Foragers typically attempt to consume food resources that offer the greatest energy gain for the least cost, switching between habitats as the most profitable food resource changes over time. Optimal foraging models require accurate data on the gains and costs associated with each food resource to successfully predict temporal shifts. Whilst previous studies have shown that seasonal changes in food quantity and quality can drive habitat shifts, few studies have shown the effects on habitat choice of seasonal changes in metabolic foraging costs. In this study we combined field and literature data to construct an optimal foraging model to examine the effect of seasonal changes in food quantity, food quality and foraging costs on the timing of a switch from terrestrial to aquatic habitat by non-breeding mute swans Cygnus olor in a shallow river catchment. Feeding experiments were used to quantify the functional response of swans to changes in aquatic plant biomasses. By sequentially testing alternative models with fixed or variable values for food quantity, food quality and foraging cost, we found that we needed to include seasonal variance in foraging costs in the model to accurately predict the observed habitat switch date. However, we did not need to include seasonal variance in food quantity and food quality, as accurate predictions could be obtained with fixed values for these two parameters. Therefore, the seasonal changes in foraging costs were the key factor influencing the behavioural decision to switch feeding habitats. These seasonal changes in foraging costs were driven by changes in water velocity; the profitability of aquatic foraging was negatively related to water velocity, as faster water required more energy to be expended in swimming. Our results demonstrate the importance of incorporating seasonal variation in foraging costs into our understanding of the foraging decisions of animals.

The determinants of local species richness in plant communities have been the subject of much debate. Is species richness the result of stochastic events such as dispersal processes, or do local environmental filters sort species into communities according to their ecological niches? Recent studies suggest that these two processes simultaneously limit species richness, although their relative importance may vary in space and time. Understanding the limiting factors for species richness is especially important in light of the ongoing global warming, as new species establish in resident plant communities as a result of climate-driven migration. We examined the relative importance of dispersal and environmental filtering during seedling recruitment and plant establishment in an alpine plant community subjected to seed addition and long-term experimental warming. Seed addition increased species richness during the seedling recruitment stage, but this initial increase was cancelled out by a corresponding decrease in species richness during plant establishment, suggesting that environmental filters limit local species richness in the long term. While initial recruitment success of the sown species was related to both abiotic and biotic factors, long-term establishment was controlled mainly by biotic factors, indicating an increase in the relative importance of biotic interactions once plants have germinated in a microhabitat with favourable abiotic conditions. The relative importance of biotic interactions also seemed to increase with experimental warming, suggesting that increased competition within the resident vegetation may decrease community invasibility as the climate warms.

Over the last decades, many species have been forced to track their shifting climate envelopes, and at the same time man-induced landscape fragmentation has led to the global decrease of natural habitat availability and connectivity. The interaction between these two co-occurring global environmental changes might have very strong effects on biodiversity that are still understudied. Species-specific responses to these environmental changes critically depend on individual dispersal, either to track suitable climatic conditions or to cope with landscape fragmentation. Here we study how dispersal in an ectotherm is affected by interactions between landscape fragmentation and weather conditions. We show that both the emigration rates out of suitable habitats and the topology of the trajectory of dispersing individuals were affected by temperature and landscape fragmentation. The emigration rate was temperature-dependent in fragmented landscapes, with butterflies emigrating more at high temperatures. The emigration rate was temperature insensitive in more continuous landscapes. Move length was farther at low temperatures and less at high temperatures in fragmented landscapes. Move length was less at low temperatures and farther at high temperatures in more continuous landscapes. To our knowledge only two recent studies have documented patterns of interactions between climate and fragmentation, despite the fact that they are the two main drivers of biodiversity loss worldwide. Here, we go a step further by providing mechanistic explanations to such patterns.

Understanding the factors that regulate biodiversity over spatial and temporal gradients is an important scientific objective with ramifications for theory and conservation. Species composition is known to vary across spatial gradients, but how this spatial variation is linked to temporal dynamics is less well studied. Our objective was to understand how Shannon (α) diversity, spatial species turnover (Bray–Curtis dissimilarity), and temporal species turnover (Bray–Curtis dissimilarity) varied with regard to three topographic gradients (aspect, slope and elevation) over one growing season. In April, June and August of 2011, the herbaceous layer was sampled in 320 1-m² plots within Big Everidge Hollow, an old-growth forest in southeastern Kentucky. Multiple regression models revealed that Shannon diversity was linearly related to aspect (negative) and slope (positive), but unimodally related to elevation, indicating steep, mid-elevation, and south-facing plots were most diverse. Distance decay analysis showed that significant spatial species turnover occurred across all three topographic gradients, but aspect and elevation had a greater influence on compositional dissimilarity than slope. Mean temporal species turnover was significantly greater (p < 0.001) between April and June (0.39 ± 0.02 SE) than between June and August (0.20 ± 0.01). April-to-June turnover was related to aspect (linear) and elevation (quadratic; r²= 0.23, p < 0.0001), suggesting greater temporal species turnover occurred on north-facing and mid-elevation plots during this period; however, June-to-August turnover was weakly related to slope only (positive linear; r²= 0.08, p = 0.006). Environmental heterogeneity generated by topography is one of many factors that may constrain or promote biodiversity through space and across time, and a solid understanding of these spatiotemporal patterns of diversity can benefit both conservation and ecological theory.

The potential connection between exploitation and interference competition was recognized long ago but has not been evaluated. We measured the levels of both forms of competition for the protist Didinium preying upon Paramecium. Across populations, exploitation intensity was tightly linked to interference intensity, and the form of this relationship follows from a simple model of interaction speeds. The variation in interference competition was as large across populations of Didinium as has been observed previously across species from a variety of taxa including birds, mammals, insects, crustaceans, flatworms and protists. The link between exploitation and interference competition alters our understanding of how interference competition influences population dynamics. Instead of simply stabilizing systems, variation in interference levels can shift population dynamics through qualitatively different regimes because of its association with exploitation competition. Strong interference competition pushes a system to a regime of deterministic extinction, but intermediate interference generates a system that is stable with a high competitive ability. This may help to explain why the distribution of interference values is unimodal and mostly intermediate in intensity.

Some conceptual models seeking to explain the coexistence of multiple species in hyperdiverse settings predict that species will not be randomly distributed with respect to each other. In stark contrast the ‘stochastic geometry’ model assumes that a species fine-scale spatial distribution is independent of that of other species in the community. Empirical tests in temperate and tropical forests have provided support for both perspectives. Using point pattern analyses we assessed the prevalence of heterospecific associations between > 10 500 pairs of species and > 3400 pairs of plant functional types (PFTs) in four biodiverse shrubland communities in southwestern Australia. After controlling for first-order effects, spatial associations between species and PFTs were rare, but were most prevalent at the least species-rich of the four sites considered. Individuals tended to have fewer species in their local neighbourhoods than expected under a null model of random relabelling, with this departure most pronounced at the site with fewest species. The consistency of neighbourhood composition experienced by individuals of the same species is, as a result, less than the average under random mixing. Our results demonstrate that the frequency of heterospecific spatial associations is both rare in speciose systems and declines with species richness, and provide further empirical support for the stochastic geometry assumption in species-rich communities.

Personality and metabolic rate are predicted to show covariance on methodological and functional grounds, but empirical studies at the individual level are rare, especially in natural populations. Here we assess the relationship between exploration behaviour, an important axis of personality, and basal metabolic rate (BMR) for 680 free-living great tits Parus major, studied over three years. We find that exploration behaviour is weakly negatively related to BMR among female, but not male, birds. Moreover, we find exploration behaviour to be independent of methodological aspects of BMR measurements (e.g. activity levels, time to acclimatize) which have been suggested to be indicative of personality-related activity or stress levels during measurement. This suggests that the weak link between exploration behaviour and BMR found here is functional rather than methodological. We therefore test the hypothesis that selection favours covariance between exploration behaviour and metabolic rate, but find no evidence for correlational survival or fecundity selection. Our data therefore provide at best only very weak evidence for a functional link between personality and metabolic rate, and we suggest that studies of personality and metabolic strategies, or personality and daily energy expenditure, are required to further resolve the link between personality and metabolic rate.

The total effect of predators on prey is a combination of direct consumption, and non-consumptive effects (NCEs), such as predator-induced changes to prey morphology, behaviour and life history. Past research into NCEs has tended to focus on pair-wise interactions between predators and prey, while in natural ecosystems, species exist in complex communities with several trophic levels made up of multiple autotrophic and heterotropic species. To address how predator NCEs alter the photosynthetic and heterotrophic components of communities, we exposed microbial microcosms to one of three predator treatments: live predators (full predator effect), freeze-killed predators (NCEs only) or no predators (control), and incubated them under either 12 h:12 h light:dark conditions or continual darkness. Under 12 h:12 h light:dark conditions, NCEs-only communities never differed from predator-free communities, but differed from live predator communities. Under conditions of continual darkness, the structure of NCEs-only communities differed from predator-free controls, but not from live predator communities, suggesting NCEs can be strong enough to structure communities. Predation threat may cause certain prey to induce defences, such as reductions in movement, which make them less competitive in a community setting. This reduction in competitive ability could lead to these species being driven to extinction through interspecific competition, resulting in similar communities to those in which live predators are present. Heterotrophic species whose rates of resource acquisition depend on movement rates may be affected to a greater extent than autotrophs by predator-induced reductions in movement, accounting for our observed differences in predator NCEs in ‘dark’ and ‘light’ communities. Our results suggest that the community-level consequences of fear are greater in the dark.

Phenotypes of plants, and thus their ecology and evolution, can be affected by the environmental conditions experienced by their parents, a phenomenon called parental effects or transgenerational plasticity. However, whether such effects are just passive responses or represent a special type of adaptive plasticity remains controversial because of a lack of solid tests of their adaptive significance. Here, we investigated transgenerational effects of different nutrient environments on the productivity, carbon storage and flowering phenology of the perennial plant Plantago lanceolata, and whether these effects are influenced by seasonal variation in the maternal environment. We found that maternal environments significantly affected the offspring phenotype, and that plants consistently produced more biomass and had greater root carbohydrate storage if grown under the same environmental conditions as experienced by their mothers. The observed transgenerational effects were independent of the season in which seeds had matured. We therefore conclude that transgenerational effects on biomass and carbon storage in P. lanceolata are adaptive regardless of the season of seed maturation.

