiple plant hosts interlinked by common mycelial networks, it may be possible to reveal the impact of multiple, simultaneous colonizations by competing symbionts on carbon and nutrient flows across ecologically important scales., Plants resources must be distributed between contrasting, concurrent symbionts. Here, we review and outline major gaps in knowledge surrounding the allocation of plant resources to two contrasting below‐ground root symbionts, the mutualistic arbuscular mycorrhizal fungi, and the parasitic plant parasitic nematodes. Based on recent developments in other symbiotic relationships, we propose hypotheses for the distribution of plant resources between these contrasting below‐ground partners and how this may affect the host.

Mutualism is a form of symbiosis whereby both parties benefit from the relationship. An example is cleaning symbiosis, which has been observed in terrestrial and marine environments. The most recognized form of marine cleaning symbiosis is that of cleaner fishes and their clients.Cleaner species set up cleaning stations on the reef, and other species seek out their services. However, it is not well understood how the presence of cleaning stations influence movements of large highly mobile species. We examined the role of cleaning stations as a driver of movement and habitat use in a mobile client species.Here, we used a combination of passive acoustic telemetry and in‐water surveys to investigate cleaning station attendance by the reef manta ray Mobula alfredi. We employed a novel approach in the form of a fine‐scale acoustic receiver array set up around a known cleaning area and tagged 42 rays. Within the array, we mapped structural features, surveyed the distribution of cleaner wrasse, and observed the habitat use of the rays.We found manta ray space use was significantly associated with blue‐streak cleaner wrasse Labroides dimidiatus distribution and hard coral substrate. Cleaning interactions dominated their habitat use at this site, taking precedence over other life history traits such as feeding and courtship.This study has demonstrated that cleaning symbiosis is a driver for highly mobile, and otherwise pelagic, species to visit inshore reef environments. We suggest that targeted and long‐term use of specific cleaning stations reflects manta rays having a long‐term memory and cognitive map of some shallow reef environments where quality cleaning is provided. We hypothesize that animals prefer cleaning sites in proximity to productive foraging regions., This study investigates the role of mutualism in determining site selection in marine megafauna. The aim was to determine the role of cleaning stations as a driver of movement and habitat use in a mobile client species. Through a combination of in‐water observations and a novel application of fine‐scale passive acoustic tracking, we demonstrated that cleaning symbiosis is a driver for highly mobile, and otherwise pelagic species to visit inshore reef environments. We suggest that targeted and long‐term use of specific cleaning stations reflects manta rays having a long‐term memory and cognitive map of some shallow reef environments where quality cleaning is provided. We hypothesize that animals preference cleaning sites in proximity to productive foraging regions.

Invasive ants shape assemblages and interactions of native species, but their effect on fundamental ecological processes is poorly understood. In East Africa, Pheidole megacephala ants have invaded monodominant stands of the ant-tree Acacia drepanolobium, extirpating native ant defenders and rendering trees vulnerable to canopy damage by vertebrate herbivores. We used experiments and observations to quantify direct and interactive effects of invasive ants and large herbivores on A. drepanolobium photosynthesis over a 2-year period. Trees that had been invaded for ≥ 5 years exhibited 69% lower whole-tree photosynthesis during key growing seasons, resulting from interaction between invasive ants and vertebrate herbivores that caused leaf- and canopy-level photosynthesis declines. We also surveyed trees shortly before and after invasion, finding that recent invasion induced only minor changes in leaf physiology. Our results from individual trees likely scale up, highlighting the potential of invasive species to alter ecosystem-level carbon fixation and other biogeochemical cycles.

Plant-fungal symbioses can have strong consequences for ecological communities and are sensitive to variation in abiotic factors. While the functions of mycorrhizal fungi are well established, the role of other root-colonizing fungi such as dark septate endophytes (DSE), which lack specialized structures for nutrient transfer, are less clear. DSE are ubiquitous in extreme and stressful ecosystems, including marine environments, and some studies suggest a potential role in plant nutrition. However, the response of DSE to nutrient availability and physical stress has rarely been tested in the field. We conducted a 10 mo field experiment to investigate how a symbiosis between the salt marsh plant Spartina alterniflora and DSE fungi responded to increased resources (nutrient addition) and physical stress (salt addition). Plant stem density and height increased in response to nutrient enrichment, consistent with past experiments in nutrient-limited marsh systems. Nutrient additions also increased S. alterniflora percent cover, but this effect was negated with elevated salinity. Nutrient addition decreased colonization by DSE hyphae by nearly half (8.8%, vs. 15.7% at ambient levels). Nutrients did not decrease DSE microsclerotia, which were marginally increased with the combination of added nutrients and salinity. These results are consistent with the view that plant-DSE interactions are based in part on enhanced nutritional condition of plants by fungi. In addition, there was a positive relationship between plant shoot growth and root colonization by DSE, suggesting a benefit of the association for the plants. Our results suggest that the poorly understood plant-DSE symbioses may be important in intertidal environments.

