Heterospecific Competitors Research Articles

Arbuscular mycorrhizal fungi equalize differences in plant fitness and facilitate plant species coexistence through niche differentiation.

Mycorrhizal fungi are essential to the establishment of the vast majority of plant species but are often conceptualized with contradictory roles in plant community assembly. On the one hand, host-specific mycorrhizal fungi may allow a plant to be competitively dominant by enhancing growth. On the other hand, host-specific mycorrhizal fungi with different functional capabilities may increase nutrient niche partitioning, allowing plant species to coexist. Here, to resolve the balance of these two contradictory forces, we used a controlled greenhouse study to manipulate the presence of two main types of mycorrhizal fungus, ectomycorrhizal fungi and arbuscular mycorrhizal fungi, and used a range of conspecific and heterospecific competitor densities to investigate the role of mycorrhizal fungi in plant competition and coexistence. We find that the presence of arbuscular mycorrhizal fungi equalizes fitness differences between plants and stabilizes competition to create conditions for host species coexistence. Our results show how below-ground mutualisms can shift outcomes of plant competition and that a holistic view of plant communities that incorporates their mycorrhizal partners is important in predicting plant community dynamics.

Assessing biological invasion predatory impacts through interaction strengths and morphological trophic profiling

AbstractBiological invasions are a major stressor on ecosystems worldwide, but tools to predict their predatory impact remain limited. Here, we quantified invader impacts using two complementary approaches: functional responses (to reveal per capita and multiple predator interaction strengths) and ecomorphology (to reveal trophic profiles and competitive overlap). We compared Mozambique tilapia Oreochromis mossambicus, a native southern African cichlid, and a near-trophically analogous invasive congener, the Nile tilapia Oreochromis niloticus. Both Nile tilapia and Mozambique tilapia exhibited a potentially prey population destabilizing Type II functional response. In both single and multiple predator pairings, invasive Nile tilapia had significantly greater prey consumption rates than native Mozambique tilapia, and thereby a greater predatory impact than its native congeneric. Attack rates were greater for Nile tilapia than Mozambique tilapia, with both species showing more similar handling times and maximum feeding rates. No evidence for multiple predator effects was detected within or between these species, and therefore impacts of both species increased additively in the presence of conspecific or heterospecific competitors. Morphological trait analyses found general differences between these two species, with the invasive Nile tilapia having distinctively larger lower jaw closing force, gill resistance and gill raker length, which facilitated greater feeding capacities over the native species. Trophic profiles predicted using morphological trait differences showed high dietary overlap and served as evidence for potential exploitative competition between the two species. These results reveal superior interaction strengths and ecomorphological trait profiles of an invasive over native species which could influence impact and native species replacement dynamics. Novel applications of functional response and ecomorphology provide complementary insights into predatory and competitive impacts from invasive species, aiding impact prediction across environmental contexts.

Adult poison dart frogs avoid potential heterospecific competitors using their sense of smell

AbstractOlfaction is the oldest sense in the animal kingdom. It is used during a multitude of behaviours, such as the encounter of food, the detection of predators, the recognition of habitat‐related cues or the communication with conspecifics. While the use of olfaction and chemical communication has been studied widely in some animals, it is barely known in others. Anurans (frogs and toads), for example, are well known to use acoustic and visual senses, but their chemical sense is still largely understudied. Studies concerning the chemical sense in anurans have been mostly based on the use of semiochemicals in juvenile stages, while the information on adult anurans remains limited. In this study, we analysed the behavioural response of the Neotropical poison frog Ranitomeya sirensis (Sira poison frog, Dendrobatidae) when presented with the odours of prey, novel/prey‐luring fruit, habitat, conspecific faeces and heterospecifics. For this, we offered each of the odours by placing them into one of two testing tubes fixed in an arena, with the other tube left empty as a control. We then measured the time the frogs spent in the vicinity of the odour versus the control tube and calculated a response index. While the frogs did not show a significant avoidance or attraction towards most of the tested odours, they showed a strong response towards the heterospecific odour, which was significantly avoided. This is the first evidence of a poison dart frog responding towards the odours of adult heterospecific frogs. We consider potential reasons for this strong negative reaction, such as the interspecific competition avoidance hypothesis, and discuss our results in the context of other animal species being deterred or attracted by heterospecific chemical cues.