As human-aided range expansions and climate change alter the distributions of plants and their herbivores, predicting and addressing novel species interactions will become increasingly pressing for community ecologists. In this context, a key, surprisingly understudied question is: when an exotic plant is introduced, which herbivores will adopt this new potential host? Whether the plant is a weed, an ornamental, or a crop, the development versus non-development of a novel plant–insect interaction can have profound effects for both economic and conservation applications. In this paper, we sketch mechanistic and statistical frameworks for predicting these interactions, based on how plant and herbivore traits as well as shared evolutionary history can influence detection, recognition, and digestion of novel plants. By emphasizing mechanisms at each of these steps, we hope to clarify different aspects of novel interactions and why they may or may not occur. We also emphasize prediction and forecasting, as a major goal is to know in advance which interactions will develop from the many plant or insect introductions that occur in natural and man-made systems.

Temperature can regulate a number of important biological processes and species interactions. For example, environmental temperature can alter insect herbivore consumption, growth and survivorship, suggesting that temperature-driven impacts on herbivory could influence plant community composition or nutrient cycling. However, few studies to date have examined whether rising temperature influences herbivore preference and performance among multiple plant species, which often dictates their impact at the community level. Here, we assessed the effects of temperature on the performance and preference of the generalist herbivore Popillia japonica among nine plant species. We show that, on average, consumption rates and herbivore performance increased at higher temperatures. However, there was considerable variation among plant species with consumption and performance increasing on some plant species at higher temperatures but decreasing on others. Plant nutritional quality appeared to influence these patterns as beetles increased feeding on high-nitrogen plants with increasing temperature, suggesting stronger nitrogen limitation. In addition to changes in feeding rates, feeding preferences of P. japonica shifted among temperatures, a pattern that was largely explained by differential deterrence of plant chemical extracts at different temperatures. In fact, temperature-induced changes in the efficacy of plant chemical extracts led P. japonica to reduce its diet breadth at higher temperatures. Our results indicate that rising temperatures will influence herbivore feeding behavior by altering the importance of plant nutritional and chemical traits, suggesting that climate change will alter the strength and sign of plant–insect interactions.

Understanding the various processes contributing to community assembly is among the central aims of ecology. As a means of exploring this topic we quantified the relative influences of habitat filtering and competition in establishing patterns of community functional trait diversity across a landscape of lakes. Habitat filtering has been invoked in shaping community structure when co-occurring taxa are more similar in their traits than expected by chance (under-dispersion), and competition has been inferred as a structuring agent when co-occurring taxa are less similar (over-dispersion). We tested these hypotheses in crustacean zooplankton communities using a functional trait-based approach based on five traits defining zooplankton feeding and habitat preferences across 51 lakes spanning several large limnological gradients. In general, zooplankton communities were functionally less diverse than random assemblages created from the same regional species pool. Furthermore, functional diversity was strongly correlated with variables related to lake productivity, suggesting that access to resources was the chief habitat filtering process constraining zooplankton functional diversity. This pattern was driven by the predominantly herbivorous cladocerans as opposed to the more commonly omnivorous, and sometimes carnivorous, copepods.

Canopy structural parameters are often used to give adequate representation of vegetated ecosystems for various purposes including primary productivity, climate system, water and carbon gas exchanges, and radiation extinction. Canopy structural parameters are usually described using several pseudo-synonymous terms, often measuring different components of vegetation canopies. Standardization in the definitions has fallen short, leading to confusion of terms even in standard text books making the comparison of historic measures futile. Here we clarify concepts that have been used for fractional canopy element cover and openness measures. The fractional canopy element cover and openness concepts considered are canopy closure, canopy cover, canopy openness, crown closure, crown completeness, crown cover, crown porosity, site openness and tilt openness. New methodologies are presented to obtain large scale fractional canopy element cover and openness measures using hemispherical photography. The new methodologies and variations in definitions of fractional canopy element cover and openness concepts are demonstrated using photographic measurements in complex topography. The results indicate that both fractional canopy element cover and openness parameters can be estimated with a few point-based measurements using hemispherical photography. Hemispherical photography is therefore less time, labour and resource intensive, as compared to point based measuring techniques of canopy element cover and openness.

Forcibly removing species from ecosystems has important consequences for the remaining assemblage, leading to changes in community structure, ecosystem functioning and secondary (cascading) extinctions. One key question that has arisen from single- and multi-trophic ecosystem models is whether the secondary extinctions that occur within competitive communities (guilds) are also important in multi-trophic ecosystems? The loss of consumer–resource links obviously causes secondary extinction of specialist consumers (topological extinctions), but the importance of secondary extinctions in multi-trophic food webs driven by direct competitive exclusion remains unknown. Here I disentangle the effects of extinctions driven by basal competitive exclusion from those caused by trophic interactions in a multi-trophic ecosystem (basal producers, intermediate and top consumers). I compared food webs where basal species either show diffuse (all species compete with each other identically: no within guild extinctions following primary extinction) or asymmetric competition (unequal interspecific competition: within guild extinctions are possible). Basal competitive exclusion drives extra extinction cascades across all trophic levels, with the effect amplified in larger ecosystems, though varying connectance has little impact on results. Secondary extinction patterns based on the relative abundance of the species lost in the primary extinction differ qualitatively between diffuse and asymmetric competition. Removing asymmetric basal species with low (high) abundance triggers fewer (more) secondary extinctions throughout the whole food web than removing diffuse basal species. Rare asymmetric competitors experience less pressure from consumers compared to rare diffuse competitors. Simulations revealed that diffuse basal species are never involved in extinction cascades, regardless of the trophic level of a primary extinction, while asymmetric competitors were. This work highlights important qualitative differences in extinction patterns that arise when different assumptions are made about the form of direct competition in multi-trophic food webs.

The extent to which ecological communities are coherent entities as opposed to mere intersections of individual species distributions has long been one of the fundamental questions of ecology. Gradient analysis is one commonly used tool for addressing this question; however, all such studies have used organisms from a single taxon or guild. This risks missing important connections due to non-competitive interactions, which should be more likely to occur between members of different guilds. Such organisms are unlikely to compete for resources and can have complementary niches that promote non-competitive interactions. We examined the abundances of taxa in four interacting guilds along an elevation gradient in a forest in the southern Appalachian mountains. A causal discovery algorithm was used to investigate the relative frequencies of interguild and intraguild interactions. These were approximately equally common once taxonomic richness was taken into account. We used elements of metacommunity structure analysis to study the extent to which species distributions are non-independent and tested the hypothesis that combinations of two or more interacting guilds exhibit more coherence than single guilds. In this analysis, all guilds other than collembola were classified as Clementsian or quasi-Clementsian. (Collembola were classified as random.) When sets of multiple guilds were examined, Clementsian and quasi-Clementsian structures predominated. We also compared boundary conjunction, measured as Morisita’s index (MI) for these sets of guilds to the weighted average of the guilds’ MI values. Only sets of directly interacting guilds had higher-than-baseline boundary conjunction values, and such boundary conjunction values are found in all but one set of directly interacting guilds. Our results highlight the importance of inter-guild interactions in structuring patterns of cooccurrence. Trophic interactions and plant–fungus symbioses (mutualistic and/or pathogen–host) appear particularly important.

The structure of mutualistic networks provides insights into ecological and coevolutionary dynamics of interacting species. However, the spatial effect has only recently been incorporated as a factor structuring mutualistic networks. In this study, we evaluated how the topological structure and species turnover of ant–plant mutualistic networks vary over a spatial gradient. We showed that although the ant and plant composition of networks changed over space, the central core of generalist species and the structure of networks remained unaltered on a geographic distance of up to 5099 m in the southern Brazilian Amazon. This finding indicates that independently of variation in local and landscape environmental factors, the nonrandom pattern organization of these interacting assemblages do not change. Finally, we suggest that a stable core can increase the potential for coevolutionary convergence of traits among species from both sides of the interaction within the community. These findings contribute to our understanding of the maintenance of biodiversity and coevolutionary processes.

The fluctuations of natural populations have deep impact on important ecological issues, such as pest outbreaks, fisheries or the formation of harmful algal blooms (HABs). However, consensus on the appropriate descriptor of such fluctuations is still lacking. Here, using 16 to 20 years of weekly data on marine microbial population abundance comprising more than 200 species, we analysed the distribution of the population fluctuations. We found that population fluctuations of all groups and in 12 out of 17 species were not Gaussian (D’Agostino test p < 0.05) but instead were adequately described by Levy-stable distributions (LSD). Consistent with ecological theories, the LSD characteristic parameter (α) characterizes as highly volatile those groups known to form HABs, such as dinoflagellates (α= 1.48), and as lowly volatile, nanoflagellates (α= 1.92), a group which can be subjected to predatory control. Moreover, zooplankton groups composed of species with sexual reproduction and complex life cycles such as crustacean and Appendicullaria also showed departures from Gaussian population fluctuations and adequate fits to LSD. Our results suggest that heavy-tailed population fluctuations are widespread, implying that extreme population fluctuations are more likely than previously expected, a fact that has important consequences for the predictability of population outbreaks.

Soil-dwelling insects commonly co-occur and feed simultaneously on belowground plant parts, yet patterns of damage and consequences for plant and insect performance remain poorly characterized. We tested how two species of root-feeding insects affect the performance of a perennial plant and the mass and survival of both conspecific and heterospecific insects. Because root damage is expected to impair roots’ ability to take up nutrients, we also evaluated how soil fertility alters belowground plant–insect and insect–insect interactions. Specifically, we grew common milkweed Asclepias syriaca in low or high nutrient soil and added seven densities of milkweed beetles Tetraopes tetraophthalmus, wireworms (mainly Hypnoides abbreviatus), or both species. The location and severity of root damage was species-specific: Tetraopes caused 59% more damage to main roots than wireworms, and wireworms caused almost seven times more damage to fine roots than Tetraopes. Tetraopes damage decreased shoot, main root and fine root biomass, however substantial damage by wireworms did not decrease any component of plant biomass. With the addition of soil nutrients, main root biomass increased three times more, and fine root biomass increased five times more when wireworms were present than when Tetraopes were present. We detected an interactive effect of insect identity and nutrient availability on insect mass. Under high nutrients, wireworm mass decreased 19% overall and was unaffected by the presence of Tetraopes. In contrast, Tetraopes mass increased 114% overall and was significantly higher when wireworms were also present. Survival of wireworms decreased in the presence of Tetraopes, and both species’ survival was negatively correlated with conspecific density. We conclude that insect identity, density and soil nutrients are important in mediating the patterns and consequences of root damage, and suggest that these factors may account for some of the contradictory plant responses to belowground herbivory reported in the literature.

