The complex circuitry of interactions determining coexistence among plants and mycorrhizal fungi.

We present a mechanistic model of coexistence among a mycorrhizal fungus and one or two plant species that compete for a single nutrient. Plant-fungal coexistence is more likely if the fungus is better at extracting the environmental nutrient than the plant and the fungus acquires carbon from the plant above a minimum rate. When they coexist, their interaction can shift from mutualistic to parasitic at high nutrient availability. The fungus is a second nutrient source for plants and can promote the coexistence of two plant competitors if one is better at environmental nutrient extraction and the other is better at acquiring the nutrient from the fungus. Because it extracts carbon from both plants, the fungus also serves as a conduit of apparent competition between the plants. Consequently, the plant with the lower environmental nutrient extraction rate can drive the plant with the higher environmental nutrient extraction rate extinct at high carbon supply rates. This model illustrates mechanisms to explain several observed patterns, including shifts in plant-mycorrhizal growth responses and coexistence along nutrient gradients, equivocal results among experiments testing the effect of mycorrhizal fungi on plant diversity, and differences in plant diversity among ecosystems dominated by different mycorrhizal groups.

Spatial prey availability and pulsed reproductive tactics: Encounter risk in a canid-ungulate system.

Predation risk is a function of spatiotemporal overlap between predator and prey, as well as behavioural responses during encounters. Dynamic factors (e.g. group size, prey availability and animal movement or state) affect risk, but rarely are integrated in risk assessments. Our work targets a system where predation risk is fundamentally linked to temporal patterns in prey abundance and behaviour. For neonatal ungulate prey, risk is defined within a short temporal window during which the pulse in parturition, increasing movement capacity with age and antipredation tactics have the potential to mediate risk. In our coyote-mule deer (Canis latrans-Odocoileus hemionus) system, leveraging GPS data collected from both predator and prey, we tested expectations of shared enemy and reproductive risk hypotheses. We asked two questions regarding risk: (A) How does primary and alternative prey habitat, predator and prey activity, and reproductive tactics (e.g. birth synchrony and maternal defence) influence the vulnerability of a neonate encountering a predator? (B) How do the same factors affect behaviour by predators relative to the time before and after an encounter? Despite increased selection for mule deer and intensified search behaviour by coyotes during the peak in mule deer parturition, mule deer were afforded protection from predation via predator swamping, experiencing reduced per-capita encounter risk when most neonates were born. Mule deer occupying rabbit habitat (Sylvilagus spp.; coyote's primary prey) experienced the greatest risk of encounter but the availability of rabbit habitat did not affect predator behaviour during encounters. Encounter risk increased in areas with greater availability of mule deer habitat: coyotes shifted their behaviour relative to deer habitat, and the pulse in mule deer parturition and movement of neonatal deer during encounters elicited increased speed and tortuosity by coyotes. In addition to the spatial distribution of prey, temporal patterns in prey availability and animal behavioural state were fundamental in defining risk. Our work reveals the nuanced consequences of pulsed availability on predation risk for alternative prey, whereby responses by predators to sudden resource availability, the lasting effects of diversionary prey and inherent antipredation tactics ultimately dictate risk.

Sperm-dependent asexual species and their role in ecology and evolution.

Sexual reproduction is the primary mode of reproduction in eukaryotes, but some organisms have evolved deviations from classical sex and switched to asexuality. These asexual lineages have sometimes been viewed as evolutionary dead ends, but recent research has revealed their importance in many areas of general biology. Our review explores the understudied, yet important mechanisms by which sperm-dependent asexuals that produce non-recombined gametes but rely on their fertilization, can have a significant impact on the evolution of coexisting sexual species and ecosystems. These impacts are concentrated around three major fields. Firstly, sperm-dependent asexuals can potentially impact the gene pool of coexisting sexual species by either restricting their population sizes or by providing bridges for interspecific gene flow whose type and consequences substantially differ from gene flow mechanisms expected under sexual reproduction. Secondly, they may impact on sexuals' diversification rates either directly, by serving as stepping-stones in speciation, or indirectly, by promoting the formation of pre- and postzygotic reproduction barriers among nascent species. Thirdly, they can potentially impact on spatial distribution of species, via direct or indirect (apparent) types of competition and Allee effects. For each such mechanism, we provide empirical examples of how natural sperm-dependent asexuals impact the evolution of their sexual counterparts. In particular, we highlight that these broad effects may last beyond the tenure of the individual asexual lineages causing them, which challenges the traditional perception that asexual lineages are short-lived evolutionary dead ends and minor sideshows. Our review also proposes new research directions to incorporate the aforementioned impacts of sperm-dependent asexuals. These research directions will ultimately enhance our understanding of the evolution of genomes and biological interactions in general.