For successful colonization of host roots, ectomycorrhizal (EM) fungi must overcome host defense systems, and defensive phenotypes have previously been shown to affect the community composition of EM fungi associated with hosts. Secondary metabolites, such as terpenes, form a core part of these defense systems, but it is not yet understood whether variation in these constitutive defenses can result in variation in the colonization of hosts by specific fungal species.We planted seedlings from twelve maternal families of Scots pine (Pinus sylvestris) of known terpene genotype reciprocally in the field in each of six sites. After 3 months, we characterized the mycorrhizal fungal community of each seedling using a combination of morphological categorization and molecular barcoding, and assessed the terpene chemodiversity for a subset of the seedlings. We examined whether parental genotype or terpene chemodiversity affected the diversity or composition of a seedling's mycorrhizal community.While we found that terpene chemodiversity was highly heritable, we found no evidence that parental defensive genotype or a seedling's terpene chemodiversity affected associations with EM fungi. Instead, we found that the location of seedlings, both within and among sites, was the only determinant of the diversity and makeup of EM communities.These results show that while EM community composition varies within Scotland at both large and small scales, variation in constitutive defensive compounds does not determine the EM communities of closely cohabiting pine seedlings. Patchy distributions of EM fungi at small scales may render any genetic variation in associations with different species unrealizable in field conditions. The case for selection on traits mediating associations with specific fungal species may thus be overstated, at least in seedlings., Host defense genotype has previously been shown to affect associations with ectomycorrhizal (EM) fungi, but the extent to which secondary metabolites such as terpenes affect these associations is unknown. We conducted a reciprocal transplant field experiment to test the effect of varying terpene genotype on the EM community composition of individual seedlings and found that the only the location of a seedling affects community composition. Other processes affecting EM community composition at small scales, such as environmental preference and interspecific interactions, may thus preclude the expression of any effect of genotype in wild conditions.

Background Within tropical shallow-water coral reefs, marine sponges provide microhabitats for a wide range of fauna. Although there have been numerous studies and reports of symbiotic relationships amongst sponges and their associated fauna, those pieces of information are isolated and disconnected. For this reason, based on the available literature, we compiled a species-interaction dataset of coral reef marine sponge-associated fauna known to date. New information We introduce a dataset that includes 67 literature items that report 101 species of sponge hosts clustered in 12 Orders having a host/guest interaction with 284 guest species from six Phyla present in the Northwestern Tropical Atlantic coral reefs. This dataset consists of two types of information: 1. Machine-readable data and 2. Human-readable data. These two types of coding improve the scope of the dataset and facilitate the link between machine platforms and human-friendly displays. We also created an interactive visualisation of the species-interactions dataset and of a dynamic Chord Diagram of the host-guest species connections to generate a user-friendly link between the user and the dataset.

Pseudomyrmex concolor is an Amazonian ant that works as an induced biotic defence of its host plant. Nevertheless, there is no information about the division of labour in P. concolor and how it impacts its ant–plant mutualism. We hypothesized that the spatio-temporal variation in P. concolor workers might be reflected in labour division associated with colony/plant defence. For this, we took morphometric measures of ants present on the stem and leaves of Tachigali myrmecophila. They were marked, and the frequency with which they remained in the plant regions and behavioural responses to herbivores presence throughout the day was recorded. The results showed that the spatio-temporal variation influences the division of labour in the P. concolor colony, with workers showing local high fidelity over time and differentiated responses to the same behavioural stimulus, suggesting that the workers perform tasks depending on where they are located in the host plant. Probably workers of P. concolor exhibit age polyethism, with individuals performing different functions in the colony throughout their lifetime and changing their tasks in the colony as they get older and depending on their spatial location.

