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Showing papers in "Oecologia in 2019"


Journal ArticleDOI
TL;DR: This work focuses on spiders as a model group of generalist predators, among the most abundant and most diverse natural enemies in agroecosystems, and reviews their functional traits that influence food-web dynamics and pest suppression at organisational levels ranging from individuals to communities.
Abstract: The role of generalist predators in biological control remains controversial as they may not only reduce pest populations but also disrupt biocontrol exerted by other natural enemies. Here, we focus on spiders as a model group of generalist predators. They are among the most abundant and most diverse natural enemies in agroecosystems. We review their functional traits that influence food-web dynamics and pest suppression at organisational levels ranging from individuals to communities. At the individual and population levels, we focus on hunting strategy, body size, life stage, nutritional target, and personality (i.e., consistent inter-individual differences in behaviour). These functional traits determine the spider trophic niches. We also focus on the functional and numerical response to pest densities and on non-consumptive effects of spiders on pests. At the community level, we review multiple-predator effects and effect of alternative prey on pest suppression. Evidence for a key role of spiders in pest suppression is accumulating. Importantly, recent research has highlighted widespread non-consumptive effects and complex intraguild interactions of spiders. A better understanding of these effects is needed to optimize biocontrol services by spiders in agroecosystems.

108 citations


Journal ArticleDOI
TL;DR: Evidence for personality-dependent space use, movement, and occupation of individual spatial niches in bank voles is found, besides dietary niche specialization also spatial dimensions of ecological niches vary among individuals within populations, which may have important consequences for ecological interactions within- and between species.
Abstract: Personality-dependent space use and movement might be crucially influencing ecological interactions, giving way to individual niche specialization. This new approach challenges classical niche theory with potentially great ecological consequences, but so far has only scarce empirical support. Here, we investigated if and how consistent inter-individual differences in behavior predict space use and movement patterns in free-ranging bank voles (Myodes glareolus) and thereby contribute to individual niche specialization. Individuals were captured and marked from three different subpopulations in North-East Germany. Inter-individual differences in boldness and exploration were quantified via repeated standardized tests directly in the field after capture. Subsequently, space use and movement patterns of a representative sample of the behavioral variation (n = 21 individuals) were monitored via automated VHF telemetry for a period of four days, yielding on average 384 locations per individual. Bolder individuals occupied larger home ranges and core areas (estimated via kernel density analyses), moved longer distances, spatially overlapped with fewer conspecifics and preferred different microhabitats based on vegetation cover compared to shyer individuals. We found evidence for personality-dependent space use, movement, and occupation of individual spatial niches in bank voles. Thus, besides dietary niche specialization also spatial dimensions of ecological niches vary among individuals within populations, which may have important consequences for ecological interactions within- and between species.

81 citations


Journal ArticleDOI
TL;DR: The notion that temperature is a major determinant of body condition for arid-zone birds is supported, as the strong temperature dependence of foraging success and body condition among male hornbills provisioning nests raises the possibility that male behavioural trade-offs translate into equally strong effects of hot weather on female condition and nest success.
Abstract: Avian responses to high environmental temperatures include retreating to cooler microsites and/or increasing rates of evaporative heat dissipation via panting, both of which may affect foraging success. We hypothesized that behavioural trade-offs constrain the maintenance of avian body condition in hot environments, and tested predictions arising from this hypothesis for male Southern Yellow-billed Hornbills (Tockus leucomelas) breeding in the Kalahari Desert. Operative temperatures experienced by the hornbills varied by up to 13 °C among four microsite categories used by foraging males. Lower prey capture rates while panting and reductions associated with the occupancy of off-ground microsites, resulted in sharp declines in foraging efficiency during hot weather. Consequently, male body mass (Mb) gain between sunrise and sunset decreased with increasing daily maximum air temperature (Tmax), from ~ 5% when Tmax < 25 °C to zero when Tmax = 38.4 °C. Overnight Mb loss averaged ~ 4.5% irrespective of Tmax, creating a situation where nett 24-h Mb loss approached 5% on extremely hot days. These findings support the notion that temperature is a major determinant of body condition for arid-zone birds. Moreover, the strong temperature dependence of foraging success and body condition among male hornbills provisioning nests raises the possibility that male behavioural trade-offs translate into equally strong effects of hot weather on female condition and nest success. Our results also reveal how rapid anthropogenic climate change is likely to substantially decrease the probability of arid-zone birds like hornbills being able to successfully provision nests while maintaining their own condition.

65 citations


Journal ArticleDOI
TL;DR: This study shows that an ambush predator’s temporal activity and space use patterns interact to create diel refugia and shape the antipredator behaviors of its prey.
Abstract: The landscape of fear is an important driver of prey space use. However, prey can navigate the landscape of fear by exploiting temporal refuges from predation risk. We hypothesized that diel patterns of predator and prey movement and space use would be inversely correlated due to temporal constraints on predator habitat domain. Specifically, we evaluated habitat selection and activity of the vicuna and its only predator, the puma, during three diel periods: day, dawn/dusk, and night. Pumas selected the same habitats regardless of diel period—vegetated and rugged areas that feature stalking cover for pumas—but increased their activity levels during dawn/dusk and night when they benefit from reduced detection by prey. Vicunas avoided areas selected by pumas and reduced activity at night, but selected vegetated areas and increased activity by day and dawn/dusk. Vicuna habitat selection and movement strategies appeared to reduce the risk of encountering pumas; movement rates of pumas and vicunas were negatively correlated across the diel cycle, and habitat selection was negatively correlated during dawn/dusk and night. Our study shows that an ambush predator’s temporal activity and space use patterns interact to create diel refugia and shape the antipredator behaviors of its prey. Importantly, it is likely the very nature of ambush predators’ static habitat specificity that makes predator activity important to temporally varying perceptions of risk. Prey which depend on risky habitats for foraging appear to mitigate risk by feeding when they can more easily detect predators and when predators are least active.