Ultimate causes of phenotypic plasticity in visual appearance are frequently related to increasing the degree of crypsis in a way specific to the environment. The cues used to elicit such plastic responses may be both direct (i.e. straightforward background matching) as well as indirect. In the latter case, cues other than the visual signals from of the environment are used to predict the phenotype best corresponding to the particular situation. On the basis of a series of laboratory experiments we show that the remarkable variability in the visual appearance of the larvae of the geometrid moth Ematurga atomaria, though genetically based in part, involves a substantial environmental component. Using multiple correspondence analysis, we transformed the multidimensional variation in colour and pattern into two dimensions interpretable as patterning and darkness. Plastic changes in the darkness of the larvae were elicited by direct cues: the larvae were darker when reared on dark host-plants. Host-specific degree of patterning was also induced in absolute darkness which indicates the use of an indirect cue. This was unexpected because the study species is broadly polyphagous, and thus not likely to have evolved adaptations specific to particular host-plant species. Indeed, the larvae of E. atomaria originating from geographic populations using different host-plants showed analogous plastic responses which indicates that the link between the indirect cue and visual appearance of the host needs not to be specific to plant species. In an additional experiment, we showed that surface roughness is a likely candidate to serve as the proximate cue for determination of some pattern elements, a case not reported for insect larvae earlier.

Pollen and seed dispersal are the two key processes in which plant genes move in space, mostly mediated by animal dispersal vectors in tropical forests. Due to the movement patterns of pollinators and seed dispersers and subsequent complex spatial patterns in the mortality of offspring, we have little knowledge of how pollinators and seed dispersers affect effective gene dispersal distances across successive recruitment stages. Using six highly polymorphic microsatellite loci and parentage analyses, we quantified pollen dispersal, seed dispersal, and effective paternal and maternal gene dispersal distances from pollen- and seed-donors to offspring across four recruitment stages within a population of the monoecious tropical tree Prunus africana in western Kenya. In general, pollen-dispersal and paternal gene dispersal distances were much longer than seed-dispersal and maternal gene dispersal distances, with the long-distance within-population gene dispersal in P. africana being mostly mediated by pollinators. Seed dispersal, paternal and maternal gene dispersal distances increased significantly across recruitment stages, suggesting strong density- and distance-dependent mortality near the parent trees. Pollen dispersal distances also varied significantly, but inconsistently across recruitment stages. The mean dispersal distance was initially much (23-fold) farther for pollen than for seeds, yet the pollen-to-seed dispersal distance ratio diminished by an order of magnitude at later stages as maternal gene dispersal distances disproportionately increased. Our study elucidates the relative changes in the contribution of the two processes, pollen and seed dispersal, to effective gene dispersal across recruitment. Overall, complex sequential processes during recruitment contribute to the genetic make-up of tree populations. This highlights the importance of a multistage perspective for a comprehensive understanding of the impact of animal-mediated pollen and seed dispersal on small-scale spatial genetic patterns of long-lived tree species.

The size distribution of trees in natural forests is a fundamental attribute of forest structure. Previous attempts to model tree size distributions using simple functions (such as power or Weibull functions) have had limited success, typically overestimating the number of large stems observed. We describe a model which assumes that the dominant mortality process is asymmetric competition when trees are smaller, and size-independent processes (e.g. disturbance) when trees are larger. This combination of processes leads to a size distribution which takes the form of a power distribution in the small tree phase and a Weibull distribution in the large tree phase. Analyses of data from four large-scale (≥ 24 ha each) subtropical and temperate forest plots totalling 99 ha and approximately 0.4 million trees provide support for this model in two respects: (a) the combined function provided unbiased predictions and (b) power-law functions fitted to small trees had exponents that deviated from the universal exponent of –2 predicted by metabolic scaling theory, gradually decreasing from subtropical evergreen to temperate deciduous forests along the latitudinal gradient.

Bark beetle population dynamics is thought to be primarily driven by bottom-up forces affecting insect performance and host tree resistance. Although there are theoretical predictions and empirical evidences that predation and parasitism may play an important role in driving bark beetle population fluctuations, long-term studies testing the role of both biotic and abiotic controls on population dynamics are still rare. The aim of the study was to quantify the relative importance of predation, negative density feedback and abiotic factors in driving Ips typographus population dynamics. We analyzed a unique time series of population density of I. typographus and its main predator Thanasimus formicarius over almost two decades in four regions across Sweden. We used a discrete population model and a multi-model inference approach to evaluate the importance of both bottom up and top down factors. We found that availability of breeding substrates in the form of storm-felled trees was the main outbreak trigger, while strong intra-specific competition for host trees was the main endogenous regulating factor. Although temperature-related metrics are known to have strong individual effect on I. typographus development and number of generations, they did not emerge as important drivers of population dynamics. A positive effect of low summer rainfall was evident only in the region located in the southernmost and warmest part of the spruce distribution range in Sweden. Predator density did not emerge as an important prey regulating factor. As the reported damage from storms seems to have increased across whole Europe, spruce forests are expected to be increasingly susceptible to large outbreaks of I. typographus with important economic and ecological consequences for boreal ecosystems. However, the observed negative density feedback seems to be a natural regulating mechanism that impedes a strong long-term propagation of the outbreaks.

Studies of trait-mediated indirect interactions (TMIIs) typically focus on effects higher predators have on per capita consumption by intermediate consumers of a third, basal prey resource. TMIIs are usually evidenced by changes in feeding rates of intermediate consumers and/or differences in densities of this third species. However, understanding and predicting effects of TMIIs on population stability of such basal species requires examination of the type and magnitude of the functional responses exhibited towards them. Here, in a marine intertidal system consisting of a higher-order fish predator, the shanny Lipophrys pholis, an intermediate predator, the amphipod Echinogammarus marinus, and a basal prey resource, the isopod Jaera nordmanni, we detected TMIIs, demonstrating the importance of habitat complexity in such interactions, by deriving functional responses and exploring consequences for prey population stability. Echinogammarus marinus reacted to fish predator diet cues by reducing activity, a typical anti-predator response, but did not alter habitat use. Basal prey, Jaera nordmanni, did not respond to fish diet cues with respect to activity, distribution or aggregation behaviour. Echinogammarus marinus exhibited type II functional responses towards J. nordmanni in simple habitat, but type III functional responses in complex habitat. However, while predator cue decreased the magnitude of the type II functional response in simple habitat, it increased the magnitude of the type III functional response in complex habitat. These findings indicate that, in simple habitats, TMIIs may drive down consumption rates within type II responses, however, this interaction may remain de-stabilising for prey populations. Conversely, in complex habitats, TMIIs may strengthen regulatory influences of intermediate consumers on prey populations, whilst potentially maintaining prey population stability. We thus highlight that TMIIs can have unexpected and complex ramifications throughout communities, but can be unravelled by considering effects on intermediate predator functional response types and magnitudes.

Photoautotroph nitrogen (N) and phosphorus (P) tissue concentrations can influence ecosystem function via processes including growth, decomposition, and consumption, and may reflect traits maintaining coexistence. Studies in terrestrial systems have led to hypotheses that latitudinal trends in the N and P content of leaves may be driven by soil substrate age, environmental temperature, or season length; however, terrestrial patterns alone cannot differentiate these mechanisms. Here, we demonstrate that broad geographical patterns of N and P in freshwater and marine multicellular photoautotrophs are concordant with those in terrestrial ecosystems. Our > 6800 record database reveals that mean tissue N and P increase with latitude in all ecosystems, but P increases more rapidly, causing N:P to decline; mean N:P scaling within individuals also is identical among systems, despite very different evolutionary environments. A partitioning of the variance in these data suggests that species composition and local environmental context likely lead to the variation observed within a latitudinal band. However, the consistency of trends in photosynthetic tissue chemistry across Earth’s ecosystems suggests that biogeographical gradients in insolation and growing season length may constrain tissue N and P, whereas global trends in temperature, nutrient supply, and soil substrate age are unlikely to generate the consistent latitudinal trends among ecosystems. Thus, this cross-ecosystem comparison suggests a new hypothesis, global patterns of insolation, while also providing a new perspective on other mechanisms that have been hypothesized to underlie latitudinal trends in photosynthetic tissue chemistry.

The restriction of invasion biology to non-native species has been laid down as one founding principle of the discipline by many researchers. However, this split between native and non-native species is highly controversial. Using a phenomenological approach and a more pragmatic examination of biological invasions, the present paper discusses how this dichotomy has restricted the relevance of the field, both from theoretical and practical viewpoints. We advocate the emergence of a broader disciplinary field.

The response of semiarid grasslands to small, non-colonial herbivores has received little attention, focusing primarily on the effects of granivore assemblages on annual plant communities. We studied the long-term effects of both small and large herbivores on vegetation structure and species diversity of shortgrass steppe, a perennial semiarid grassland considered marginal habitat for small mammalian herbivores. We hypothesized that 1) large generalist herbivores would affect more abundant species and proportions of litter-bare ground-vegetation cover through non-selective herbivory, 2) small herbivores would affect less common species through selective but limited consumption, and 3) herbivore effects on plant richness would increase with increasing aboveground net primary production (ANPP). Plant community composition was assessed over a 14-year period in pastures grazed at moderate intensities by cattle and in exclosures for large (cattle) and large-plus-small herbivores (additional exclusion of rabbits and rodents). Exclusion of large herbivores affected litter and bare ground and basal cover of abundant, common and uncommon species. Additional exclusion of small herbivores did not affect uncommon components of the plant community, but had indirect effects on abundant species, decreased the cover of the dominant grass Bouteloua gracilis and total vegetation, and increased litter and species diversity. There was no relationship between ANPP and the intensity of effects of either herbivore body size on richness. Exclusion of herbivores of both body sizes had complementary and additive effects which promoted changes in vegetation composition and physiognomy that were linked to increased abundance of tall and decreased abundance of short species. Our findings show that small mammalian herbivores had disproportionately large effects on plant communities relative to their small consumption of biomass. Even in small-seeded perennial grasslands with a long history of intensive grazing by large herbivores, non-colonial small mammalian herbivores should be recognized as an important driver of grassland structure and diversity.