Predator home range size mediates indirect interactions between prey species in an arctic vertebrate community.

Indirect interactions are widespread among prey species that share a common predator, but the underlying mechanisms driving these interactions are often unclear, and our ability to predict their outcome is limited. Changes in behavioural traits that impact predator space use could be a key proximal mechanism mediating indirect interactions, but there is little empirical evidence of the causes and consequences of such behavioural-numerical response in multispecies systems. Here, we investigate the complex ecological relationships between seven prey species sharing a common predator. We used a path analysis approach on a comprehensive 9-year data set simultaneously tracking predator space use, prey densities and prey mortality rate on key species of a simplified Arctic food web. We show that high availability of a clumped and spatially predictable prey (goose eggs) leads to a twofold reduction in predator (arctic fox) home range size, which increases local predator density and strongly decreases nest survival of an incidental prey (American golden plover). On the contrary, a scattered cyclic prey with potentially lower spatial predictability (lemming) had a weaker effect on fox space use and an overall positive impact on the survival of incidental prey. These contrasting effects underline the importance of studying behavioural responses of predators in multiprey systems and to explicitly integrate behavioural-numerical responses in multispecies predator-prey models.

Source-sink migration of natural enemies drives maladaptation of victim populations in sink habitats.

Natural enemies are critical drivers of species biogeography, and they may often limit the evolutionary adaptation and persistence of victim populations in sink habitats. Source-sink migration is also a major determinant of adaptation in sink habitats. Here, we specifically suggest that source-sink migration of enemies reduces evolutionary adaptation of victim populations in sink habitats. The underlying mechanisms may include depressed population size (which limits the supply of genetic variation) and enforced resistance evolution in victims (which shows a trade-off with growth performance). We experimentally tested this hypothesis using a model microbial system, bacterium Pseudomonas fluorescens (victim) and its lytic bacteriophage (enemy). The ancestral bacterial strain had lower growth performance at a cold temperature (10 °C, considered as sink habitat) than at its optimal temperature (28 °C, source habitat). Evolutionary adaptation took place in bacterial populations that evolved alone in the cold environment. When phages were present, no significant abiotic adaptation was observed. Crucially, phage immigration from source populations caused maladaptation, i.e., decreased growth performance relative to the ancestral genotype, although this was not the case when there was simultaneous immigration of phage and bacteria. Therefore, enemy-mediated intraspecific apparent competition could lead to prosperity in core habitats causing hardship in edge habitats.

Contrasting seed traits of co-existing seeds lead to a complex neighbor effect in a seed-rodent interaction.

Scatter-hoarding rodents play important roles in seed dispersal and predation in many forest ecosystems. Existing studies have shown that the seed foraging preference of rodents is directly affected by seed traits and indirectly affected by the traits of other co-existing seeds nearby (i.e., neighbor effect). Plant seeds exhibit a combination of diverse seed traits, including seed size, chemical defense, and nutrient content. Therefore, it is difficult to evaluate the influence of each single seed trait on such neighbor effects. Here, by using artificial seeds, we investigated the impacts of contrasts in seed size, tannin content, and nutrient content on neighbor effects. We tracked 9000 tagged artificial seeds from 30 seed-seed paired treatments in a subtropical forest in southwest China. The contrast in seed size between paired seeds created obvious neighbor effects measured through three seed dispersal related indicators: the proportion of seeds being removed, the proportion of seeds cached, and the distance transported by rodents. However, the magnitudes and the signs of the neighbor effects differed among pairs, including both apparent mutualism and apparent competition, depending on the contrast in seed size between paired seeds. The contrasts of tannin and nutrient content between paired seeds showed relatively few neighbor effects. Our results suggest that the contrast in seed traits between the target seed and its neighboring seeds should be considered when studying rodent-seed interactions. Furthermore, we expect that similar complex neighbor effects may also exist in other plant-animal interactions, such as pollination and herbivory.