62 citations


Journal ArticleDOI
TL;DR: Robust evidence is presented that N deposition threatened biodiversity by reducing both species richness and evenness, a process which could be alleviated by mowing.
Abstract: Increasing availability of reactive nitrogen (N) threatens plant diversity in diverse ecosystems. While there is mounting evidence for the negative impacts of N deposition on one component of diversity, species richness, we know little about its effects on another one, species evenness. It is suspected that ecosystem management practice that removes nitrogen from the ecosystem, such as hay-harvesting by mowing in grasslands, would mitigate the negative impacts of N deposition on plant diversity. However, empirical evidence is scarce. Here, we reported the main and interactive effects of N deposition and mowing on plant diversity in a temperate meadow steppe with 4-year data from a field experiment within which multi-level N addition rates and multiple N compounds are considered. Across all the types of N compounds, species richness and evenness significantly decreased with the increases of N addition rate, which was mainly caused by the growth of a tall rhizomatous grass, Leymus chinensis. Such negative impacts of N addition were accumulating with time. Mowing significantly reduced the dominance of L. chinensis, and mitigated the negative impacts of N deposition on species evenness. We present robust evidence that N deposition threatened biodiversity by reducing both species richness and evenness, a process which could be alleviated by mowing. Our results highlight the changes of species evenness in driving the negative impacts of N deposition on plant diversity and the role of mowing in mediating such negative impacts of N deposition.

58 citations


Journal ArticleDOI
TL;DR: It is concluded that a clear understanding of the effects of habitat loss and fragmentation on functional diversity will improve predictions of the resiliency and resistance of plant communities to varying scales of disturbance.
Abstract: Habitat loss and fragmentation result in significant landscape changes that ultimately affect plant diversity and add uncertainty to how natural areas will respond to future global change. This uncertainty is important given that the loss of biodiversity often includes losing key ecosystem functions. Few studies have explored the effects of landscape changes on plant functional diversity and evidence so far has shown far more pervasive effects than previously reported by species richness and composition studies. Here we present a review on the impact of habitat loss and fragmentation on (1) individual functional traits—related to persistence, dispersal and establishment—and (2) functional diversity. We also discuss current knowledge gaps and propose ways forward. From the literature review we found that studies have largely focused on dispersal traits, strongly impacted by habitat loss and fragmentation, while traits related to persistence were the least studied. Furthermore, most studies did not distinguish habitat loss from spatial fragmentation and were conducted at the plot or fragment-level, which taken together limits the ability to generalize the scale-dependency of landscape changes on plant functional diversity. For future work, we recommend (1) clearly distinguishing the effects of habitat loss from those of fragmentation, and (2) recognizing the scale-dependency of predicted responses when functional diversity varies in time and space. We conclude that a clear understanding of the effects of habitat loss and fragmentation on functional diversity will improve predictions of the resiliency and resistance of plant communities to varying scales of disturbance.

46 citations


Journal ArticleDOI
TL;DR: Overall, plant growth was promoted and flowering was accelerated upon exposure to fungal volatiles, irrespectively of fungal CO2 emission levels, and data indicate that plant development and resistance can be modulated in response to exposure tofungalvolatiles.
Abstract: Plants are ubiquitously exposed to a wide diversity of (micro)organisms, including mutualists and antagonists. Prior to direct contact, plants can perceive microbial organic and inorganic volatile compounds (hereafter: volatiles) from a distance that, in turn, may affect plant development and resistance. To date, however, the specificity of plant responses to volatiles emitted by pathogenic and non-pathogenic fungi and the ecological consequences of such responses remain largely elusive. We investigated whether Arabidopsis thaliana plants can differentiate between volatiles of pathogenic and non-pathogenic soil-borne fungi. We profiled volatile organic compounds (VOCs) and measured CO2 emission of 11 fungi. We assessed the main effects of fungal volatiles on plant development and insect resistance. Despite distinct differences in VOC profiles between the pathogenic and non-pathogenic fungi, plants did not discriminate, based on plant phenotypic responses, between pathogenic and non-pathogenic fungi. Overall, plant growth was promoted and flowering was accelerated upon exposure to fungal volatiles, irrespectively of fungal CO2 emission levels. In addition, plants became significantly more susceptible to a generalist insect leaf-chewing herbivore upon exposure to the volatiles of some of the fungi, demonstrating that a prior fungal volatile exposure can negatively affect plant resistance. These data indicate that plant development and resistance can be modulated in response to exposure to fungal volatiles.

43 citations


Journal ArticleDOI
TL;DR: Assessment of wild dogs’ potential to adapt to high ambient temperatures through flexible timing of hunting behavior suggested that nocturnal activity helps this cursorial hunter to cope with high daytime temperatures, but wild dogs appear not to use this coping strategy when they are raising pups, suggesting that their resource needs may not be fulfilled during the pup-rearing period.
Abstract: Climate change is widely accepted to be one of the greatest threats to species globally. Identifying the species most at risk is, therefore, a conservation priority. Some species have the capacity to adapt to rising temperatures through changing their phenology, behavior, distribution, or physiology, and, therefore, may be more likely to persist under rising temperatures. Recent findings suggest that the African wild dog Lycaon pictus may be impacted by climate change, since reproductive success is consistently lower when pup-rearing coincides with periods of high ambient temperature. We used GPS collars, combined with generalized linear mixed-effects models, to assess wild dogs’ potential to adapt to high ambient temperatures through flexible timing of hunting behavior. On days with higher maximum temperatures, wild dogs showed lower daytime activity and greater nocturnal activity, although nocturnal activity did not fully balance the decrease in daytime activity, particularly during the denning period. Increases in nocturnal activity were confined mainly to moonlit nights, and were seldom observed when packs were raising pups. Our findings suggest that nocturnal activity helps this cursorial hunter to cope with high daytime temperatures. However, wild dogs appear not to use this coping strategy when they are raising pups, suggesting that their resource needs may not be fulfilled during the pup-rearing period. Given that moonlight availability—which will not change as the climate changes—constrains wild dogs’ nocturnal activity, the species may have insufficient behavioral plasticity to mitigate increasing diurnal temperatures. These findings raise concerns about climate change impacts on this endangered species, and highlight the need for behavior to be considered when assessing species’ vulnerability to climate change.