Maternal Care Behavior and Its Consequences in Competition.

Parental care behavior has evolved as a life history strategy to improve reproductive success, particularly in organisms facing challenging environments. However, the variation in maternal care, such as egg-guarding behavior in response to the social environment and the associated ecological consequence of competition, remains largely unknown. This study addresses a gap in current knowledge by examining the plasticity of maternal care behavior in the predatory mite C. eruditus and its impact on offspring survival and intra- and interspecific competition. Our results demonstrated that the reproductive females frequently exhibit egg-guarding behaviors, with enhanced maternal care efforts when the interspecific competitor is present. Egg masses are significantly more vulnerable to predation in the absence of maternal care. Guarding females increased egg survival rates and adversely influenced the survival of both con- and heterospecific competitors, with higher mortality rates being detected. Our findings highlight the ecological significance of maternal care behaviors and suggest that releasing C. eruditus and Neoseiulus cucumeris (Oudemans) together is not recommended for pest management in storage products.

Goliath is not the enemy: Fiddler crab species of similar size are stronger competitors than their larger adversaries in the climate change context

Many intertidal species are expanding their distribution limits towards the poles as a consequence of increased temperature due to climate change. At the same time, some intertidal habitats have been reduced due to rising sea levels. Fiddler crabs inhabit mangroves, and different species occupy well-established zones. Due to the restricted mangrove areas and consequent latitudinal expansion of the species, the distribution and density of fiddler crabs may be subject to change. The overlap forces several species to cohabit in a smaller territorial space. Fiddler crab species can vary in size, which could decisively affect territorial competition. Using Leptuca urguayensis as a reference, we tested how competing species of different sizes (larger size: L. thayeri and Minucapanema; similar size: L. cumulanta and L. leptodactyla) might interact in future climate change scenarios. First, we showed that within the territory of L. uruguayensis, the species L. cumulanta and L. thayeri occurred mainly in the lower mesolittoral zone, while L. leptodactyla and M.panema predominated in the upper mesolittoral zone. In a laboratory experiment, we evaluated whether the density and size of the heterospecific competitors could alter the agonistic behavior of Leptuca uruguayensis. Our results indicated that increasing the density can increase the agonistic behavior of L. uruguayensis with a greater intensity against species of a similar size. In a second field experiment, we demonstrated that the presence of a burrow enhances the intensity of agonistic interactions between L. uruguayensis and species of similar size. Therefore, in a scenario of territorial expansion of fiddler crabs due to climate change, it is expected that similarly sized species represent a greater competitive threat to populations of L. uruguayensis.

Female treefrog preference for breeding sites matches offspring performance in the presence of two anuran competitors.

Preference-performance theory predicts that females should select breeding sites that maximize offspring performance. Amphibians have been a model system for investigating habitat selection, yet most studies have focused on habitat selection in response to predators and conspecifics. We investigated female oviposition site selection and larval performance in eastern gray treefrogs (Hyla versicolor) among pools with and without one of two ecologically distinct heterospecific larvae, the green frog (Rana clamitans) and the American bullfrog (R. catesbeiana). Pools without heterospecifics were used on more nights and had more eggs deposited compared to pools with heterospecifics. In a competition experiment, treefrog larval performance matched this female preference: tadpoles developed faster and grew larger in the absence of either heterospecific. Although we hypothesized that bullfrogs would more strongly affect female treefrog preference and offspring performance because of previous work demonstrating that bullfrogs have stronger negative effects on other tadpoles, both heterospecifics elicited similar responses. The effects of heterospecifics on anuran breeding site selection are understudied and not well understood, and our results demonstrate that female selection of breeding sites is an adaptive behavior for offspring in the presence of heterospecific competitors.

Wolf-Dog-Human: Companionship Based on Common Social Tools.