Recovery of a marine keystone predator transforms terrestrial predator-prey dynamics.

Sea otters (Enhydra lutris) and wolves (Canis lupus) are two apex predators with strong and cascading effects on ecosystem structure and function. After decades of recovery from near extirpation, their ranges now overlap, allowing sea otters and wolves to interact for the first time in the scientific record. We intensively studied wolves during 2015 to 2021 in an island system colonized by sea otters in the 2000s and by wolves in 2013. After wolf colonization, we quantified shifts in foraging behavior with DNA metabarcoding of 689 wolf scats and stable isotope analyses, both revealing a dietary switch from Sitka black-tailed deer (Odocoileus hemionus), the terrestrial in situ primary prey, to sea otters. Here we show an unexpected result of the reintroduction and restoration of sea otters, which became an abundant marine subsidy for wolves following population recovery. The availability of sea otters allowed wolves to persist and continue to reproduce, subsequently nearly eliminating deer. Genotypes from 390 wolf scats and telemetry data from 13 wolves confirmed island fidelity constituting one of the highest known wolf population densities and upending standardly accepted wolf density predictions based on ungulate abundance. Whereas marine subsidies in other systems are generally derived from lower trophic levels, here an apex nearshore predator became a key prey species and linked nearshore and terrestrial food webs in a recently deglaciated and rapidly changing ecosystem. These results underscore that species restoration may serve as an unanticipated nutrient pathway for recipient ecosystems even resulting in cross-boundary subsidy cascades.

From bottom-up to top-down control of invertebrate herbivores in a retrogressive chronosequence.

In the long-term absence of disturbance, ecosystems often enter a decline or retrogressive phase which leads to reductions in primary productivity, plant biomass, nutrient cycling and foliar quality. However, the consequences of ecosystem retrogression for higher trophic levels such as herbivores and predators, are less clear. Using a post-fire forested island-chronosequence across which retrogression occurs, we provide evidence that nutrient availability strongly controls invertebrate herbivore biomass when predators are few, but that there is a switch from bottom-up to top-down control when predators are common. This trophic flip in herbivore control probably arises because invertebrate predators respond to alternative energy channels from the adjacent aquatic matrix, which were independent of terrestrial plant biomass. Our results suggest that effects of nutrient limitation resulting from ecosystem retrogression on trophic cascades are modified by nutrient-independent variation in predator abundance, and this calls for a more holistic approach to trophic ecology to better understand herbivore effects on plant communities.

Optimal prey switching: Predator foraging costs provide a mechanism for functional responses in multi-prey systems.

Foragers must balance the costs and gains inherent in the pursuit of their next meal. Classical functional response formulations describe consumption rates driven by prey density and are naive to predator foraging costs. Here, we integrated foraging costs into functional responses to add mechanism and precision to foundational ideas. Specifically, using a model system with a single predator and two prey, we express a functional response emerging from variable energy and time costs of each predation phase: searching, attacking, or consuming prey. The utility of our model is explored through a focused example where prey can exert variable influence on predator foraging costs through antipredator traits. Dissimilarity between prey in their foraging costs influence the energy gain rate of the predator through optimal prey switching. We found that a small subset of prey antipredator traits and density conditions generated a stabilizing Type III (sigmoidal) functional response-the pattern often thought to typify a generalist predator switching between prey species. The sigmoid functional response occurred for highly profitable prey only when the costly prey (1) were at a high density and (2) their antipredator traits increased energy or time costs following an encounter. We outline testable predictions regarding foraging costs from our model. We provide guidance on how to apply optimal foraging theory to empirical scenarios where predator foraging costs vary due to prey type, predator type, or environmental conditions. Our framework represents a synergy of foundational and contemporary theory across disciplines, facilitating the discovery of shared principles and context-dependent variation across varied predator-prey systems.

Predator discrimination of prey promotes the predator-mediated coexistence of prey species.

The predator discrimination of prey can affect predation intensity and the prey density dependence of predators, which has the potential to alter the coexistence of prey species. We used a predator-prey population dynamics model accounting for the predator's adaptive diet choice and predator discrimination of prey to investigate how the latter influences prey coexistence. The model revealed that (i) prey species that are perceived as belonging to the same species by a predator are attacked in the same manner, and it is more difficult for them to coexist than those that are recognized as different prey species, and (ii) prey species that are not discriminated by a predator-and therefore cannot coexist-may coexist in the presence of an alternative predator that does discriminate between them. These results suggest that prey diversity, which favours the predator discrimination of prey, and the different capabilities of predators to identify prey species both enhance prey coexistence.