40 citations


Journal ArticleDOI
TL;DR: The results indicate that induction of ethylene signaling by potyviruses mediates aphid attraction to infected plants and virus spread; however, additional factors may contribute to plant–vector dynamics in complex communities.
Abstract: Plant pathogens can influence host characteristics such as volatile emissions, nutrient composition or plant color, modulating vector and non-vector insect dynamics in the ecosystem. While previous research has focused on insect attraction and dispersal to infected plants, little is known about mechanisms mediating these interactions. Here, we investigate the role of ethylene in green peach aphid (Myzus persicae) attraction to potyvirus-infected plants. In our experiments, we utilized two different potyviruses, Potato virus Y (PVY) and Turnip mosaic virus, in lab and field experiments. Consistent with previous studies, we show that greater numbers of aphids settle on potyvirus-infected plants in the lab and greater numbers of aphids are found in PVY-infected potato (Solanum tuberosum) fields compared to controls. In laboratory experiments, inhibition of ethylene signaling in plants either chemically or genetically prevented aphids from preferentially settling on potyvirus-infected plants. Virus spread was reduced in lab arenas by over 80% when ethylene signaling was inhibited chemically. Despite this, ethylene inhibition had no significant impact on virus spread in field mesocosms. Our results indicate that induction of ethylene signaling by potyviruses mediates aphid attraction to infected plants and virus spread; however, additional factors may contribute to plant-vector dynamics in complex communities. Specific components of ethylene signaling may be important targets for future management of vector-borne viruses and research on mechanisms mediating plant-vector-virus interactions.

36 citations


Journal ArticleDOI
TL;DR: It is suggested that continued uneven seasonal warming will advance cessation of primary growth in autumn, even when leaf senescence is delayed, as well as the need to consider differences among plant functional types when forecasting the future behaviour of ecosystems.
Abstract: Ongoing global warming is causing phenological shifts that affect photosynthesis and growth rates in temperate woody species. However, the effects of seasonally uneven climate warming—as is occurring in much of Europe, where the winter/spring months are warming twice as fast than the summer/autumn months—on autumn growth cessation (completion of overwintering buds) and leaf senescence, and possible carry-over effects between phenophases, remain under-investigated. We conducted experiments in which we exposed saplings of canopy and understory species to 4 °C warming in winter/spring, summer/autumn, or all year to disentangle how the timing of bud break, bud set completion, and leaf senescence is affected by seasonally uneven warming. All-year warming led to significantly delayed leaf senescence, but advanced bud set completion; summer/autumn warming only delayed leaf senescence; and winter/spring warming advanced both bud set and senescence. The non-parallel effects of warming on bud completion and leaf senescence show that leaf senescence alone is an inadequate proxy for autumn growth cessation in trees and counterintuitively suggest that continued uneven seasonal warming will advance cessation of primary growth in autumn, even when leaf senescence is delayed. Phenological responses to warming treatments (earlier spring onset, later autumn senescence) were more than twice as high in understory species than in canopy species, which can partly be explained by the absence of carry-over effects among phenophases in the former group. This underscores the need to consider differences among plant functional types when forecasting the future behaviour of ecosystems.

35 citations


Journal ArticleDOI
TL;DR: The validation of the osmometer method in an herb-dominated ecosystem paves the way for rapid community-scale surveys of drought tolerance across plant functional groups, which could improve trait-based predictions of ecosystem responses to climate change.
Abstract: Community-scale surveys of plant drought tolerance are essential for understanding semi-arid ecosystems and community responses to climate change. Thus, there is a need for an accurate and rapid methodology for assessing drought tolerance strategies across plant functional types. The osmometer method for predicting leaf osmotic potential at full turgor (πo), a key metric of leaf-level drought tolerance, has resulted in a 50-fold increase in the measurement speed of this trait; however, the applicability of this method has only been tested in woody species and crops. Here, we assess the osmometer method for use in herbaceous grassland species and test whether πo is an appropriate plant trait for understanding drought strategies of herbaceous species as well as species distributions along climate gradients. Our model for predicting leaf turgor loss point (πTLP) from πo (πTLP = 0.80πo–0.845) is nearly identical to the model previously presented for woody species. Additionally, πo was highly correlated with πTLP for graminoid species (πtlp = 0.944πo–0.611; r2 = 0.96), a plant functional group previously flagged for having the potential to cause erroneous measurements when using an osmometer. We report that πo, measured with an osmometer, is well correlated with other traits linked to drought tolerance (namely, leaf dry matter content and leaf vulnerability to hydraulic failure) as well as climate extremes linked to water availability. The validation of the osmometer method in an herb-dominated ecosystem paves the way for rapid community-scale surveys of drought tolerance across plant functional groups, which could improve trait-based predictions of ecosystem responses to climate change.