Trophic species in food webs are often aggregated into fewer groups, using theoretical and empirical approaches, either for modelling tractability or because of the lack of data resolution. Heterogeneities in the resolution of food webs used in the literature have led to question their use to establish general topological rules. Despite an increasing number of studies relating topology to ecosystem functioning, we still have no idea on how species’ aggregation affects our perception of network functionalities. Therefore, we re-examined the conclusions drawn from an experimental manipulation relating top-predator foraging behaviour and biomass to food-web topology (Lazzaro et al. 2009) by aggregating a 74-species network according to different criteria (taxonomy, trophic similarity, size, expertise). We found that initial significant effects and functional properties were preserved over a large portion of the aggregation gradient (2/3) despite strong variations in the topological descriptor values along the gradient. Aggregation tended to produce more type II errors (false positive) than type I errors, advocating that most effects in aggregated networks are not methodological artefacts. Aggregation by taxonomy, trophic similarity and expertise better preserved functional properties (down to 38, 30 and 17 nodes, respectively) than aggregation by size (down to 40 nodes). Our results suggest that it is possible to relate the structure of aggregated networks to ecosystem properties provided that the methodological approaches are standardized and the level of lumping does not a exceed a reasonable threshold.

While the recent inclusion of parasites into food-web studies has highlighted the role of parasites as consumers, there is accumulating evidence that parasites can also serve as prey for predators. Here we investigated empirical patterns of predation on parasites and their relationships with parasite transmission in eight topological food webs representing marine and freshwater ecosystems. Within each food web, we examined links in the typical predator–prey sub web as well as the predator–parasite sub web, i.e. the quadrant of the food web indicating which predators eat parasites. Most predator– parasite links represented ‘concomitant predation’ (consumption and death of a parasite along with the prey/host; 58–72%), followed by ‘trophic transmission’ (predator feeds on infected prey and becomes infected; 8–32%) and predation on free-living parasite life-cycle stages (4–30%). Parasite life-cycle stages had, on average, between 4.2 and 14.2 predators. Among the food webs, as predator richness increased, the number of links exploited by trophically transmitted parasites increased at about the same rate as did the number of links where these stages serve as prey. On the whole, our analyses suggest that predation on parasites has important consequences for both predators and parasites, and food web structure. Because our analysis is solely based on topological webs, determining the strength of these interactions is a promising avenue for future research.

Empirical data on the signals and processes that direct animal movement during dispersal in heterogeneous landscapes are scarce. Our understanding could benefit from utilising simulation approaches and searching for simple rules across species and landscapes. This study sought to identify general movement behaviours that optimise butterfly movements during hilltopping. This widespread dispersal-like phenomenon in butterflies, where males and virgin females ascend to mountain summits for the purpose of mating, benefits from the uniqueness of knowing the purpose (mating) and the orientation signal (topography). We used an individual-based simulation model to search for movement rules that can optimise mating success, mating time, and the success of mated females in subsequently finding habitat patches across differently structured landscapes. We found that a strong response to topography was optimal for males and virgin females to reach summits, but slight deviation from ‘perfect’, namely a certain level of randomness in response to topography, was inherently essential to avoid the risk of being trapped on local summits. The optimal response of mated females deviated only slightly from a random movement, indicating a potential weak response to topography which could be easily overlooked by field studies. Notably, the parameter values identified by the model as optimal corresponded with the observed behaviour of butterflies in the field. Finally, the optimal movement behaviours were affected by the lifespan of butterflies and the spatial distribution of host plant patches, but less so by landscape structure. We therefore suggest that modelling approaches that build on simple biological rules can facilitate the development and parameterisation of models for understanding and potentially predicting dispersal and connectivity in complex landscapes, also in circumstances where observed patterns seem complex and empirical data are scarce.

Identifying the relative importance of regional and local processes to local species diversity is a central issue to many questions in basic and applied ecology. One widely-used method is to plot local species richness against its regional richness to infer whether regional or local processes determine local diversity. However, this method increases the tendency to find regional prevalence as suggested by a recent simulation. We reanalyzed studies in the literature with an unbiased method and found no prevalence of either regional or local processes. In addition, almost 40% of the studies and 50% of the ecology textbook examples using the traditional method were misclassified. Our findings reinforce the need of alternative, novel tools identified by for instance metacommunity theory to go beyond the studies of local–regional relationships in the ecological literature that focus on the interdependence of regional and local processes.

Nature is often more diverse than expected with multiple species appearing to occupy the same niche. This observation is especially perplexing when the co-occurring species are cryptic (i.e. only distinguishable via molecular markers), because phenotypic similarity is expected to correspond with strong niche overlap. One way that phenotypically similar species can coexist is if fine-scale phenotypic differences affect how species interact with other members of the community that ultimately results in performance tradeoffs. An alternative explanation for co-occurrence is that phenotypic similarity leads to ecological equivalence allowing species to co-occur for long periods. We tested whether three phenotypically similar amphipod species that co-occur exhibit performance tradeoffs that may allow them to stably coexist in lakes. We found that despite their similarity the three species differed in how well they performed in competition with each other and their ability to avoid predation by fish and invertebrate predators. In some species comparisons, performance tradeoffs were apparent with species that perform well against heterospecifics performing poorly against predators and vice versa. We also found evidence for direct antagonistic interactions among amphipod species, in the form of wounding, which may play a role in structuring amphipod assemblages. Finally, the two species with the most similar phenotypes showed comparable responses to competitors and predators, which suggests that long-term co-occurrence via ecological equivalence may also be important in this system. Collectively, our results suggest that a mix of performance tradeoffs and ecological equivalence may allow for higher diversity than expected in amphipod assemblages.

A key attribute of riverine food webs is the downstream movement of invertebrates via the water column, or invertebrate drift. Causes of drift include benthic predation, food limitation, and perhaps passive entry, which may occur when invertebrates lose their purchase on stream substrate. However, the relative importance of drift causes is unknown, as is whether the relative importance of drift causes varies across space. Combining observational data on invertebrate herbivore and predator guild densities with in-stream experiments, we evaluated the relative importance of benthic predation, food limitation, and passive entry as proximate causes of drift for the herbivore guild across the canopy gradient of a montane stream. We found that 1) benthic predation and food limitation were both more important as causes of herbivore drift than passive entry; 2) drift caused by food limitation did not vary with riparian canopy, whereas herbivore density decreased with increasing riparian canopy, and 3) per capita drift increased linearly with increasing density, while per capita drift decreased in a negative hyperbolic fashion with increasing food, indicating that herbivore drift is proportional to herbivore density, and inversely proportional to food. We conclude that invertebrate herbivore drift was overwhelmingly an active process to improve fitness, and that herbivore food did not vary across the canopy gradient, likely because increased herbivory from larger herbivore populations at sunnier sites prevented food from accumulating.

Pulse subsidies account for a substantial proportion of resource availability in many systems, having persistent and cascading effects on consumer population dynamics, and energy flow within and across ecosystem boundaries. Although the importance of pulsed resource subsidies is well-established, the mechanisms that regulate resource fluxes across ecosystem boundaries are not well understood. The aim of our study was to determine the extent that marsh consumers regulated a marsh prey subsidy to estuarine consumers in the oligohaline reaches of an Everglades estuary. We characterized a marsh pulsed subsidy of cyprinodontoid, invertebrate and sunfish prey that move into the upper estuary from adjacent drying marshes. In response to the prey pulse, we examined the numerical, fitness and dietary responses of three focal consumers in the upper estuary; two marsh species (largemouth bass and bowfin) that accompanied the subsidy as a result of marsh drying, and one estuarine consumer (snook). At the onset of marsh drying and the prey subsidy, estuarine consumers switched diets to consume the larger marsh prey (sunfishes), while bass and bowfin maintained similar diets (cyprinodontoids and invertebrates respectively) than pre and post subsidy. From the consumption of this subsidy, bass (marsh species) and snook (estuarine species) exhibited fitness gains while bowfin did not. Although both marsh and estuarine consumers benefitted from the subsidy, we found evidence that freshwater consumers shunted some of the subsidy away from snook. Of the prey sampled in consumer stomachs, 41% of marsh prey biomass was eaten by marsh consumers, while 59% was consumed by the estuarine consumer. We conclude that the amount of the marsh prey available to estuarine consumers may be greater in the absence of marsh consumers, thus the magnitude of the prey subsidy could depend on the dynamics of the marsh consumers from donor communities.

Understanding how diversity interacts with energy supply is of broad ecological interest. Most studies to date have investigated patterns within trophic levels, reflecting a lack of food webs which include information on energy flow. We added parasites to a published marine energy-flow food web, to explore whether parasite diversity is correlated with energy flow to host taxa. Parasite diversity was high with 36 parasite taxa affecting 40 of the 51 animal taxa. Adding parasites increased the number of trophic links per species, trophic link strength, connectance, and food chain lengths. There was evidence of an asymptotic relationship between energy flowing through a food chain and parasite diversity, although there were clear outliers. High parasite diversity was associated with host taxa which were highly connected within the food web. This suggests that energy flow through a taxon may favour parasite diversity, up to a maximal value. The evolutionary and energetic basis for that limitation is of key interest in understanding the basis for parasite diversity in natural food webs and thus their role in food web dynamics.

Dispersal is a key process in metacommunity dynamics, allowing the maintenance of diversity in complex community networks. Geographic distance is usually used as a surrogate for connectivity implying that communities that are closely located are considered more prone to exchange individuals than distant communities. However, in some natural systems, organisms may be subjected to directional dispersal (air or water flows, particular landscape configuration), possibly leading close communities to be isolated from each other and distant communities to be connected. Using geographic distance as a proxy for realised connectivity may then yield misleading results regarding the role of dispersal in structuring communities in such systems. Here, we quantified the relative importance of flow connectivity, geographic distance, and environmental gradients to explain polychaete metacommunity structure along the coasts of the Gulf of Lions (northwest Mediterranean Sea). Flow connectivity was estimated by Lagrangian particle dispersal simulations. Our results revealed that this metacommunity is strongly structured by the environment at large spatial scales, and that both flow connectivity and geographic distance play an important role within homogeneous environments at smaller spatial scales. We thus strongly advocate for a wider use of connectivity measures, in addition to geographic distance, to study spatial patterns of biological diversity (e.g. distance decay) and to infer the processes behind these patterns at different spatial scales.

Pulsed resource subsidies can have profound effects on recipient communities. The effects of resource pulses are often mediated by increases in the density of consumer populations. Here we investigate several mechanisms linking experimental pulses of seaweed deposition to population-level responses in the brown anole Anolis sagrei. Subsidized lizards grew approximately 30% faster than lizards in seaweed-removal plots, but there was no effect of seaweed subsidies on survival or body condition. Breeding is strongly seasonal in A. sagrei, resulting in a limited reproductive window of opportunity. Accelerated growth allows subsidized females to reach sexual maturity earlier and thereby exploit more of this window, which is projected to double fecundity in their first year of life. These results show how changes in an individual trait can translate pulses of resource input into reproductive output. Further, they highlight the importance of seasonal timing in mechanistically linking individual-, population- and community-level responses to pulsed resource subsidies.