Extending the gleaner-opportunist trade-off.

Species exhibit various trade-offs that can result in stable coexistence of competitors. The gleaner-opportunist trade-off to fluctuations in resource abundance is one of the most intuitive, yet also misunderstood, coexistence-promoting trade-offs. Here, we review its history as an ecological concept, discuss extensions to the classical theory and outline opportunities to advance its understanding. The mechanism of coexistence between species that grow relatively faster than their competitors in a low-resource environment (i.e. a gleaner) versus a high-resource environment (i.e. an opportunist) was first proposed in the 1970s. Stable coexistence could emerge between gleaners and opportunists if the opportunist species (dominant in unstable environments) dampens resource fluctuations via relatively convex functional responses, while the gleaner species (dominant in stable environments) promotes fluctuations, or diminishes them less than the opportunist does, via relatively saturating functional responses. This fluctuation-dependent coexistence mechanism has since been referred to by various names, including the Armstrong-McGehee mechanism and relative nonlinearity of competition. Several researchers have argued this mechanism likely plays a relatively minor role in species coexistence owing in part to the restricted range of conditions that allow it to operate. More recent theoretical research, however, suggests that relative nonlinearity can operate over wider conditions than previously thought. Here, we identify several novel, or little explored, extensions to the gleaner-opportunist trade-off that can yield species coexistence under phenomena as diverse as fluctuations in predation/pathogen pressure, multiple resources, phenotypic plasticity and rapid evolution, amongst other phenomena. While the original definition of the gleaner-opportunist trade-off may be imperfect as a collective for these extensions, we argue that a subtle reframing of the trade-off focusing on species' performance in equilibrium versus fluctuating conditions (irrespective of preferences for high or low resources, predation pressure or other competitive factors) reveals their fundamental commonality in stable coexistence via relative nonlinearity. An extended framing shines a light on the potential ubiquity of this canonical trade-off in nature and on the breadth of theoretical and empirical terrain that remains to be trodden.

Temporally auto-correlated predator attacks structure ecological communities.

For species primarily regulated by a common predator, the P* rule of Holt & Lawton (Holt & Lawton, 1993. Am. Nat.142, 623-645. (doi:10.1086/285561)) predicts that the prey species that supports the highest mean predator density (P*) excludes the other prey species. This prediction is re-examined in the presence of temporal fluctuations in the vital rates of the interacting species including predator attack rates. When the fluctuations in predator attack rates are temporally uncorrelated, the P* rule still holds even when the other vital rates are temporally auto-correlated. However, when temporal auto-correlations in attack rates are positive but not too strong, the prey species can coexist due to the emergence of a positive covariance between predator density and prey vulnerability. This coexistence mechanism is similar to the storage effect for species regulated by a common resource. Negative or strongly positive auto-correlations in attack rates generate a negative covariance between predator density and prey vulnerability and a stochastic priority effect can emerge: with non-zero probability either prey species is excluded. These results highlight how temporally auto-correlated species' interaction rates impact the structure and dynamics of ecological communities.

Indirect facilitation between prey promotes asymmetric apparent competition.

Apparent competition is one mechanism that can contribute to the complex dynamics observed in natural systems, yet it remains understudied in empirical systems. Understanding the dynamics that shape the outcome of processes like apparent competition is vital for appreciating how they influence natural systems. We empirically evaluated the role of indirect trophic interactions in driving apparent competition in a model laboratory system. Our experimental system was designed to let us evaluate combined direct and indirect interactions among species. Here we describe the results of a factorial experiment using two noncompeting prey (Colpidium kleini, a heterotroph, and Chlamydomonas reinhardtii, an autotroph) consumed by a generalist predator Euplotes eurystomus to explore the dynamics of apparent competition. To gain intuition into the potential mechanism driving the asymmetry in the observed results, we further explored the system using structural equation modelling. Our results show an important role of positive interactions and indirect effects contributing to apparent competition in this system with a marked asymmetrical outcome favouring one prey, Chlamydomonas. The selected structural equation supports a role of indirect facilitation; although Chlamydomonas (a photoautotroph) and Colpidium (a bacterivore) use different resources and therefor do not directly compete, Colpidium reduces bacteria that may compete with Chlamydomonas. In addition, formation of colonies by Chlamydomonas in response to predation by Euplotes provides an antipredator defence not available to Colpidium. Asymmetric apparent competition may be more common in natural systems than the symmetric interaction originally proposed in classic theory, suggesting that exploration of the mechanisms driving the asymmetry of the interaction can be a fruitful area of further research to better our understanding of interspecific interactions and community dynamics.