Journal ArticleDOI
TL;DR: The relationships between the estimated performance regimes and environmental temperature limits suggest that animals in confined habitats, including many juvenile elasmobranchs within nursery habitats, are likely to experience a reduction of performance under a warming climate, although their flatter thermal response will likely dampen this impact.
Abstract: Temperature is one of the most influential drivers of physiological performance and behaviour in ectotherms, determining how these animals relate to their ecosystems and their ability to succeed in particular habitats. Here, we analysed the largest set of acceleration data compiled to date for elasmobranchs to examine the relationship between volitional activity and temperature in 252 individuals from 8 species. We calculated activation energies for the thermal performance response in each species and estimated optimum temperatures using an Arrhenius breakpoint analysis, subsequently fitting thermal performance curves to the activity data. Juveniles living in confined nursery habitats not only spent substantially more time above their optimum temperature and at the upper limits of their performance breadths compared to larger, less site-restricted animals, but also showed lower activation energies and broader performance curves. Species or life stages occupying confined habitats featured more generalist behavioural responses to temperature change, whereas wider ranging elasmobranchs were characterised by more specialist behavioural responses. The relationships between the estimated performance regimes and environmental temperature limits suggest that animals in confined habitats, including many juvenile elasmobranchs within nursery habitats, are likely to experience a reduction of performance under a warming climate, although their flatter thermal response will likely dampen this impact. The effect of warming on less site-restricted species is difficult to forecast since three of four species studied here did not reach their optimum temperature in the wild, although their specialist performance characteristics may indicate a more rapid decline should optimum temperatures be exceeded.

Journal ArticleDOI
TL;DR: This study predicts southern stingrays in marine reserves would reduce the use of deep forereef habitats in the favor of shallow flats where the risk of shark encounters is lower and provides evidence that roving predators can induce pronounced habitat shifts in prey that rely on crypsis and refuging, rather than active escape, in high-visibility, heterogeneous marine habitats.
Abstract: Emerging conservation efforts for the world’s large predators may, if successful, restore natural predator–prey interactions. Marine reserves, where large predators tend to be relatively common, offer an experimental manipulation to investigate interactions between large-bodied marine predators and their prey. We hypothesized that southern stingrays—large, long-lived and highly interactive mesopredators—would invest in anti-predator behavior in marine reserves where predatory large sharks, the primary predator of stingrays, are more abundant. Specifically, we predicted southern stingrays in marine reserves would reduce the use of deep forereef habitats in the favor of shallow flats where the risk of shark encounters is lower. Baited remote underwater video was used to survey stingrays and reef sharks in flats and forereef habitats of two reserves and two fished sites in Belize. The interaction between “protection status” and “habitat” was the most important factor determining stingray presence. As predicted, southern stingrays spent more time interacting with baited remote underwater videos in the safer flats habitats, were more likely to have predator-inflicted damage inside reserves, and were less abundant in marine reserves but only in the forereef habitat. These results are consistent with a predation-sensitive habitat shift rather than southern stingray populations being reduced by direct predation from reef sharks. Our study provides evidence that roving predators can induce pronounced habitat shifts in prey that rely on crypsis and refuging, rather than active escape, in high-visibility, heterogeneous marine habitats. Given documented impacts of stingrays on benthic communities it is possible restoration of reef shark populations with reserves could induce reef ecosystem changes through behavior-mediated trophic cascades.

Journal ArticleDOI
TL;DR: The research revealed strong effects of an ecosystem engineer on beetle assemblages while highlighting the potential role of apex predators in creating and modifying physical habitats for carrion-dependent species.
Abstract: Ecosystem engineers create physical changes in abiotic and biotic material, and through this process control the availability of resources for other species. Predators that abandon large portions of their prey may be ecosystem engineers that create habitat for carrion-dependent invertebrates that utilize carcasses during critical life-history periods. Between 04-May-2016 and 04-Oct-2016, we sampled beetle assemblages at 18 carcasses of prey killed by pumas and matching control sites in the southern Greater Yellowstone Ecosystem, USA, to measure the extent to which beetle families utilized these carcass “habitats”. We used generalized linear-mixed models and linear-mixed effect models to examine changes in beetle abundance, species richness, and Simpson’s Index of Diversity. We estimated kill rates and carrion production rates for individual pumas to better assess the impact of pumas on invertebrate communities. We collected 24,209 beetles representing 215 species. We identified eight beetle families that had significantly higher abundance at carcasses than control sites. Carcasses had a statistically large to very large effect (determined using Cohen’s d) on beetle abundance, richness, and diversity for the initial 8 weeks of sampling. Our research revealed strong effects of an ecosystem engineer on beetle assemblages while highlighting the potential role of apex predators in creating and modifying physical habitats for carrion-dependent species. This suggests that there may be consequences for invertebrate communities where apex predators exist at reduced numbers or have been eradicated. The ecological role of invertebrates is often overlooked, yet they are essential taxa that provide critical ecological services upon which we depend.

Journal ArticleDOI
TL;DR: It was found that fruit development time was positively correlated with fruit size and together the morphological fruit traits showed an important relationship with the growth form, but with a strong phylogenetic signal.
Abstract: In tropical dry forests, a high interspecific variation in the strategies of fruiting phenology has been documented. Therefore, phenological responses may be mediated by influence of environmental variables, functional plant attributes or phylogenetic inertia. During 2 years, we recorded the fruiting phenology of 151 species belonging to 5 different growth forms of a Neotropical dry forest in Mexico. We evaluated the relationships between fruiting phenology, abiotic factors (precipitation, temperature, day-length) and functional attributes (growth form, dispersal syndrome, size and time for fruit development) using phylogenetic least squares models (PGLS). More species had ripe fruits during the dry season (92%) than during rainy months and dispersed their seeds by autochory and endozoochory. We found that fruit development time was positively correlated with fruit size and together the morphological fruit traits (size and dispersal syndrome) showed an important relationship with the growth form, but with a strong phylogenetic signal. Environmental seasonality had a strong influence on fruit ripening time, without a relevant association to the phylogeny of plant species. However, the phenological response to the environment (rainfall and day-length) at the community level was mediated by growth form. In woody species, we documented a high interspecific fruiting variation linked with the different dispersal syndromes. In herbaceous species, fruiting phenology is a trait restricted by the duration of their life cycle by rainfall seasonality, which in turn might have selected some traits (e.g., dry fruit, presence of spines, explosive dehiscence) for maximizing seed dispersal during the dry season.