Although Bergmann’s rule – stating that among closely related species, the bigger ones will inhabit the colder climates/higher latitudes – was formulated for inter-specific comparisons, most analyses that tested this pattern in mammals were on an intra-specific level. To date, no large-scale taxonomy-driven cross-species evaluation of the pattern predicted by Bergmann exists. Here we show, in a dataset comprising 3561 mammal species from 26 orders, that while there is no significant correlation between latitude and body mass using conventional methods, this correlation is highly significant when the phylogenetic structure of the dataset is accounted for, thus supporting Bergmann’s claim that the rule only applies to closely related species. Analyses of different subsets indicate that the Bergmann’s rule is evident across a variety of latitude ranges. In many taxonomic subsets, when analysed alone, there is no significant correlation between body mass and latitude. In combination with both the significant relationship in the overall dataset and with results of intra-specific analyses from the literature, this suggests that Bergmann’s rule describes a fundamental principle within mammals, but that its expression has been modified by a variety of factors during mammalian diversification yet to be resolved.

Aboveground net primary productivity (ANPP) varies in response to temporal fluctuations in weather. Temporal stability of community ANPP may be increased by increasing plant species richness, but stability often varies at a given richness level implying a dependence on abundances and functional properties of member species. We measured stability in ANPP during 11 years in field plots (Texas, USA) in which we varied the richness and relative abundances of perennial grassland species at planting. We sought to identify species abundance patterns and functional traits linked to the acquisition and processing of essential resources that could be used to improve richness-based predictions of community stability. We postulated that community stability would correlate with abundance-weighted indices of traits that influence plant responses to environmental variation. Annual precipitation varied by a factor of three leading to large inter-annual variation in ANPP. Regression functions with planted and realized richness (species with > 1% of community ANPP during the final four years) explained 32% and 25% of the variance in stability, respectively. Regression models that included richness plus the fraction of community ANPP produced by the two most abundant species in combination with abundance-weighted values of either the fraction of sampled root biomass at 20–45 cm depth, leaf dry matter content (LDMC), or response to greater-than-average precipitation of plants grown in monocultures explained 58–69% (planted richness) and 58–64% (realized richness) of the variance in stability. Stability was greatest in communities that were not strongly dominated by only two species and in which plants rooted shallowly, had high values of LDMC, or responded to the wettest year with a minimal increase in ANPP. Our results indicate that the temporal stability of grassland ANPP may depend as much on species abundances and functional traits linked to plant responses to precipitation variability as on species richness alone.

In many group-living animals, within-group associations are determined by familiarity, i.e. familiar individuals, independent of genetic relatedness, preferentially associate with each other. The ultimate causes of this behaviour are poorly understood and rigorous documentation of its adaptive significance is scarce. Limited attention theory states that focusing on a given task has interrelated cognitive, behavioural and physiological costs with respect to the attention paid to other tasks. In multiple signal environments attention has thus to be shared among signals. Assuming that familiar neighbours require less attention than unfamiliar ones, associating with familiar individuals should increase the efficiency in other tasks and ultimately increase fitness. We tested this prediction in adult females of the group-living, plant-inhabiting predatory mite Phytoseiulus persimilis. We evaluated the influence of social familiarity on within-group association behaviour, activity, predation and reproduction. In mixed groups (familiar and unfamiliar), familiar predator females preferentially associated with each other. In pure groups (either familiar or unfamiliar), familiar predator females produced more eggs than unfamiliar females at similar predation rates. Higher egg production was correlated with lower activity levels, indicating decreased restlessness. In light of limited attention theory, we argue that the ability to discriminate between familiar and unfamiliar individuals and preferential association with familiar individuals confers a selective advantage because familiar social environments are cognitively and physiologically less taxing than unfamiliar social environments.

Facilitation by nurse plants plays an important role in determining community composition in severe environments. Although the unidirectional effect of nurses on beneficiary species has received considerable research interest, nurse-mediated interactions among beneficiary species (so-called indirect interactions) are less known. Consequently, community composition in nurse plant systems is generally considered as a simple consequence of the facilitative effect of the nurse even though beneficiary species may significantly contribute to community assembly and modulate the direct nurse effects on the community. In an observational study we assessed nurse effects and nurse-mediated beneficiary interactions in two contrasting nurse plant systems in dry environments using a newly developed framework. We quantified plant–plant interaction intensity using the relative interaction index (RII) at the community and species level for three Retama sphaerocarpa shrub size-classes in a semiarid shrubland and four Arenaria tetraquetra cushion plant communities differing in aspect and elevation in dry alpine gravel habitats. The observed RII was split into nurse and beneficiary effects, and related to individual mass, species frequency and abundance using generalized linear mixed models. Results showed predominantly positive nurse effects and negative beneficiary interactions. The effect size of nurse plants, however, was significantly higher than the effect size of beneficiary species in both systems. Individual plant mass and abundance of species was dependent on the combined effects of nurse and beneficiary species whereas species occurrence was related to nurse effects only. Despite evident differences, the semiarid and alpine nurse plant systems showed strong functional parallelisms. We found interdependence between the effects of nurse and beneficiary species on beneficiary plant assemblages emphasizing their combined role on community assembly in both systems. Our results highlight the need to consider indirect interactions to understand fully plant community dynamics.

The ecological attributes of two species may be similar through convergent evolution or common ancestry. The extent of similarity by descent can be evaluated by comparing them with their most closely-related outgroup in a given phylogeny. I describe a method of nested sister-group analysis for estimating ecological similarity based on landscape features or on co-distribution. The phylogeny is dissected into triplets, each comprising two sister taxa and their outgroup. For a triplet at any phylogenetic level, the similarity of sister groups with respect to some given character can be compared with their joint similarity to the outgroup to give a single test of similarity by descent. Each comparison is independent, and the full set of triplets provides a complete accounting of phylogenetic variation at all levels. This procedure was applied to 188 moderately abundant species of dicots in two independent surveys from adjoining districts of midland England, supplemented by physical surveys of landscape attributes obtained from digitized maps of the same districts. The co-distribution of sister species was consistently more positive than the co-distribution of random species pairs, demonstrating the existence of a phylogenetic signal at some level. When sister species are compared with their most closely-related outgroup, however, neither landscape attributes nor co-distribution showed any overall similarity arising from common ancestry, in the sense that ecological attributes are not generally conserved after lineage splitting. Instead, the distribution of similarity is strikingly similar to random data. The lack of ecological similarity between closely-related groups was attributed to rapid character change at or shortly after the splitting of lineages, coupled with a lack of correlation between successive lineage splits.

Communication contributes to mediate the interactions between plants and the animals that disperse their genes. As yet, seasonal patterns in plant–animal communication are unknown, even though many habitats display pronounced seasonality e.g. when leaves senescence. We thus hypothesized that the contrast between fruit displays and their background vary throughout the year in a seasonal habitat. If this variation is adaptive, we predicted higher contrasts between fruits and foliage during the fruiting season in a cerrado–savanna vegetation, southeastern Brazil. Based on a six-year data base of fruit ripening and a one-year data set of fruit biomass, we used reflectance measurements and contrast analysis to show that fruits with distinct colors differed in the beginning of ripening and the peak of fruit biomass. Black, and particularly red fruits, that have a high contrast against the leaf background, were highly seasonal, peaking in the wet season. Multicolored and yellow fruits were less seasonal, not limited to one season, with a bimodal pattern for yellow ones, represented by two peaks, one in each season. We further supported the hypothesis that seasonal changes in fruit contrasts can be adaptive because fruits contrasted more strongly against their own foliage in the wet season, when most fruits are ripe. Hence, the seasonal variation in fruit colors observed in the cerrado–savanna may be, at least partly, explicable as an adaptation to ensure high conspicuousness to seed dispersers.

Taylor’s power law, i.e. that the slope for the increase in variance with mean population size is between 1 and 2 at a logarithmic scale, provides one of the few quantitative relationships in population ecology, yet the underlying ecological mechanisms are only poorly understood. Stochastic theory of population dynamics predicts that demographic and environmental stochasticity will affect the slope differently. In a stable environment under the influence of demographic stochasticity alone the slope will be equal to 1. In large populations in which demographic variance will have a negligible effect on the dynamics the slope will approach 2. In addition, the slope will also be influenced by how the strength of density dependence is related to mean population size. To disentangle the relative contribution of these processes we estimate the mean-variance relationship for a large number of populations of British birds. The variance in population size of most species decreased with the mean due to decreased influence of demographic stochasticity at larger population sizes. Interspecific differences in demographic stochasticity was the main factor influencing variation in slopes of Taylor’s power law among species through a significant negative relationship between the slope and demographic variance. In addition, slopes were influenced by interspecific variation in life history parameters such as adult survival and clutch size. These analyses show that Taylor’s power law is generated from an interplay between stochastic and density dependent factors, modulated by life history.

Intra and interspecific variation in frugivore behaviour can have important consequences for seed dispersal outcomes. However, most information comes from among-species comparisons, and within-species variation is relatively poorly understood. We examined how large intraspecific differences in the behaviour of a native disperser, blackbuck antelope Antilope cervicapra, influence dispersal of a woody invasive, Prosopis juliflora, in a grassland ecosystem. Blackbuck disperse P. juliflora seeds through their dung. In lekking blackbuck populations, males defend clustered or dispersed mating territories. Territorial male movement is restricted, and within their territories males defecate on dung-piles. In contrast, mixed-sex herds range over large areas and do not create dung-piles. We expected territorial males to shape seed dispersal patterns, and seed deposition and seedling recruitment to be spatially localized. Territorial males had a disproportionately large influence on seed dispersal. Adult males removed twice as much fruit as females, and seed arrival was disproportionately high on territories. Also, because lek-territories are clustered, seed arrival was spatially highly concentrated. Seedling recruitment was also substantially higher on territories compared with random sites, indicating that the local concentration of seeds created by territorial males continued into high local recruitment of seedlings. Territorial male behaviour may, thus, result in a distinct spatial pattern of invasion of grasslands by the woody P. juliflora. An ex situ experiment showed no beneficial effect of dung and a negative effect of light on seed germination. We conclude that large intraspecific behavioural differences within frugivore populations can result in significant variation in their effectiveness as seed dispersers. Mating strategies in a disperser could shape seed dispersal, seedling recruitment and potentially plant distribution patterns. These mating strategies may aid in the spread of invasives, such as P. juliflora, which could, in turn, negatively influence the behaviour and ecology of native dispersers.