Wolves, dogs and humans share extremely social and cooperative minds. These similarities are rooted in phylogenetic homology and in the convergence of neuronal and physiological mechanisms, particularly the brain, in the functioning and communication of basic affects and in the mechanisms of stress and calming. The domesticated wolves called dogs are particularly close companion animals. Both Palaeolithic humans and wolves were hypercursorial hunters, cooperating in complex and prosocial ways within their clans with respect to hunting, raising offspring, and defending against conspecific and heterospecific competitors and predators. These eco-social parallels have shaped the development of similar social mindsets in wolves and humans. Over the millennia of domestication, this social match was fine-tuned, resulting in the socio-cognitive specialists humans and dogs, possessing amazingly similar social brains and minds. Therefore, it can be concluded that the quality of their relationships with their human masters is a major factor in the wellbeing, welfare and even health of dogs, as well as in the wellbeing of their human partners. Based on their strikingly similar social brains and physiologies, it can be further concluded that anthropomorphically applying human empathy to dogs in an educated manner may not be as inappropriate as previously thought.

The effect of sociality on competitive interactions among birds.

Sociality can provide many benefits, including increased foraging success, reproductive opportunities and defence against predation. How does sociality influence the dominance hierarchies of ecological competitors? Here, we address this question using a large citizen science dataset of competitive interactions among birds foraging at backyard feeders, representing a network of over 55 000 interactions among 68 common species. We first show that species differ in average group size (the number of conspecifics observed together) as a fundamental measure of sociality. When analysing heterospecific competition, we find that sociality is inversely related to dominance. On average, a single individual from a solitary species is more likely to displace a size-matched opponent than a single individual from a social species. Yet, we find that social species gain an increase in their competitive advantage when in the presence of their conspecifics, which may occur as a result of dynamics within their groups. Finally, we show that more social species have relatively fewer dominance interactions with heterospecifics, and more with conspecifics. Overall, these results demonstrate that sociality can influence competition in ecological networks. More social species have decreased competitive ability as individuals, but they may gain competitive ability in groups.

The effect of heterospecific and conspecific competition on inter-individual differences in tungara frog tadpole (Engystomops pustulosus) behavior.

Repeated social interactions with conspecifics and/or heterospecifics during early development may drive the differentiation of behavior among individuals. Competition is a major form of social interaction and its impacts can depend on whether interactions occur between conspecifics or heterospecifics and the directionality of a response could be specific to the ecological context that they are measured in. To test this, we reared tungara frog tadpoles (Engystomops pustulosus) either in isolation, with a conspecific tadpole or with an aggressive heterospecific tadpole, the whistling frog tadpole (Leptodactylus fuscus). In each treatment, we measured the body size and distance focal E. pustulosus tadpoles swam in familiar, novel and predator risk contexts six times during development. We used univariate and multivariate hierarchical mixed effect models to investigate the effect of treatment on mean behavior, variance among and within individuals, behavioral repeatability and covariance among individuals in their behavior between contexts. There was a strong effect of competition on behavior, with different population and individual level responses across social treatments. Within a familiar context, the variance in the distance swam within individuals decreased under conspecific competition but heterospecific competition caused more variance in the average distance swam among individuals. Behavioral responses were also context specific as conspecific competition caused an increase in the distance swam within individuals in novel and predator risk contexts. The results highlight that the impact of competition on among and within individual variance in behavior is dependent on both competitor species identity and context.

Integrating eco-evolutionary dynamics and modern coexistence theory.

Community ecology typically assumes that competitive exclusion and species coexistence are unaffected by evolution on the time scale of ecological dynamics. However, recent studies suggest that rapid evolution operating concurrently with competition may enable species coexistence. Such findings necessitate general theory that incorporates the coexistence contributions of eco-evolutionary processes in parallel with purely ecological mechanisms and provides metrics for quantifying the role of evolution in shaping competitive outcomes in both modelling and empirical contexts. To foster the development of such theory, here we extend the interpretation of the two principal metrics of modern coexistence theory-niche and competitive ability differences-to systems where competitors evolve. We define eco-evolutionary versions of these metrics by considering how invading and resident species adapt to conspecific and heterospecific competitors. We show that the eco-evolutionary niche and competitive ability differences are sums of ecological and evolutionary processes, and that they accurately predict the potential for stable coexistence in previous theoretical studies of eco-evolutionary dynamics. Finally, we show how this theory frames recent empirical assessments of rapid evolution effects on species coexistence, and how empirical work and theory on species coexistence and eco-evolutionary dynamics can be further integrated.