Invasive grass indirectly alters seasonal patterns in seed predation.

Invasive species threaten ecosystems globally, but their impacts can be cryptic when they occur indirectly. Invader phenology can also differ from that of native species, potentially causing seasonality in invader impacts. Yet, it is unclear if invader phenology can drive seasonal patterns in indirect effects. We used a field experiment to test if an invasive grass (Imperata cylindrica) caused seasonal indirect effects by altering rodent foraging and seed predation patterns through time. Using seeds from native longleaf pine (Pinus palustris), we found seed predation was 25% greater, on average, in invaded than control plots, but this effect varied by season. Seed predation was 24-157% greater in invaded plots during spring and autumn months, but invasion had no effect on seed predation in other months. One of the largest effects occurred in October when longleaf pine seeds are dispersed, suggesting potential effects on tree regeneration. Thus, seasonal patterns in indirect effects from invaders may cause underappreciated impacts on ecological communities.

Complex networks of parasites and pollinators: moving towards a healthy balance.

Parasites are viewed as a major threat to wild pollinator health. While this may be true for epidemics driven by parasite spillover from managed or invasive species, the picture is more complex for endemic parasites. Wild pollinator species host and share a species-rich, generalist parasite community. In contrast to the negative health impacts that these parasites impose on individual hosts, at a community level they may act to reduce competition from common and abundant pollinator species. By providing rare species with space in which to exist, this will act to support and maintain a diverse and thus healthier pollinator community. At this level, and perhaps paraxodically, parasites may be good for pollinators. This stands in clear contrast to the obvious negative impacts of epidemic and spillover parasites on wild pollinator communities. Research into floral resources that control parasites could be best employed to help design landscapes that provide pollinators with the opportunity to moderate their parasite community, rather than attempting to eliminate specific parasites from wild pollinator communities. This article is part of the theme issue 'Natural processes influencing pollinator health: from chemistry to landscapes'.

Complex community-wide consequences of consumer sexual dimorphism.

Sexual dimorphism is a ubiquitous source of within-species variation, yet the community-level consequences of sex differences remain poorly understood. Here, we analyse a bitrophic model of two competing resource species and a sexually reproducing consumer species. We show that consumer sex differences in resource acquisition can have striking consequences for consumer-resource coexistence, abundance and dynamics. Under both direct interspecific competition and apparent competition between two resource species, sexual dimorphism in consumers' attack rates can mediate coexistence of the resource species, while in other cases can lead to exclusion when stable coexistence is typically expected. Slight sex differences in total resource acquisition also can reverse competitive outcomes and lead to density cycles. These effects are expected whenever both consumer sexes require different amounts or types of resources to reproduce. Our results suggest that consumer sexual dimorphism, which is common, has wide-reaching implications for the assembly and dynamics of natural communities.

Evolution of prudent predation in complex food webs.

Prudent predators catch sufficient prey to sustain their populations but not as much as to undermine their populations' survival. The idea that predators evolve to be prudent has been dismissed in the 1970s, but the arguments invoked then are untenable in the light of modern evolution theory. The evolution of prudent predation has repeatedly been demonstrated in two-species predator-prey metacommunity models. However, the vigorous population fluctuations that these models predict are not widely observed. Here we show that in complex model food webs prudent predation evolves as a result of consumer-mediated ('apparent') competitive exclusion of resources, which disadvantages aggressive consumers and does not generate such fluctuations. We make testable predictions for empirical signatures of this mechanism and its outcomes. Then we discuss how these predictions are borne out across freshwater, marine and terrestrial ecosystems. Demonstrating explanatory power of evolved prudent predation well beyond the question of predator-prey coexistence, the predicted signatures explain unexpected declines of invasive alien species, the shape of stock-recruitment relations of fish, and the clearance rates of pelagic consumers across the latitudinal gradient and 15 orders of magnitude in body mass. Specific research to further test this theory is proposed.