Journal ArticleDOI
TL;DR: A strong, positive, linear relationship between lichen biodiversity and habitat quality is recovered: lower levels of disturbance correlate to higher species diversity.
Abstract: The impacts of disturbance on biodiversity and distributions have been studied in many systems. Yet, comparatively less is known about how lichens–obligate symbiotic organisms–respond to disturbance. Successful establishment and development of lichens require a minimum of two compatible yet usually unrelated species to be present in an environment, suggesting disturbance might be particularly detrimental. To address this gap, we focused on lichens, which are obligate symbiotic organisms that function as hubs of trophic interactions. Our investigation was conducted in the southern Appalachian Mountains, USA. We conducted complete biodiversity inventories of lichens (all growth forms, reproductive modes, substrates) across 47, 1-ha plots to test classic models of responses to disturbance (e.g., linear, unimodal). Disturbance was quantified in each plot using a standardized suite of habitat quality variables. We additionally quantified woody plant diversity, forest density, rock density, as well as environmental factors (elevation, temperature, precipitation, net primary productivity, slope, aspect) and analyzed their impacts on lichen biodiversity. Our analyses recovered a strong, positive, linear relationship between lichen biodiversity and habitat quality: lower levels of disturbance correlate to higher species diversity. With few exceptions, additional variables failed to significantly explain variation in diversity among plots for the 509 total lichen species, but we caution that total variation in some of these variables was limited in our study area. Strong, detrimental impacts of disturbance on lichen biodiversity raises concerns about conservation and land management practices that fail to incorporate complete estimates of biodiversity, especially from ecologically important organisms such as lichens.

Journal ArticleDOI
TL;DR: It is suggested that increasing floral resources at high impervious sites may counteract the negative effects of impervious surface on bee diversity and pollination services in cities similar to Toledo, OH.
Abstract: Current declines in the abundance and diversity of bees and other pollinators has created uncertainty in their ability to reliably deliver pollination services. Recent studies examining urban bee diversity have provided conflicting results, with some studies identifying parts of cities with high bee diversity and others documenting reduced diversity with high levels of urbanization, with potential effects on surrounding agricultural areas. However, these studies have not specifically investigated pollination services, or examined the influence of local habitat conditions on these services. We surveyed urban gardens and city parks across the metropolitan region of Toledo, Ohio (USA) to understand how urbanization (impervious surface) and local habitat characteristics (herbaceous cover, floral abundance and color, tree abundance, canopy cover, soil moisture, garden size) impact bee communities (abundance, diversity, composition) and pollination services (visitation frequency). We collected 729 bees representing 19 genera and 57 species. We found that bee community composition was strongly associated with percent impervious surface. Bee abundance declined with increased canopy cover and impervious surface, while declines in bee diversity with increasing impervious surface were greatly reduced by increases in floral resources. Visitation rates were positively correlated with bee abundance and diversity, declining with increased impervious surface, but increasing with floral resource availability. These results suggest that increasing floral resources at high impervious sites may counteract the negative effects of impervious surface on bee diversity and pollination services in cities similar to Toledo, OH.

Journal ArticleDOI
TL;DR: Strong negative correlation between trophic level (CSIA of nitrogen) and the proportion of plant EAA ( CSIA of carbon) suggests that both novel methods can indicate the degree of microbivory in detritivores.
Abstract: Quantification of the bacterial, fungal, and plant energy channels to the nutrition of detritivores is methodologically challenging. This is especially true for earthworms that ingest large amounts of litter and soil mixed with microorganisms. Novel methods such as compound-specific stable isotope analysis (CSIA) of C and N of individual amino acids promise major progress in this field in comparison with bulk stable isotope analysis (bulk SIA). Here, we combine CSIA and bulk SIA of carbon and nitrogen to quantify the linkage of epigeic and endogeic earthworm species to different energy channels across boreal and temperate forest ecosystems. The results showed pronounced flux of energy directly from plants to earthworms (33–50% of essential amino acids, EAA) refining the position of earthworms in soil food webs as both competitors and consumers of microorganisms. Epigeic earthworm species primarily relied on plant litter and endogeic species primarily relied on bacteria and soil organic matter. The linkage of both groups to plant or microbial energy channel was likely driven by the quality of detritus. Both bulk 15N and 13C enrichments were related to the trophic level of earthworms. Furthermore, 15N enrichment was related to the proportions of bacterial and plant EAA in the diet. Strong negative correlation between trophic level (CSIA of nitrogen) and the proportion of plant EAA (CSIA of carbon) suggests that both novel methods can indicate the degree of microbivory in detritivores. CSIA of amino acids provide detailed and baseline-independent information on basal resources and trophic levels of detritivores.

Journal ArticleDOI
TL;DR: Sex- and age-based differences in habitat use reflected nursing mothers’ need for high-quality forage while also protecting their young from predation, and have implications for conservation and management of giraffes and other large-bodied, herd-forming ungulates in heterogeneous environments subject to anthropogenic threats.
Abstract: Fission–fusion dynamics hypothetically enable animals to exploit dispersed and ephemeral food resources while minimizing predation risk. Disentangling factors affecting group size and composition of fission–fusion species facilitates their management and conservation. We used a 6-year data set of 2888 group formations of Masai giraffes in Tanzania to investigate determinants of social group size and structure. We tested whether ecological (lion density, vegetation structure, and prevalence of primary forage plants), anthropogenic (proximity to human settlements), temporal (rainy or dry season), and social (local giraffe density, adult sex ratio, and proportion of calves) factors explained variation in group size and sex- and age-class composition. Food availability rather than predation risk mediated grouping dynamics of adult giraffes, while predation risk was the most important factor influencing congregations with calves. Smallest group sizes occurred during the food-limiting dry season. Where predation risk was greatest, groups with calves were in bushlands more than in open grasslands, but the groups were smaller in size, suggesting mothers adopted a strategy of hiding calves rather than a predator-detection-and-dilution strategy. Groups with calves also were farther from towns but closer to traditional human compounds (bomas). This may be due to lower predator densities, and thus reduced calf predation risk, near bomas but higher human disturbance near towns. Sex- and age-based differences in habitat use reflected nursing mothers’ need for high-quality forage while also protecting their young from predation. Our results have implications for conservation and management of giraffes and other large-bodied, herd-forming ungulates in heterogeneous environments subject to anthropogenic threats.