Animals often alternate between searching for food locally and moving over larger distances depending on the amount of food they find. This ability to switch between movement modes can have large implications on the fate of individuals and populations, and a mechanism that allows animals to find the optimal balance between alternative movement strategies is therefore selectively advantageous. Recent theory suggests that animals are capable of switching movement mode depending on heterogeneities in the landscape, and that different modes may predominate at different temporal scales. Here we develop a conceptual model that enables animals to use either an area-concentrated food search behavior or undirected random movements. The model builds on the animals’ ability to remember the profitability and location of previously visited areas. In contrast to classical optimal foraging models, our model does not assume food to be distributed in large, well-defined patches, and our focus is on animal movement rather than on how animals choose between foraging patches with known locations and value. After parameterizing the fine-scale movements to resemble those of the harbor porpoise Phocoena phocoena we investigate whether the model is capable of producing emergent home ranges and use pattern-oriented modeling to evaluate whether it can reproduce the large-scale movement patterns observed for porpoises in nature. Finally we investigate whether the model enables animals to forage optimally. We found that the model was indeed able to produce either stable home ranges or movement patterns that resembled those of real porpoises. It enabled animals to maximize their food intake when fine-tuning the memory parameters that controlled the relative contribution of area concentrated and random movements.

Species–energy theory can account for spatial variation in the abundance and community composition of animals, though the mechanisms of species–energy theory are under contention. We evaluated three competing mechanisms at the local spatial scale by conducting an in vivo light manipulation over supplemental ant nests placed in the leaf litter of a Costa Rican tropical rainforest. We found that the light environment did not alter the 10% rate of occupation of the supplemental nests, but light did alter the size of colonies and the genus-level composition of the community. Light levels in the foraging range were positively associated with colony sizes of all ants, whereas light levels directly on the nest site were predictive of the occurrence of ant genera. Colonies of specialized predators, dacetine ants, were larger in more shaded foraging environments, and the functional group of generalized myrmicines exhibited an opposite pattern, with smaller-sized colonies in response to shading. Responses of twig-dwelling ants to the light environment were most consistent with the metabolic cost hypothesis as a mechanism of species–energy theory. We found mixed support for the thermal energy availability hypothesis, and scant support for the chemical energy hypothesis, as the litter depth, a measure of prey density, was not predictive of ant responses. In summary, at the local scale, we found patterns in colony size and life history are governed by light-dependent mechanisms.

The success of a biological invasion can depend upon other invasions; and in some cases, an earlier invader may fail to spread until facilitated by a second invader. Our study documents a case whereby an invasive parasite has remained patchily distributed for decades due to the fragmented nature of available hosts; but the recent arrival of a broadly distributed alternative invasive host species provides an opportunity for the parasite to expand its range considerably. At least 20 years ago, endoparasitic pentastomids (Raillietiella frenata) were brought with their native host, the invasive Asian house gecko Hemidactylus frenatus, to the port city of Darwin in tropical Australia. These geckos rarely disperse away from human habitation, restricting the transmission of their parasites to urban environments – and thus, their pentastomids have remained patchily distributed and have only been recorded in scant localities, primarily surrounding Darwin. The recent range expansion of the invasive cane toad Rhinella marina into the Darwin area has provided an alternative host for this pentastomid. Our results show that the cane toad is a competent host for Ra. frenata– toads shed fully embryonated pentastomid eggs in their faeces – and that pentastomids are now common in cane toads near Darwin. Likely reflecting the tendency for the parasite's traditional definitive host (the Asian house gecko) and only known intermediate host (the cockroach) to reside around buildings, we found the prevalence of this parasite follows an urban distribution. Because cane toads are widely distributed through urban and rural habitat and can shed viable pentastomid eggs, the toad invasion is likely to facilitate the parasite's spread across the tropics, into areas (and additional susceptible hosts) that were previously inaccessible to it.

There is a growing recognition of the need to integrate non-trophic interactions into ecological networks for a better understanding of whole-community organization. To achieve this, the first step is to build networks of individual non-trophic interactions. In this study, we analyzed a network of interdependencies among bird species that participated in heterospecific foraging associations (flocks) in an evergreen forest site in the Western Ghats, India. We found the flock network to contain a small core of highly important species that other species are strongly dependent on, a pattern seen in many other biological networks. Further, we found that structural importance of species in the network was strongly correlated to functional importance of species at the individual flock level. Finally, comparisons with flock networks from other Asian forests showed that the same taxonomic groups were important in general, suggesting that species importance was an intrinsic trait and not dependent on local ecological conditions. Hence, given a list of species in an area, it may be possible to predict which ones are likely to be important. Our study provides a framework for the investigation of other heterospecific foraging associations and associations among species in other non-trophic contexts.

The use of species’ traits is increasing in ecological research. Many studies obtain trait data from a single source, implicitly assuming the accuracy of these data. I critically evaluate this assumption by measuring agreement among sources for trait data. I evaluate inter-source agreement for 22 traits (anatomical, behavioural, life-history and niche-related) among five authoritative data sources (two field guides, two atlases and one online resource) for 263 Canadian butterfly species. This represents the first quantitative comparison of trait data among field guides or atlases. Traits varied considerably in their agreement among sources. Some traits such as wingspan and overwinter stage were fairly consistent among sources, whereas other traits such as habitat breadth were remarkably inconsistent among sources. These findings call into question the reliability of research that relies on a single source for trait data. I offer several recommendations for how trait researchers can account for inter-source variation in trait data.

Dispersal is a central life-history trait for most animals and plants: it allows to colonize new habitats, escape from competition or avoid inbreeding. Yet, not all species are mobile enough to perform sufficient dispersal. Such passive dispersers may use more mobile animals as dispersal vectors. If multiple potential vectors are available, an active choice can allow to optimize the dispersal process and to determine the distribution of dispersal distances, i.e. an optimal dispersal kernel.

The costs and benefits of symbiotic interactions may vary with host and symbiont ontogeny. Effects of symbionts at different stages of host development or on different host demographic rates do not contribute equally to fitness. Although rarely applied, a population dynamics approach that integrates over the host life cycle is therefore necessary for capturing the net costs or benefits and, thus, the mutualistic or parasitic nature of symbioses. Using the native, disturbance-specialist grass Agrostis hyemalis, we asked how a symbiotic endophyte affected the population dynamics of its host and how imperfect vertical transmission influenced symbiont frequency in a late successional environment. A size-structured integral projection model (IPM) parameterized with experimental field data showed that greater rates of individual growth and reproduction for endophyte-symbiotic (E+) hosts outweighed their lower rates of survival, leading to a net positive effect of symbiosis on equilibrium plant population growth (slower rate of extinction). Given that populations under going successional transitions are unlikely to be at an equilibrium size structure, we also conducted transient analysis that showed an initial short-term cost to endophyte symbiosis. We used a megamatrix approach to link E− and E+ IPMs via imperfect vertical transmission and found that this parameter strongly influenced the frequency of symbiosis via complex interactions with host demographic rates. Overall, our population dynamics approach improves the ability to characterize the outcome of symbiotic interactions, and results suggest that particular attention should be paid to interactions between the rate of vertical transmission and host demography.

Public information offers a valuable means for social foragers to determine the relative quality of foraging patches. Despite much evidence that foragers use public information based on others’ feeding behavior, no experiments have examined whether foragers might use public information based on others’ competitive behavior, particularly the collective commotion that can be generated by aggregations. Such commotion could potentially provide a rich source of public information: as foragers compete in a patch with an especially high value resource, their heightened competition intensity could enable eavesdropping foragers to target this superior patch, based simply on its higher level of collective commotion. To test the hypothesis that the level of collective commotion is used as public information by eavesdropping foragers I conducted field experiments on terrestrial hermit crabs Coenobita compressus. These animals engage in collective competitive interactions in foraging patches for food and shells, generating variable levels of commotion across different quality patches. By experimentally manipulating the level of collective commotion in sham aggregations in the wild I show that a higher level of commotion is exploited by eavesdropping foragers to differentially target more valuable patches. Broadly, these results highlight an underappreciated significance of competitive by-products and higher- order collective pheno mena as forms of public information for foragers.

Predation has important cascading impacts on primary producer biomass and community composition in many ecosystems. While most studies have focused on the consequences of interspecific or density differences in predators, it is recognized that phenotypic variation within species can have strong and cascading community and ecosystem consequences at lower trophic levels. In coastal New England lakes, both the presence and life history form of the zooplanktivorous fish alewife, Alosa pseudoharengus, have strong influence on the biomass, size structure and community composition of crustacean zooplankton communities. Here we test the hypothesis that alewife presence and life history will have cascading impacts on phytoplankton biomass and community composition in a mesocosm experiment that previously reported strong biomass and compositional differences of crustacean zooplankton communities among alewife treatments. We show that alewife life history led to small but statistically significant differences in phytoplankton community composition among treatments. This compositional difference was driven primarily by an increase in the density of two edible phytoplankton genera associated with lower zooplankton biomass in the anadromous alewife treatment. Our results show that intraspecific variation in a predator can have cascading effects on primary producer communities. However we did not observe significant differences in total algal biomass.

The tritrophic model featuring plants consumed by herbivores consumed by parasitoids or predators has become the primary paradigm used to describe herbivore dynamics. However, interactions involving herbivores can be habitat- specific and plants often provide habitat, as well as food. Structural complexity of the habitat may favor predators or may allow herbivore prey to escape detection and capture. This study considered the spatial and temporal dynamics of an arctiid caterpillar, Platyprepia virginalis. The tritrophic model that includes only a tachinid parasitoid that attacks P. virginalis and the caterpillars’ primary host-plant, Lupinus arboreus, has failed to provide much insight into this system. Instead, we found that ants killed and removed many small caterpillars. Protecting caterpillars from ants increased their survival three-fold and five-fold in assays conducted during two years. Caterpillars were more likely to survive in short-term assays at sites that naturally had a deeper cover of dead and living plant material. Experiments with baits showed that ant recruitment declined as litter depth increased on average. These survey results indicated that ant predation was an important source of mortality for young caterpillars and that the presence of thick litter reduced this mortality. These results were corroborated in an experiment that manipulated litter depth and ant access to caterpillars. Previous findings that other defoliating caterpillars increased litter depth and benefitted P. virginalis are also consistent with this hypothesis. Litter acts as an important non-trophic resource, allowing caterpillars to avoid predation by ants such that wet sites with deep litter act as source populations for caterpillars. Our results show strong effects of both trophic and non-trophic interactions since plants indirectly provided limiting habitat and this heterogeneous habitat strongly affected risk of predation and ultimately caterpillar abundance and distribution.