The effects of intra- and interspecific competitions on personality and individual plasticity in two sympatric brown frogs

We studied how individuals modify their behavior in response to inter- and intraspecific competitors and how these changes affected the pattern of variation between populations and species. As study models, we used tadpoles of two brown frogs, Rana latastei and R. dalmatina. Since R. latastei is always sympatric to R. dalmatina, whereas R. dalmatina is sympatric to R. latastei only in the periphery of its range, we predicted a stronger response to heterospecifics in R. latastei than in R. dalmatina and, within each species, in syntopic than in allotopic populations. To test these predictions, we raised tadpoles, from either syntopic or allotopic populations, in either syntopy or allotopy and repeatedly tested them in open field trials in the presence of a caged conspecific, a caged heterospecific, or an empty cage. As predicted, we found that, on average, R. latastei tadpoles modified their behavior across treatments more than R. dalmatina tadpoles and individuals from the syntopic population changed more than their conspecifics from the allotopic population. In both species, the pattern of variation at the individual level mirrored that at the population and species levels providing no evidence for an individual-by-environment interaction (Itimes E). Besides these differences, however, individuals of the two species also showed unpredicted and context-independent behavioral differences, suggesting that there might be more to interspecific behavioral variation than the effect of selection by heterospecific competitors.Significance statementDoes the distribution range of a species influence the evolution of plastic behaviors to heterospecific competitors? And how do differences in plasticity affect animal personality? To answer these questions, we raised tadpoles of two brown frog species, Rana dalmatina and R. latastei, and studied how the amount and the type of their swimming varied with the presence of the other species. R. latastei, whose small distribution range fully overlaps with that of R. dalmatina, plastically responds to it, whereas R. dalmatina, which is sympatric to R. latastei only in the periphery of its broader range, does not. These interspecific differences mirrored those among individuals: tadpoles of both species show repeatable behaviors, but only those of R. latastei plastically changed their behavior with the presence of the other species; however, neither R. latastei nor R. dalmatina show among-individual variation in plasticity.

Climate warming may weaken stabilizing mechanisms in old forests

AbstractPlant competition may intensify with climate warming, but whether this will occur equally for conspecific and heterospecific competition remains unknown. Competitive shifts have the potential to instigate community change because the relative strengths of conspecific and heterospecific negative density dependence mediate the stabilizing mechanisms underpinning species coexistence. We examined a mature temperate forest to assess both direct and indirect climate effects at multiple scales: individual species, interspecies relationships, and community stability mechanisms. Our coupled approach (1) quantified tree mortality risk dependence on the interactive effects of competition, climatic water deficit, snowpack, and soil moisture for 28,913 trees over 8 years (3149 mortalities), then (2) used a climate‐projection ensemble to forecast changes in conspecific and heterospecific competition from 2020 to 2100. We predict that projected climate warming will destabilize the foundational forest community by increasing the strength of heterospecific competition at a greater rate and to a greater degree than conspecific competition for four of five abundant tree species, particularly on dry microsites. Modeling showed that these findings were most pronounced after the year 2038, at which point snowpacks were projected to be too small to ameliorate the effects of drought on competitive interactions. Our finding that heterospecific competition is more sensitive than conspecific competition to climate warming may indicate the impending loss of ecosystem functioning. We join the growing body of work showing a predominance of indirect drought effects, yet coupled climate models still fail to consider how changing community dynamics may impact forest cover and, in turn, disrupt forest–climate carbon feedbacks. Ecosystems sharing characteristics with our example forest—those with low species richness and therefore a limited biodiversity insurance effect—may be similarly vulnerable to climate‐mediated destabilization. In such communities, increased heterospecific competition among even a small number of species can more easily destabilize communities without recourse from redundant species. This study of an overlooked but vital mechanism of community change can be adapted by research in a range of ecosystems to improve the understanding of climate change consequences.