Journal ArticleDOI
TL;DR: Results provide clear experimental evidence that greater resource availability early in colony development increases overall productivity, and indicate that colonies may pursue different allocation strategies in response to the resource environment, investing in more or better workers.
Abstract: The temporal distribution of resources is an important aspect of habitat quality that can substantially impact population success. Although it is widely accepted that floral resources directly influence wild bee population sizes, we lack experimental data evaluating how resource availability affects colony growth via demographic mechanisms. To achieve this, we tracked marked individuals in bumble bee (Bombus vosnesenskii) colonies to evaluate whether worker survival and reproduction responded to experimentally elevated forage early in colony development. Specifically, we assessed the effect of early resource environment on worker and sexual offspring production, and the survival and body size of individual workers. We also assessed whether responses of colonies differed when exposed to higher or lower resource environments at a relatively smaller (~ 10 workers) or larger (~ 20 workers) size. Resource supplementation always resulted in greater total offspring and male production; however, the influence of supplementation on worker production and quality depended on colony size at the start of supplementation. Among colonies that were initially smaller, colonies that were supplemented produced fewer but larger bodied and longer lived workers compared to control counterparts. Among colonies that were initially larger, colonies that were supplemented produced more workers than corresponding controls, but without changes to worker quality. Collectively, these results provide clear experimental evidence that greater resource availability early in colony development increases overall productivity, and indicate that colonies may pursue different allocation strategies in response to the resource environment, investing in more or better workers.

Journal ArticleDOI
TL;DR: The effect of filter treatment differed among the three species and appeared to depend on litter quality, producing the greatest effect in recalcitrant litter (F. sylvatica).
Abstract: Sunlight can accelerate the decomposition process through an ensemble of direct and indirect processes known as photodegradation. Although photodegradation is widely studied in arid environments, there have been few studies in temperate regions. This experiment investigated how exposure to solar radiation, and specifically UV-B, UV-A, and blue light, affects leaf litter decomposition under a temperate forest canopy in France. For this purpose, we employed custom-made litterbags built using filters that attenuated different regions of the solar spectrum. Litter mass loss and carbon to nitrogen (C:N) ratio of three species: European ash (Fraxinus excelsior), European beech (Fagus sylvatica) and pedunculate oak (Quercus robur), differing in their leaf traits and decomposition rate, were analysed over a period of 7–10 months. Over the entire period, the effect of treatments attenuating blue light and solar UV radiation on leaf litter decomposition was similar to that of our dark treatment, where litter lost 20–30% less mass and had a lower C:N ratio than under the full-spectrum treatment. Moreover, decomposition was affected more by the filter treatment than mesh size, which controlled access by mesofauna. The effect of filter treatment differed among the three species and appeared to depend on litter quality (and especially C:N), producing the greatest effect in recalcitrant litter (F. sylvatica). Even under the reduced irradiance found in the understorey of a temperate forest, UV radiation and blue light remain important in accelerating surface litter decomposition.

Journal ArticleDOI
TL;DR: This study conducted a hand-pollination experiment with ten co-flowering species to determine whether the effects of heterospecific pollen receipt are mediated by the pollen donor or recipient species alone, or whether the results are determined by the interaction between them.
Abstract: Evaluation of pollen transfer in wild plant communities revealing heterospecific pollen receipt is common, yet experimental hand pollinations have revealed high among-species variation in the magnitude of its effect on recipient fitness. The causes of this among-species variation are unknown, however, prompting the investigation of underlying factors. Here, we conducted a hand-pollination experiment with ten co-flowering species to determine whether the effects of heterospecific pollen receipt are mediated by the pollen donor or recipient species alone, or whether the effects are determined by the interaction between them. We further assessed species traits potentially mediating interactive effects in heterospecific pollen receipt by evaluating the relationship between heterospecific pollen effect size and three different predictors reflecting a unique combination of pollen donor and recipient characteristics. Our results show, for the first time, that the magnitude of the heterospecific pollen receipt effect is determined by the specific combination of donor and recipient species (i.e., interactive effects). However, we were unable to uncover the specific combination of traits mediating these effects. Overall, our study provides strong evidence that an understanding of heterospecific pollen receipt effects based on recipient or donor characteristics alone may be insufficient. This study is an important step toward an understanding of consequences of heterospecific pollen receipt in co-flowering communities.

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TL;DR: The idea, across native and exotic species, that plant species turnover is the primary mechanism underlying effects of nutrient enrichment on allocation to growth and defense in plant communities is supported.
Abstract: High-resource environments typically favor quick-growing, poorly defended plants, while resource-poor environments typically favor slow-growing, well-defended plants. The prevailing hypothesis explaining this pattern states that, as resource availability increases, well-defended, slow-growing species are replaced by poorly defended, fast-growing species. A second hypothesis states that greater resource availability increases allocation to growth at the expense of defense, within species. Regardless of mechanism, if exotic species are released from enemies relative to natives, shifts in allocation to growth and defense both within and among species could differ by geographic provenance. To test whether resource availability alters growth or defense, within and among species, and whether any such effects differ between natives and exotics, we manipulated soil nutrient supply and access of aboveground insect herbivores and fungal pathogens under field conditions to individuals of six native and six exotic grass species that co-occurred in a North Carolina old field. The prevailing hypothesis’ prediction—that species-level enemy impact increases with species’ nutrient responsiveness—was confirmed. Moreover, this relationship did not differ between native and exotic species. The second hypothesis’ prediction—that individual-level enemy impact increases with nutrient supply, after accounting for species-level variation in performance—was not supported. Together, these results support the idea, across native and exotic species, that plant species turnover is the primary mechanism underlying effects of nutrient enrichment on allocation to growth and defense in plant communities.