It is suggested that specialized mutualisms are more vulnerable to climate change. Ant-gardens (AGs) are a complex and specialized mutualistic system represented by epiphytic plants that specifically inhabit the arboreal nest built by canopy ants in tropical forests. Different ant-epiphyte ensembles constitute the AGs throughout the Neotropics. However, neither the environmental factors that determine their geographical distribution nor the effects of climate change on this canopy biological system are known. Here, we estimated the ecological niche and elevational distribution of the Neotropical AGs as an entity (regardless of species composition), and individually for six AG ant and 16 AG epiphyte species in order to determine and compare their current and future distributions (vulnerability), using two unrelated Global Circulation Models for the year 2070 under two Representative Concentration Pathways (RCP4.5: optimistic and RCP8.5: pessimistic). The current potential distribution of the AGs is discontinuous from Tamaulipas, Mexico, to Rio Grande do Sul, Brazil, in low elevation areas with high mean annual temperatures (> 25 °C) and precipitation (> 2400 mm). In contrast, the individual distributions of the AG ants and epiphytes tended not to follow to this climatic profile and were segregated by both latitude and elevation. The geographic distribution of most AG ant and epiphyte species diminished under climate change, while that of the AGs increased, even under the pessimistic scenario. This suggests that AGs allow the species that comprise them to broaden their ecological niche and be more resistant to climate change than they would be outside of this system.

Microbial mutualists provide substantial benefits to hosts that feed back to enhance the fitness of the associated microbes. In many systems, beneficial microbes colonize symbiotic organs, specialized host structures that house symbionts and mediate resources exchanged between parties. Mutualisms are characterized by net benefits exchanged among members of different species, however, inequalities in the magnitude of these exchanges could result in evolutionary conflict, destabilizing the mutualism. We investigated joint fitness effects of root nodule formation, the symbiotic organ of legumes that house nitrogen-fixing rhizobia in planta. We quantified host and symbiont fitness parameters dependent on the number of nodules formed using near-isogenic Lotus japonicus and Mesorhizobium loti mutants, respectively. Empirically estimated fitness functions suggest that legume and rhizobia fitness is aligned as the number of nodules formed increases from zero until the host optimum is reached, a point where aligned fitness interests shift to diverging fitness interests between host and symbiont. However, fitness conflict was only inferred when analyzing wild-type hosts along with their mutants dysregulated for control over nodule formation. These data demonstrate that to avoid conflict, hosts must tightly regulate investment into symbiotic organs maximizing their benefit to cost ratio of associating with microbes.

Supercolonial ants are among the largest cooperative units in nature, attaining extremely high densities. How these densities feed back into their population growth rates and how abundance and extrinsic factors interact to affect their population dynamics remain open questions. We studied how local worker abundance and extrinsic factors (rain, tree density) affect population growth rate and spread in the invasive big-headed ant, which is disrupting a keystone mutualism between acacia trees and native ants in parts of East Africa. We measured temporal changes in big-headed ant (BHA) abundance and rates of spread over 20 months along eight transects, extending from areas behind the front with high BHA abundances to areas at the invasion front with low BHA abundances. We used models that account for negative density dependence and incorporated extrinsic factors to determine what variables best explain variation in local population growth rates. Population growth rates declined with abundance, however, the strength of density dependence decreased with abundance. We suggest that weaker density dependence at higher ant abundances may be due to the beneficial effect of cooperative behavior that partially counteracts resource limitation. Rainfall and tree density had minor effects on ant population dynamics. BHA spread near 50 m/year, more than previous studies reported and comparable to rates of spread of other supercolonial ants. Although we did not detect declines in abundance in areas invaded a long time ago (> 10 years), continued monitoring of abundance at invaded sites may help to better understand the widespread collapse of many invasive ants.

Hitchhikers (phoretic organisms) need vehicles to disperse out of unsuitable habitats. Therefore, finding vehicles with the right functional attributes is essential for phoretic organisms. To locate these vehicles, phoretic organisms employ cues within modalities, ranging from visual to chemical senses. However, how hitchhikers discriminate between individual vehicles has rarely been investigated. Using a phoretic nematode community associated with an obligate fig-fig wasp pollination mutualism, we had earlier established that hitchhiking nematodes make decisions based on vehicle species identity and number of conspecific hitchhikers already present on the vehicle. Here we investigate if hitchhikers can differentiate between physiological states of vehicles. We asked whether phoretic nematodes choose between live or dead vehicles present in a chemically crowded environment and we investigated the basis for any discrimination. We conducted two-choice and single-choice behavioral assays using single nematodes and found that plant- and animal-parasitic nematodes preferred live over dead vehicles and used volatiles as a sensory cue to make this decision. However, in single-choice assays, animal-parasitic nematodes were also attracted towards naturally dead or freeze-killed wasps. The volatile profile of the wasps was dominated by terpenes and spiroketals. We examined the volatile blend emitted by the different wasp physiological states and determined a set of volatiles that the phoretic nematodes might use to discriminate between these states which is likely coupled with respired CO2. We determined that CO2 levels emitted by single wasps are sufficient to attract nematodes, demonstrating the high sensitivity of nematodes to this metabolic product.