An observational analysis of Canada Jay (Perisoreus canadensis) foraging and caching ecology in Denali National Park and Preserve, Alaska, USA

Arctic and subarctic wildlife are among the most vulnerable species to climate change. Canada Jays (Perisoreus canadensis (Linnaeus, 1776)) are generalist residents of northern boreal forests and scatter-hoard food to insulate against food scarcity during winter. Unlike most scatter-hoarders, however, Canada Jays primarily cache perishable food, rendering their caches more susceptible to climate change induced degradation and loss. Here we use a mostly noninvasive approach to document Canada Jay foraging ecology among a population in interior Alaska, USA, including the types of food acquired, foraging and caching rates, and cache longevity and loss. We also tested for associations between foraging and caching rates with reproductive metrics to assess possible relationships among food and productivity. We found that Canada Jays have a varied diet that changed seasonally, and responded to a record-setting warm spring by directing foraging efforts away from cache recovery and towards the emergence of fresh food. We did not find evidence for relationships between foraging and caching rate with reproductive output, possibly owing to small sample sizes. We found that caches were recovered quickly (<4 weeks) and frequently lost to conspecific and heterospecific competitors. Our study suggests that Canada Jays may be better poised to respond to changes in cache integrity and food availability than has been previously recognized.

Discrimination by Ganaspis pelleranoi of Ceratitis capitata larvae previously parasitised by Diachasmimorpha longicaudata

ABSTRACT Heterospecific and conspecific host discrimination, and super- and multiparasitism by the neotropical figitid Ganaspis pelleranoi were determined using 6-d-old Ceratitis capitata larvae in choice trials, where alternatives such as hosts previously parasitised by the exotic braconid Diachasmimorpha longicaudata, or by conspecific G. pelleranoi, and/or unparasitised hosts were available. Choice assays involved both experienced and naïve figitid females, previously host-exposed and non-host-exposed, respectively. Multiparasitism was mainly avoided if the G. pelleranoi female had a choice between unparasitised and parasitised larvae. Heterospecific host discrimination was highly improved by ovipositional experience. A very limited number of experienced G. pelleranoi females visited, probed, or oviposited host larvae previously parasitised by D. longicaudata. Only 2% of figitid adults were recovered from puparia resulting from larvae previously parasitised by D. longicaudata and exposed to naïve females. No figitid adults were recovered in those treatments involving experienced females. Latency and host examination data, number of visits, probing and oviposition’s activities showed that experienced G. pelleranoi female has a higher propensity to forage unparasitised host larvae rather than larvae previously parasitised by conspecific females. Self-superparasitism was greater than conspecific superparasitism when figited females had to choose between unparasitised and conspecific parasitised larvae. The ability of G. pelleranoi females to discriminate previously parasitised medfly larvae suggests low levels of both heterospecific and conspecific competition. Combined releases of D. longicaudata and G. pelleranoi in Argentinean fruit-growing regions could be a more advantageous alternative than individual species releases.

Exotic predators are not better biocontrol agents: the harlequin ladybird is not the most voracious in Mexico.

The use of exotic species for pest biocontrol has been a common pathway for introduction and dispersal of invasive species that may have undesired outcomes. Biocontrol agents are believed to be a less damaging alternative than pesticides, but some species may also prey on or parasitize native species or outcompete them for resources. The harlequin ladybird (Harmonia axyridis) is a well-known biocontrol agent originally from Asia that has established invasive populations in 59 countries around the globe. Harlequin ladybirds are generalist predators that in addition to pests prey on an array of different species including other coccinelids’ eggs and larvae. In Mexico, native ladybirds that share ecological requirements with harlequin ladybirds are at risk of being outcompeted and predated upon. The aim of our study was to compare the foraging efficiency of harlequin ladybirds against three species of native coccinelids when preying on aphids. We investigated the foraging behaviour of ladybirds alone and in pairs with a conspecific, a native heterospecific or an exotic heterospecific. We found that the native Cycloneda sanguinea was the species that consumed the most aphids, while Hippodamia convergens was the fastest to find and consume each aphid. Harlequin ladybirds and H. convergens consumed the same number of aphids while P. vittigera consumed less. Conspecific competition was stronger than heterospecific competition. We discuss the suitability of using the exotic invasive harlequin ladybird for aphid biocontrol in comparison with native coccinelids.