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TL;DR: A meta-analysis of studies on pollen nutrients found that crude protein and the protein-to-lipid ratio in pollen strongly correlated with phylogeny, indicating that, besides producing pollen with sufficient nutrients for reproduction, the nutrient profile of zoophilous plants may have been shaped by their pollinators’ nutritional needs.
Abstract: A prime example of plant-animal interactions is the interaction between plants and pollinators, which typically receive nectar and/or pollen as reward for their pollination service. While nectar provides mostly carbohydrates, pollen represents the main source of protein and lipids for many pollinators. However, the main function of pollen is to carry nutrients for pollen tube growth and thus fertilization. It is unclear whether pollinator attraction exerts a sufficiently strong selective pressure to alter the nutritional profile of pollen, e.g., through increasing its crude protein content or protein-to-lipid ratio, which both strongly affect bee foraging. Pollen nutritional quality may also be merely determined by phylogenetic relatedness, with pollen of closely related plants showing similar nutritional profiles due to shared biosynthetic pathways or floral morphologies. Here, we present a meta-analysis of studies on pollen nutrients to test whether differences in pollen nutrient contents and ratios correlated with plant insect pollinator dependence and/or phylogenetic relatedness. We hypothesized that if pollen nutritional content was affected by pollinator attraction, it should be different (e.g., higher) in highly pollinator-dependent plants, independent of phylogenetic relatedness. We found that crude protein and the protein-to-lipid ratio in pollen strongly correlated with phylogeny. Moreover, pollen protein content was higher in plants depending mostly or exclusively on insect pollination. Pollen nutritional quality thus correlated with both phylogenetic relatedness and pollinator dependency, indicating that, besides producing pollen with sufficient nutrients for reproduction, the nutrient profile of zoophilous plants may have been shaped by their pollinators' nutritional needs.

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TL;DR: Environmental correlates of taxonomically-based and traits-based compositional distances using a comprehensive data set of diatom and macroinvertebrate communities found support for the role of environmental filtering as a driver of community dissimilarities of rather different biological groups.
Abstract: Recently, community ecology has emphasized the multi-facetted aspects of biological diversity by linking species traits and the environment. Here, we explored environmental correlates of taxonomically-based and traits-based compositional distances using a comprehensive data set of diatom and macroinvertebrate communities. We also explored the responses of different beta diversity components (i.e., overall beta diversity, turnover, and nestedness) of beta diversity facets (i.e., taxonomically and traits-based beta diversity) to environmental distances. Partial Mantel tests were used to test the relationships between beta diversity and environmental distance (while controlling for spatial distances). Taxonomically-based beta diversity varied much more than traits-based beta diversity, indicating strong functional convergence. We found that taxonomically-based beta diversity was largely driven by the turnover component. However, the nestedness component contributed more to overall traits-based beta diversity than the turnover component. Taxonomically-based beta diversity was significantly correlated with environmental distances for both diatoms and macroinvertebrates. Thus, we found support for the role of environmental filtering as a driver of community dissimilarities of rather different biological groups. However, the strength of these relationships between beta diversity and environmental distances varied depending on the biological group, facet, component, and the way which the environmental variables were selected to calculate the explanatory (distance) matrix. Our results indicated that both taxonomically and traits-based approaches are still needed to better understand patterns and mechanisms affecting the organization of biological communities in streams. This is because different facets of biological communities may be driven by different mechanisms.

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TL;DR: Call is called for the establishment of a unified framework that will guide diet reconstruction in stable isotope ecology to better guide treatment of lipids in isotope studies using carbon.
Abstract: Carbon isotopes are commonly used in trophic ecology to estimate consumer diet composition. This estimation is complicated by the fact that lipids exhibit a more depleted carbon signature (δ13C) than other macromolecules, and are often found at different concentrations among individual organisms. Some researchers argue that lipids bias diet reconstructions using stable isotopes and should be accounted for prior to analysis in food web mixing models, whereas others contend that removing lipids may result in erroneous interpretations of the trophic interactions under study. To highlight this disagreement on best practices for applying δ13C in food web studies, we sampled the recent literature to determine the frequency and method of lipid-correction. We then quantified the potential magnitude and source of bias in mixing model results from a theoretical example and case study of diet reconstruction. The literature was split nearly evenly as to whether lipid-correction was applied to δ13C data in mixing model estimates of diet composition. Comparative mixing model scenarios demonstrated that lipid-correction can substantially alter the estimated diet composition and interpretation of consumer foraging habits. Given the lack of consensus on whether or not to lipid-correct prey and/or consumers, and the associated variation in mixing model results, we call for the establishment of a unified framework that will guide diet reconstruction in stable isotope ecology. Uncertainty in the prevalence of direct routing versus de novo synthesis of lipids across ecosystems, taxa, and trophic levels must be resolved to better guide treatment of lipids in isotope studies using carbon.