Marking and avoiding poor-quality resources can be an important mechanism by which animals lacking a spatial memory can maximize their foraging efficiency. Here, we investigate the behaviour of larval Harmonia axyridis ladybeetles that leave chemical tracks as they forage. We built a model of an individual larva foraging for aphids, parameterized it using experimental data, and used the model to predict the effect of larval track production and detection on foraging efficiency, an important component of fitness. The model predicted that there is an optimal sensitivity of larvae to tracks which maximizes foraging efficiency; if the larva is too sensitive to tracks, it will avoid areas that might still contain resources, whereas if it is too insensitive, it will forage in areas that have depleted resources. Furthermore, the increased efficiency conferred by detecting tracks depends on the spatial arrangement of resources, with more aggregated resource distributions allowing greater benefits of detecting tracks. We tested the predictions of the model experimentally by measuring predation on aggregated versus dispersed soybean aphids by H. axyridis larvae whose ability to produce tracks was experimentally manipulated. The experiments corroborated the results of the model: larvae that could produce tracks consumed more aphids than those that could not, and this difference was greatest when aphids were aggregated among plants. Our results suggest that larval tracks play an important role in foraging efficiency, and we discuss implications for the evolution of larval track production and detection in ladybeetles.

The exploitation ecosystems hypothesis (EEH) proposes that 1) plant biomass reflects the primary productivity of an ecosystem modified by the regulating effect of herbivory, and 2) herbivore abundance reflects the productivity of plants modified by the regulating effect of predation. Primary productivity thus determines the number of trophic levels in an ecosystem and the extent to which bottom–up and top–down regulation influence the biomass ratios of adjacent and non-adjacent trophic levels (i.e. trophic cascading). We constructed an interactive model of plant (pasture), herbivore (red kangaroo Macropus rufus) and predator (dingo Canis lupus dingo), a system in which trophic cascades have been suggested to occur, and used it to test the effects of increasing stochastic variation in primary productivity and dingo culling on predictions of the EEH. The model contained four feedback loops: the predator–herbivore and herbivore–plant feedback loops, and the predator and plant density-dependent feedback loops. The equilibrium conditions along the primary productivity gradient reproduced the three zones of trophic dynamics predicted by the EEH, plus an additional zone at productivities above which the maximum density of a predator is achieved due to social regulation: that zone is characterized by increasing herbivore density and decreasing plant biomass. Culling dingoes produced trophic cascades that were strongly attenuated at primary productivities below which the maximum density of dingoes was attained. Results were robust to uncertainty in kangaroo off-take by dingoes and to the efficacy of dingo culling, but prey switching by dingoes from red kangaroos to reptiles would weaken trophic cascades. We conclude that social regulation of carnivores has important implications for expression of the EEH and trophic cascades, and that attenuation of trophic cascades increases with increasing stochasticity in primary productivity. Our model also provides a framework for understanding the conditions in which dingo-mediated trophic cascades might be expected to occur, and generates testable predictions about the effects of higher dingo densities (e.g. by stopping culling or reintroduction to former range) on kangaroo and pasture dynamics.

One of the most studied problems in population ecology has been to understand the relative roles of top–down and bottom–up forces in regulating animal populations. This has also been a key issue in studies of vole population dyna mics. Vole populations exhibit a wide variation of dynamics, from seasonal fluctuations to multiannual variations or cyclicity. One of the hypotheses to explain cyclic population dynamics is predation by the specialist predators. A common counterargument against the predation hypothesis has been the lack of conclusive observations of the time delay in the predators’ numerical response. We studied the interaction between voles and their specialist small mustelid predators, the stoat Mustela erminea and the least weasel Mustela n. nivalis, by modelling their interaction to data sets that cover large areas of Finland. Vole abundance was monitored with biannual trappings and their predators with snow-tracking. Results show a high dependence of the predators on the voles, and this connection is generally tighter in weasels than in stoats. Weasel abundance is affected most strongly by the vole abundance in previous spring, 8.5– 10 months earlier, while in stoats the effect of autumn abundance of voles, 2.5–6 months earlier, was the strongest. These results, together with the observation that the weasels’ effects on voles are stronger after a time lag of 6–9.5 than 2–4.5 months, indicate the existence of a time lag in weasels’ numerical response. A time lag in the predators’ numerical response is a necessary condition for the predators to drive population cycles in its prey, and therefore our results support the specialist predation hypothesis.

Human activities have led to massive influxes of pollutants, degrading the habitat of species and simplifying their biodiversity. However, the interaction between food web complexity, pollution and stability is still poorly understood. In this study we evaluate the effect exerted by accumulable pollutants on the relationship between complexity and stability of food webs. We built model food webs with different levels of richness and connectance, and used a bioenergetic model to project the dynamics of species biomasses. Further, we developed appropriate expressions for the dynamics of bioaccumulated and environmental pollutants. We additionally analyzed attributes of organisms’ and communities as determinants of species persistence (stability). We found that the positive effect of complexity on stability was enhanced as pollutant stress increased. Additionally we showed that the number of basal species and the maximum trophic level shape the complexity–stability relationship in polluted systems, and that in-degree of consumers determines species extinction in polluted environments. Our study indicates that the form of biodiversity and the complexity of interaction networks are essential to understand and project the effects of pollution and other ecosystem threats.

A general prediction from simple metapopulation models is that spatially synchronized forcing can spatially synchronize population dynamics and destabilize metapopulations. In contrast, spatially asynchronous forcing is predicted to decrease population synchrony and promote temporal stability and population persistence, especially in the presence of dispersal. Only recently have studies begun to experimentally address these predictions. Moreover, few studies have experimentally examined how such processes operate in the context of competition communities. Stabilizing processes may continue to operate when placed within a metacommunity context with multiple competing consumers but only at low to intermediate levels of dispersal. High dispersal rates can reverse these predictions and lead to destabilization. We tested this under controlled conditions using an experimental aquatic system composed of three competing species of zooplankton. Metacommunities experienced different levels of dispersal and environmental forcing in the form of spatially synchronous or asynchronous pH perturbations. We found support that dispersal can have contrasting effects on population stability depending on the degree to which population dynamics were synchronized in space. Dispersal under synchronous forcing or no forcing had either neutral of positive effects on spatial population synchrony of all three zooplankton species. In these treatments, dispersal reduced population stability at the local and metapopulation levels for two of three species. In contrast, asynchronously varying environments reduced population synchrony relative to unforced systems, regardless of dispersal level. In these treatments, dispersal enhanced temporal stability and persistence of populations not by reducing population synchrony but by enhancing population minima and spatial averaging of abundances. High dispersal rates under asynchronous forcing reduced the abundance of one species, consistent with increasing regional competition and general metacommunity theory. However, no effects on its stability or persistence were observed. Our work highlights the context-dependent effects of dispersal on population dynamics in varying environments.

The diversity of symbionts (commensals, mutualists or parasites) that share the same host species may depend on opportunities and constraints on host exploitation associated with host phenotype or environment. Various host traits may differently influence host accessibility and within-host population growth of each symbiont species, or they may determine the outcome of within-host interactions among coexisting species. In turn, phenotypic diversity of a host species may promote divergent exploitation strategies among its symbiotic organisms. We studied the distribution of two feather mite species, Proctophyllodes sylviae and Trouessartia bifurcata, among blackcaps Sylvia atricapilla wintering in southern Spain during six winters. The host population included migratory and sedentary individuals, which were unequally distributed between two habitat types (forests and shrublands). Visual mite counts showed that both mite species often coexisted on sedentary blackcaps, but were seldom found together on migratory blackcaps. Regardless of host habitat, Proctophyllodes were highly abundant and Trouessartia were scarce on migratory blackcaps, but the abundance of both mite species converged in intermediate levels on sedentary blackcaps. Coexistence may come at a cost for Proctophyllodes, whose load decreased when Trouessartia was present on the host (the opposite was not true). Proctophyllodes load was positively correlated with host wing length (wings were longer in migratory blackcaps), while Trouessartia load was positively correlated to uropygial gland size (sedentary blackcaps had bigger glands), which might render migratory and sedentary blackcaps better hosts for Proctophyllodes and Trouessartia, respectively. Our results draw a complex scenario for mite co-existence in the same host species, where different mite species apparently take advantage of, or are constrained by, divergent host phenotypic traits. This expands our understanding of bird–mite interactions, which are usually viewed as less dynamic in relation to variation in host phenotype, and emphasizes the role of host phenotypic divergence in the diversification of symbiotic organisms.

Many animals scatter-hoard seeds to ensure an even supply of food throughout the year and this behavior requires similar foraging decisions. Seed-traits have been shown to affect the final foraging decision but little is known about the decision process itself. Here, we first defined four sequential steps comprising the decision process of scatter-hoarding rodents: 1) upon encountering a seed, should it be ignored or manipulated; 2) if manipulated, should it be eaten in situ or removed elsewhere; 3) upon removal, how far away should it be carried; and finally 4) whether to eat or cache the removed seed. Using experimental seeds with controlled differences in size, tannin and nutrient content, we evaluated how different traits influence each step in this decision process. We found that different traits had distinct effects on each step. Seed size affected all four steps, while nutrient and tannin content primarily affected the first and third steps. By dissecting foraging behavior in relation to experimentally controlled seed-traits, we have created an effective framework within which to understand the unique relationship between scatter-hoarding rodents that both predate and disperse plant seeds.

Plant functional traits are increasingly used to evaluate changes in ecological and ecosystem processes. However our understanding of how functional traits coordinate across different plant structures, and the implications for trait-driven processes such as litter decomposition, remains limited. We compared the functional traits of green leaves and leaf, twig and wood litter among 27 co-occurring tree species from New Zealand, and quantified the loss of mass, N and P from the three litter types during decomposition. We hypothesised that: a) the functional traits of green leaves, and leaf, twig and wood litter are co-ordinated so that species which produce high quality leaves and leaf litter will also produce high quality twig and wood litter, and b) the decomposability of leaf, twig and wood litter is coordinated because breakdown of all three litter types is driven by similar combinations of traits. Trait variation across species was co-ordinated between leaves, twigs and wood when angiosperm and gymnosperm species were considered in combination, or when angiosperms were considered separately, but trait coordination was poor for gymnosperms. There was little coordination among the three litter types in their decomposability, especially when angiosperms and gymnosperms were considered separately; this was caused by the decomposability of each of the three litter types, at least partially, being driven by different functional traits or trait combinations. Our findings indicate that although interspecific variation in the functional traits of trees can be coordinated among leaves, twigs and wood, different or unrelated traits predict the decomposition of these different structures. Furthermore, leaf-level analyses of functional traits are not satisfactory proxies for function of whole trees and related ecological processes. As such, efforts to understand how tree species influence C, N and P dynamics in forested ecosystems through the decomposition pathway need to consider functional traits of other plant structures.