Combining social information use and comfort seeking for nest site selection in a cavity-nesting raptor

When selecting a breeding site, individuals can use social information to reduce the uncertainty regarding habitat quality. Individuals of several bird species tend to reuse nests previously occupied by conspecific or heterospecific competitors but the proximate mechanisms underlying this behaviour remain unclear. Reoccupying nests previously used by competitors could result from individuals copying competitors' choices (the ‘social information’ hypothesis). Alternatively, it could allow individuals to fulfil their need for a soft nest substrate (e.g. by improving thermal insulation or reducing egg breakage risks) at low costs, regardless of previous occupancy (the ‘comfort’ hypothesis). Here, we aimed to determine which of these nonmutually exclusive mechanisms triggered the preference for old conspecific nest material in a secondary cavity-nesting raptor that does not add lining material to its nests, the lesser kestrel, Falco naumanni . Using an experimental design forcing settling lesser kestrels to choose between two adjacent nestboxes containing different substrates, we detected a strong preference for soft substrates (peat moss or old conspecific or European roller, Coracias garrulus , nest material) over coarse mineral substrate, especially when the soft substrate also provided social information about previous nest use by a competitor. Despite the apparent absence of preference when directly comparing settlement patterns in soft substrates with and without social information, early settling individuals favoured the substrate with social information, while late settling ones favoured the substrate without social information. This could reflect intraspecific competition avoidance by late arriving individuals that may be competitively inferior to early arriving ones. This hypothesis is supported by a later laying date of young breeders in our population. Our findings suggest that both comfort seeking and social information use explain the preference for previously used nest cavities, and that nest site choices may depend on individual competitive abilities and experience. • The proximate mechanisms underlying the reuse of old nest material are diverse. • Lesser kestrels preferred soft nest substrates, with or without social cues. • Between two soft substrates, early breeders favoured the one providing social cues. • Late breeders, mostly young, preferred soft substrates with no social cues. • Lesser kestrels may adjust social information use according to individual traits.

Ecological and behavioral mechanisms of density‐dependent habitat expansion in a recovering African ungulate population

AbstractMajor disturbances can temporarily remove factors that otherwise constrain population abundance and distribution. During such windows of relaxed top‐down and/or bottom‐up control, ungulate populations can grow rapidly, eventually leading to resource depletion and density‐dependent expansion into less‐preferred habitats. Although many studies have explored the demographic outcomes and ecological impacts of these processes, fewer have examined the individual‐level mechanisms by which they occur. We investigated these mechanisms in Gorongosa National Park, where the Mozambican Civil War devastated large‐mammal populations between 1977 and 1992. Gorongosa’s recovery has been marked by proliferation of waterbuck (Kobus ellipsiprymnus), an historically marginal 200‐kg antelope species, which is now roughly 20‐fold more abundant than before the war. We show that after years of unrestricted population growth, waterbuck have depleted food availability in their historically preferred floodplain habitat and have increasingly expanded into historically avoided savanna habitat. This expansion was demographically skewed: mixed‐sex groups of prime‐age individuals remained more common in the floodplain, while bachelors, loners, and subadults populated the savanna. By coupling DNA metabarcoding and forage analysis, we show that waterbuck in these two habitats ate radically different diets, which were more digestible and protein‐rich in the floodplain than in savanna; thus, although individuals in both habitats achieved positive net energy balance, energetic performance was higher in the floodplain. Analysis of daily activity patterns from high‐resolution GPS‐telemetry, accelerometry, and animal‐borne video revealed that savanna waterbuck spent less time eating, perhaps to accommodate their tougher, lower‐quality diets. Waterbuck in savanna also had more ectoparasites than those in the floodplain. Thus, plasticity in foraging behavior and diet selection enabled savanna waterbuck to tolerate the costs of density‐dependent spillover, at least in the short term; however, the already poorer energetic performance of these individuals implies that savanna occupancy may become prohibitively costly as heterospecific competitors and predators continue to recover in Gorongosa. Our results suggest that behavior can provide a leading indicator of the onset of density‐dependent limitation and the likelihood of subsequent population decline, but that reliable inference hinges on understanding the mechanistic basis of observed behavioral shifts.