Morelet’s crocodile (Crocodylus moreletii) and hybrid crocodiles (C. moreletii × acutus) are mound-nesting crocodilians found in the lowlands of northern and southern Belize, respectively. Nests of...

According to apparent competition theory, sharing a predator should cause indirect interactions among prey that can affect the structure and the dynamics of natural communities.Though shifts in prey dominance and predator resource use along environmental gradients are rather common, empirical evidence on the role of indirect prey–prey interactions through shared predation particularly with increasing productivity, is still scarce.In an 8‐week lake mesocosm experiment, we manipulated both the addition of inorganic nutrients and the presence of generalist fish predators (crucian carp, Carassius carassius L.), to test for the effects of indirect interactions through shared predation along a productivity gradient.We found that apparent mutualism (indirect positive interaction) between benthic and pelagic prey strongly affected short‐term responses of aquatic food webs to increasing productivity in the presence of a generalist fish. Increasing productivity favoured the relative abundance of benthic prey, following trends in natural productive lake systems. This led to a shift in fish selectivity from pelagic to benthic prey driven by changes in fish behaviour, which resulted in apparent mutualism due to the lower and delayed top‐down control of pelagic prey at increasing productivity.Our results show empirical evidence that the coupling of multiple production pathways can lead to strong indirect interactions through shared predation, whereby prey dynamics on short time‐scales are highly dependent on the foraging behaviour of generalist predators. This mechanism may play an important role in short‐term responses of food webs across environmental gradients., In an aquatic mesocosm study, the authors found that switching foraging in a generalist predator can lead to short‐term apparent mutualism between its benthic and pelagic prey. Such indirect prey–prey interactions had important consequences in prey community dynamics and strongly affected the food web response to increasing productivity.

Hitchhikers (phoretic organisms) identify their vehicles using species-specific visual, chemical and vibrational cues. However, what factors influence their choice between vehicles of the same species has rarely been investigated. Hitchhikers must not only avoid overcrowded vehicles but may also need to travel with conspecifics to ensure mates at their destination. Hence, a trade-off between overcrowding and presence of conspecifics likely determines the choice of a vehicle especially when destination sites are distant, ephemeral and unique. Here, we investigate whether a trade-off between the presence of conspecifics versus overcrowding by conspecifics or heterospecifics on a vehicle affects hitchhiker choice. We also investigate the sensory modality responsible for this choice. We experimentally examine these questions using a phoretic nematode community (containing plant- and animal-parasitic taxa) obligately associated with a brood-site pollination mutualism. In this model system, nematodes co-travel with conspecifics and heterospecifics on pollinators as vehicles, between ephemeral plant brood sites to complete their developmental life cycle. In this system, hitchhiker overcrowding has proven negative impacts on vehicle and plant fitness. We expected nematodes to respond to conspecifics and heterospecific density on offered vehicles when making their choice. We found that animal-parasitic nematodes preferred vehicles containing some conspecifics within a certain density range. However, plant-parasitic nematodes preferentially boarded vehicles that were devoid of conspecifics or had few conspecifics. Plant parasites that preferred empty vehicles likely hitchhiked in pairs. Both nematode types employed volatile cues to discriminate between vehicles with different conspecific nematode densities. Our results suggest that vehicle overcrowding by conspecifics, most likely, guaranteed access to mates at the destination determined hitchhiker choice. Surprisingly, and contrary to our expectations, plant- and animal-parasitic nematodes did not respond to heterospecific crowding on vehicles and did not discriminate between vehicles with different heterospecific nematode densities. The reason for this lack of response to heterospecific presence is unknown. This study not only shows that phoretic organisms use different strategies while choosing a vehicle but also confirms that density-dependent effects can ensure the stability and persistence of phoretic interactions in a mutualism by balancing overcrowding against reproductive assurance.