Trophic niche partitioning between two prey and their incidental predators revealed various threats for an endangered species.

Documenting trophic niche partitioning and resource use within a community is critical to evaluate underlying mechanisms of coexistence, competition, or predation. Detailed knowledge about foraging is essential as it may influence the vital rates, which, in turn, can affect trophic relationships between species, and population dynamics. The aims of this study were to evaluate resource and trophic niche partitioning in summer/autumn between the endangered Atlantic-Gaspésie caribou (Rangifer tarandus caribou) population, moose (Alces americanus) and their incidental predators, the black bear (Ursus americanus) and coyote (Canis latrans), and to quantify the extent to which these predators consumed caribou. Bayesian isotopic analysis showed a small overlap in trophic niche for the two sympatric ungulates suggesting a low potential for resource competition. Our results also revealed that caribou occupied a larger isotopic niche area than moose, suggesting a greater diversity of resources used by caribou. Not surprisingly, coyotes consumed mainly deer (Odocoileus virginianus), moose, snowshoe hare (Lepus americanus), and occasionally caribou, while bears consumed mainly vegetation and, to a lesser extent, moose and caribou. As coyotes and bears also feed on plant species, we documented trophic niche overlap between caribou and their predators, as searching for similar resources can force them to use the same habitats and thus increase the encounter rate and, ultimately, mortality risk for caribou. Although the decline in the Gaspésie caribou population is mostly driven by habitat-mediated predation, we found evidence that the low level of resource competition with moose, added to the shared resources with incidental predators, mainly bears, may contribute to jeopardize the recovery of this endangered caribou population. Highlighting the trophic interaction between species is needed to establish efficient conservation and management strategies to insure the persistence of endangered populations. The comparison of trophic niches of species sharing the same habitat or resources is fundamental to evaluate the mechanisms of coexistence or competition and eventually predict the consequences of ecosystem changes in the community.

The Effect of Predator Population Dynamics on Batesian Mimicry Complexes.

AbstractUnderstanding Batesian mimicry is a classic problem in evolutionary biology. In Batesian mimicry, a defended species (the model) is mimicked by an undefended species (the mimic). Prior theories have emphasized the role of predator behavior and learning as well as evolution in model-mimic complexes but have not examined the role of population dynamics in potentially governing the relative abundances and even persistence of model-mimic systems. Here, we examined the effect of the population dynamics of predators and alternative prey on the prevalence of warning-signaling prey composed of models and mimics. Using optimal foraging theory and signal detection theory, we found that the inclusion of predator and alternative prey population dynamics could reverse traditional theoretical predictions: as alternative prey increase in numbers, mimics suffer because larger populations of predators are maintained, resulting in apparent competition. Under some circumstances, apparent competition affects model populations as well, although not as severely as it affects mimics. Our results bear on the intriguing puzzle that in nature warning signals are relatively scarce, yet experiments suggest that such signals can be highly advantageous. The availability of alternative prey and numerical responses by predators can overwhelm advantages observed in experiments to keep warning signals in model-mimic systems relatively scarce.

The Impact of Coinfection Dynamics on Host Competition and Coexistence.

AbstractParasites can mediate competition among host species in an ecological community by differentially affecting key parameters that normally give one species a competitive edge. In nature, however, coinfecting parasites that antagonize or facilitate each other-for example, by altering cross-protective host immune responses-can modulate host infection outcomes and parasite transmission relative to a single infection. Under what conditions is coinfection likely to interfere with parasite-mediated apparent competition among hosts? To address this question, we created a model of two coinfected host species. Parasites could interact indirectly by affecting host reproduction or directly by modulating recovery and disease-induced mortality of each host species to a focal infection. We grounded our model with parameters from a classic apparent competition system but allowed for multiple parasite transmission modes and interaction scenarios. Our results suggest that infection-induced mortality has an outsized effect on competition outcomes relative to recovery but that coinfection-mediated modulation of mortality can produce a range of coexistence or competitive exclusion outcomes. Moreover, while infection prevalence is sensitive to variation in parasite transmission mode, host competitive outcomes are not. Our generalizable model highlights the influence of immunological variation and parasite ecology on community ecology.