Heterospecific competitors and seasonality can affect host physiology and behavior: key factors in disease transmission

AbstractEcological and environmental factors can influence the transmission of infectious diseases. They can accomplish this via effects on host susceptibility and exposure to infection, which are governed by host physiology and behavior, respectively. To better inform disease control, more information is needed about how extrinsic factors affect physiological and behavioral processes that determine transmission. We investigated how heterospecific competitors and seasonality may influence host susceptibility and intraspecific contact rates using a directly transmitted disease system, the North American deer mouse (Peromyscus maniculatus)—Sin Nombre hantavirus (SNV) system. In grasslands of western Montana, USA, deer mice compete with dominant voles (Microtus spp.) and shrews (Sorex spp.) and experience a seasonal temperate climate. Higher SNV transmission occurs primarily during spring/summer, when changes in physiology and behavior may serve as influential contributors. We hypothesized that (1) voles, and to a lesser extent shrews, will induce chronic stress, suppress immunity, and may change contact rates of deer mice; and (2) during spring/summer, deer mice may experience chronic stress, suppressed immunity, and higher contact rates, which may help explain the reported seasonality in SNV transmission. Over two years, we trapped small mammals at four grids in western Montana. Deer mice were sampled for feces and blood and evaluated for scar numbers, demography, and body condition scores (BCSs). We evaluated stress physiology with fecal corticosterone metabolites (FCMs), neutrophil/lymphocyte (N/L) ratios and BCSs, immunity with white blood cell (WBC) counts, and contact rates with scar numbers. We found that shrew density was negatively associated with stress response FCMs, suggestive of chronic stress. Additionally, although complex interactions existed, shrew and vole densities were negatively associated with BCSs, but differentially with scar numbers. N/L ratios were higher in spring/summer, whereas WBC counts were lower in summer, suggestive of chronic stress and suppressed immunity, respectively. Our results suggest that (1) heterospecific competitors may differentially influence disease transmission via stress physiology and contact rates, and that (2) chronic stress, suppressed immunity, and higher contact rates may help explain why higher SNV transmission has been previously reported during spring/summer in Montana. Our findings may extend to other directly transmitted disease systems.

Frequency-dependent tree growth depends on climate.

Climate and competition interact to affect species' performance, such as growth and survival, and help determine species distributions and coexistence. However, it is unclear how climatic conditions modulate frequency-dependent performance, that is, how performance changes as a species becomes locally rare or common. This is critical because declines in performance as a species becomes more common (negative frequency dependence) is a signature of niche differences among species that stabilize coexistence, whereas positive frequency dependence leads to priority effects and hampers species coexistence. Here, we used dendrochronology and hierarchical models to test whether frequency-dependent growth of sugar pine (Pinus lambertiana) depends on climatic conditions. We found that growth rates were strongly dependent on annual precipitation, but no frequency dependence was evident across all years. However, there was a strong interaction between precipitation and frequency dependence, revealing stabilizing niche differences in dry years but positive frequency dependence in wet years. These differences emerged because of precipitation-driven changes in the direction and strength of both con- and heterospecific competition. Overall, these results show how stabilizing and destabilizing effects can be temporally dynamic for long-lived species and interact with climate variation.

Aphid alarm pheromone alters larval behaviour of the predatory gall midge, Aphidoletes aphidimyza and decreases intraguild predation by anthocorid bug, Orius laevigatus.

Intraguild predation is the killing and consuming of a heterospecific competitor that uses similar resources as the prey, and also benefit from preying on each other. We investigated the foraging behaviour of the gallmidge, Aphidoletes aphidimyza, a predator of aphids used for biological control that is also the intraguild prey for most other aphid natural enemies. We focus on how aphid alarm pheromone can alter the behaviour of the gallmidge, and predation by the anthocorid bug Orius laevigatus (O. laevigatus). We hypothesised that gallmidges would respond to the presence of (E)-β-farnesene (EBF) by leaving the host plant. Since feeding by Aphidoletes gallmidge larvae does not induce EBF emission by aphids, this emission indicates the presence of an intraguild predator. We found that gallmidge larvae reduced their foraging activities and left the plant earlier when exposed to EBF, particularly when aphids were also present. Contrastingly, gallmidge females did not change the time visiting plants when exposed to EBF, but lay more eggs on plants that had a higher aphid density. Lastly, EBF reduced the number of attacks of the intraguild predator, O. laevigatus, on gallmidge larvae, potentially because more gallmidges stopped aphid feeding and moved off the plant at which point O. laevigatus predated on aphids. Our work highlights the importance of understanding how intraguild predation can influence the behaviour of potential biological control agents and the impact on pest control services when other natural enemies are also present.