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TL;DR: A consistent 2H-enrichment of compounds in heterotrophically growing plants across a series of autotrophic/heterotrophic plant pairs is shown and this work contributes to a better mechanistic understanding of what the biochemical principles are that couple the carbohydrate dynamics of plants to their δ2H values and hope to foster the application of H isotopes in plant sciences.
Abstract: Hydrogen (H) isotopes of plant organic compounds are rarely employed in ecological studies. If so, these values are interpreted as being indicative of the plant source and/or leaf water. Recent observations suggest, however, that variations in hydrogen isotope fractionation that occur during the biosynthesis of plant compounds (2H-ebio) imprint valuable metabolic information into the hydrogen isotope composition (δ2H values) of plant organic compounds. Here we show a consistent 2H-enrichment of compounds in heterotrophically growing plants across a series of autotrophic/heterotrophic plant pairs. We suggest that this is due to a higher recycling of compounds in the Calvin and tricarboxylic acid cycles in heterotrophic plants that is associated with a more complete exchange of C-bound H with the surrounding 2H-enriched foliar water. Interestingly, we found that 2H-enrichment in heterotrophic plants was larger for carbohydrates than for lipids, with an average 2H-enrichment of 76 ± 9‰ in α-cellulose and 23 ± 23‰ in n-alkanes. We propose that this systematically larger 2H-enrichment for carbohydrates than for lipids is either due to different level of 2H-fractionation associated with heterotrophically produced NADPH, or to the potential uptake of lipids by heterotrophic plants. With the work we present here, we contribute to a better mechanistic understanding of what the biochemical principles are that couple the carbohydrate dynamics of plants to their δ2H values and hope to foster as such the application of H isotopes in plant sciences.

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TL;DR: The results suggest that the positive productivity–diversity relationship found may be due, at least in part, to increased food resources and niche opportunities at more productive elevations, which may also be partially due to reductions in competition that result from productivity increases.
Abstract: Photosynthetic productivity is a key determinant of the abundance and distribution of biodiversity around the world. The effect of this productivity on the distribution patterns of mammals is frequently invoked; however, it is seldom measured directly. In this study, we used Sherman live traps set in dry and rainy seasons across a 2300-m elevation gradient in southwestern Mexico to assess small rodent species distributions, and to relate these patterns to habitat structure, climate, and a well-accepted measure of photosynthetic productivity: the normalized difference vegetation index (NDVI). While habitat structure and climate helped explain some of the patterns observed, NDVI proved to be the most important contributing variable for most of the distribution models. We also found that partitioning the gradient-distribution model by trapping season revealed strong differences in terms of the effect of NDVI and the other explanatory variables. For example, lower elevations were associated with seasonal and year-round reductions in rodent diversity and were composed almost exclusively of granivore-based species assemblages. By contrast, the middle and upper elevations were more species rich, less affected by seasonality, and characterized by omnivorous species. Our results suggest that the positive productivity–diversity relationship found may be due, at least in part, to increased food resources and niche opportunities at more productive elevations. Increased diversity at the higher elevations may also be partially due to reductions in competition that result from productivity increases, as well as from the broader spectrum of feeding guild representation that it and the lack of seasonality allow.

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TL;DR: Snow burrowing is an important strategy that appears to allow grouse to mediate the possibly stressful effects of cold temperatures, and this is one of the first studies to explore how variable winter weather conditions influence stress in a free–living cold–adapted vertebrate and its ability to Mediate this relationship behaviorally.
Abstract: Animals in temperate northern regions employ a variety of strategies to cope with the energetic demands of winter. Behavioral plasticity may be important, as winter weather conditions are increasingly variable as a result of modern climate change. If behavioral strategies for thermoregulation are no longer effective in a changing environment, animals may experience physiological stress, which can have fitness consequences. We monitored winter roosting behavior of radio–tagged ruffed grouse (Bonasa umbellus), recorded snow depth and temperature, and assayed droppings for fecal corticosterone metabolites (FCM). Grouse FCM levels increased with declining temperatures. FCM levels were high when snow was shallow, but decreased rapidly as snow depth increased beyond 20 cm. When grouse used snow burrows, there was no effect of temperature on FCM levels. Snow burrowing is an important strategy that appears to allow grouse to mediate the possibly stressful effects of cold temperatures. This is one of the first studies to explore how variable winter weather conditions influence stress in a free–living cold–adapted vertebrate and its ability to mediate this relationship behaviorally. Animals that depend on the snowpack as a winter refuge will likely experience increased stress and possible fitness costs resulting from the loss of snow cover due to climate change.

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TL;DR: The low levels of intraspecific morphological variation that are found suggest that the approach of using mean species traits is valid, in this fauna at least, and it is encouraged to measure greater numbers of individuals, especially across gradients, to shed further light on intrapecific variation in this functionally important group of insects.
Abstract: The trait-based approach to ecology promises to provide a mechanistic understanding of species distributions and ecosystem functioning. Typically, trait analyses focus on average species trait values and assume that intraspecific variation is small or negligible. Recent work has shown, however, that intraspecific trait variation can often contribute substantially to total trait variation. Whilst many studies have investigated intraspecific variation in plants, very few have done so for invertebrates. There is no research on the level of intraspecific trait variation in ants (Hymenoptera: Formicidae), despite the fact that there is a growing body of literature using ant morphological trait data and demonstrating that these insects play important roles in many ecosystems and food webs. Here, we investigate the intraspecific variability of four commonly used ant morphological traits from 23 species from the Maloti-Drakensberg Mountains of southern Africa. In total, we measured 1145 different individuals and made 6870 trait measurements. Intraspecific variation accounted for only 1–4% of total trait variation for each of the four traits we analysed. We found no links between intraspecific variation, phylogeny and elevation. On average, six individuals generated robust species means but under biased sampling scenarios 20 individuals were needed. The low levels of intraspecific morphological variation that we find suggest that the approach of using mean species traits is valid, in this fauna at least. Regardless, we encourage ant trait ecologists to measure greater numbers of individuals, especially across gradients, to shed further light on intraspecific variation in this functionally important group of insects.