Sperm-dependent asexual species must coexist with a sexual species (i.e. a sperm source) to reproduce. The maintenance of this coexistence, and hence the persistence of sperm-dependent asexual species, may depend on ecological niche separation or preference by males for conspecific (i.e. sexual) mates. We first modified an analytical model to consider both of these mechanisms acting simultaneously on the coexistence of the two species. Our model indicates that a small amount of niche separation between parental species and hybrids can facilitate coexistence by weakening the requirement for male mate preference. We also estimated niche separation empirically in the Chrosomus (formerly Phoxinus) sexual-asexual system based on diet overlap between sperm-dependent asexuals and their two sexual host species. Diet overlap between the sexual species was not significant in either lake, whereas the sperm-dependent asexual had an intermediate niche that overlapped significantly, but somewhat asymmetrically, with both sexual species. These empirical results were then used to parameterize our analytical model to predict the minimum strength of male mate preference required to maintain coexistence in each lake. Some male mate preference is likely required to maintain coexistence in the Chrosomus system, but the minimum required preference depends on the severity of density dependence. Future empirical work on understanding coexistence in sperm-dependent asexual systems would benefit from taking both niche separation and mate choice into account, and from simultaneous empirical estimates of male mate choice, niche separation, and density dependence.

Snow cover is a key environmental component for tundra wildlife that will be affected by climate change. Change to the snow cover may affect the population dynamics of high-latitude small mammals, which are active throughout the winter and reproduce under the snow. We experimentally tested the hypotheses that a deeper snow cover would enhance the densities and winter reproductive rates of small mammals, but that predation by mustelids could be higher in areas of increased small mammal density. We enhanced snow cover by setting out snow fences at three sites in the Canadian Arctic (Bylot Island, Nunavut, and Herschel Island and Komakuk Beach, Yukon) over periods ranging from one to four years. Densities of winter nests were higher where snow depth was increased but spring lemming densities did not increase on the experimental areas. Lemmings probably moved from areas of deep snow, their preferred winter habitat, to summer habitat during snow melt once the advantages associated with deep snow were gone. Our treatment had no effect on signs of reproduction in winter nests, proportion of lactating females in spring, or the proportion of juveniles caught in spring, which suggests that deep snow did not enhance reproduction. Results on predation were inconsistent across sites as predation by weasels was higher on the experimental area at one site but lower at two others and was not higher in areas of winter nest aggregations. Although this experiment provided us with several new insights about the impact of snow cover on the population dynamics of tundra small mammals, it also illustrates the challenges and difficulties associated with large-scale experiments aimed at manipulating a critical climatic factor.

The scale-dependent species abundance distribution (SAD) is fundamental in ecology, but few spatially explicit models of this pattern have thus far been studied. Here we show spatially explicit neutral model predictions for SADs over a wide range of spatial scales, which appear to match empirical patterns qualitatively. We find that the assumption of a log-series SAD in the metacommunity made by spatially implicit neutral models can be justified with a spatially explicit model in the large area limit. Furthermore, our model predicts that SADs on multiple scales are characterized by a single, compound parameter that represents the ratio of the survey area to the species’ average biogeographic range (which is in turn set by the speciation rate and the dispersal distance). This intriguing prediction is in line with recent empirical evidence for a universal scaling of the species-area curve. Hence we hypothesize that empirical SAD patterns will show a similar universal scaling for many different taxa and across multiple spatial scales.

Root herbivores can influence both the performance and the behaviour of parasitoids of aboveground insect herbivores through changes in aboveground plant quality and in the composition of the plant's odour blend. Here we show that root herbivory by Delia radicum larvae did not influence the innate preferences for plant odours of the two closely related parasitoid species Cotesia glomerata and C. rubecula, but did affect their learned preferences, and did so in an opposite direction. While C. glomerata learned to prefer the odour of plants with intact roots, C. rubecula learned to prefer the odour of root-infested plants. The learned preference of C. glomerata for the odour of plants with intact roots matches our previously published result of its better performance when developing in P. brassicae hosts feeding on this plant type. In contrast, the relatively stronger learned preference of C. rubecula for the odour of root-infested plants cannot be merely explained by its performance, as the results of our present study indicate that D. radicum root herbivory did not influence the performance of C. rubecula nor of its host P. rapae. Our results stress the importance of assessing the influence of root herbivores on both innate and learned responses of parasitoids to plant odours.

Research on the role of top–down (predation) and bottom–up (food) effects in food webs has led to the understanding that the variability of these effects in space and time is a fundamental feature of natural systems. Consequently, our measurement tools must allow us to evaluate the effects from a dynamical perspective. A population-dynamics approach may be appropriate to the task. More specifically, because food and predators both affect birth rate, birth rate dynamics may be a key to understanding their impact on the population of interest. Based on the Edmondson–Paloheimo model for birth rate, we propose a new population metric to assess the relative strength of top–down vs bottom–up effects. The metric is the ratio of contributions of changes in proportion of adults and fecundity to change in birth rate. Proportion of adults reflects a top–down effect (predators are assumed to be size-selective), fecundity reflects a bottom–up effect, and birth rate appears as a common currency with which to compare the former and the latter. Using microcosm experiments and computer simulations on the cladoceran Daphnia, we calibrate the metric and show that, in both types of tests, the ratio of contributions is typically 0.5–0.7 under a strong bottom–up effect and 2.0–2.2 under a strong top–down effect. This provides experimental evidence that the ratio of contributions may allow one to distinguish a strong top–down effect from a strong bottom–up effect.

The effects of plant genotype and environmental factors on tri-trophic interactions have usually been investigated separately, limiting our ability to compare the relative strength of these effects as well as their potential to interactively shape arthropod communities. We studied the interactions among the herb Ruellia nudiflora, a seed predator, and its parasitoids using 14 maternal plant families grown in a common garden. By fertilizing half of the plants of each family and subsequently recording fruit number, seed predator number, and parasitoid number per plant, we sought to compare the strength of plant genetic effects with those of soil fertility, and determine if these factors interactively shape tri-trophic interactions. Furthermore, we evaluated if these bottom–up factors influenced higher trophic levels through changes in abundance across trophic levels (density-mediated) or changes in the function of species interactions (trait-mediated). Plant genetic effects on seed predators and parasitoids were stronger than fertilization effects. Moreover, we did not find plant genetic variation for fertilization effects on fruit, seed predator, or parasitoid abundance, showing that each factor acted independently on plant resources and higher trophic levels. Both bottom–up forces were transmitted via density-mediated effects where increased fruit number from fertilization and plant genetic effects increased seed predator and parasitoid abundance; however, seed predator attack was density-dependent, while parasitoid attack was density-independent. Importantly, there was evidence (marginally significant in one case) that fertilization modified the function of plant-seed predator and seed predator–parasitoid interactions by increasing the number of seed predators per fruit and decreasing the number of parasitoids per seed predator, respectively. These findings show that plant genetic and soil fertility effects cascaded up this simple food chain, that plant genetic effects were stronger across all trophic levels, and that these effects were transmitted independently and through contrasting mechanisms.

Despite recent findings on the ecological relevance of within population diet variation far less attention has been devoted to the role diet variation for ecological services. Seed dispersal is a key ecological service, affecting plant fitness and regeneration based on foraging by fruit-eating vertebrates. Here we used a network approach, widely used to understand how seed-dispersal is organized at the species level, to gain insights into the patterns that emerge at the individual-level. We studied the individual fruit consumption behavior of a South American didelphid Didelphis albiventris, during the cool–dry and warm–wet seasons. In species–species networks the heterogeneity in specialization levels generates patterns such as nestedness and asymmetry. Because generalist populations may be comprised of specialized individuals, we hypo thesized that network structural properties, such as nestedness, should also emerge at the individual level. We detected variation in fruit consumption that was not related to resource availability, ontogenetic or sexual factors or sampling biases. Such variation resulted in the structural patterns often found in species–species seed-dispersal networks: low connectance, a high degree of nestedness and the absence of modules. Moreover structure varied between the warm–wet and cool–dry seasons, presumably as a consequence of seasonal fluctuation in fruit availability. Our findings suggest individuals may differ in selectivity causing asymmetries in seed dispersal efficiency within the population. In this sense the realized dispersal would differ from the expected dispersal estimated from their average dispersal potential. Additionally the results suggest possible frequency-dependent effects on seed dispersal that might affect individual plant performance and plant community composition.

The coexistence of multiple seed size strategies within plant communities have been considered puzzling, based on a theoretical expectation of the existence of an optimal seed size under each set of specific environmental conditions. A model aimed at explaining the coexistence of different seed sizes has been suggested, where a seed size – seed number tradeoff is connected to a tradeoff between competition and colonization, leading to a competitive advantage in larger-seeded species and a colonization advantage in smaller-seeded species. Recently an alternative model has been suggested, based on a tradeoff between stress tolerance and fecundity, associated with the variation from large to small seeds. Here, we examine the role of seed size for recruitment in two-species contests subjected to various treatments. In a garden experiment seeds of 14 plant species were combined pair-wise into seven pairs, each with one larger-seeded species and one smaller-seeded species. Each species-pair was sown with sparse and dense seed densities and subjected to different treatments of shading and litter. Recruitment was recorded during two years. Our results showed a general advantage of larger-seeded species over smaller-seeded species. This seed size advantage increased in treatments with litter, whereas there were minor effects of shade, and no effect of seed density was found. We thus found little support for a density dependent seed size game as assumed in models of a competition-colonization tradeoff, whereas our results fit well with a model based on a tradeoff between stress tolerance and fecundity. Our experiment provides novel empirical data to theoretical models on co-existence between multiple seed size strategies.

Anthropogenically induced global climate change has important implications for marine ecosystems with unprecedented ecological and economic consequences. Climate change will include the simultaneous increase of temperature and CO₂ concentration in oceans. However, experimental manipulations of these factors at the community scale are rare. In this study, we used an experimental approach in mesocosms to analyse the combined effects of elevated CO₂ and temperature on macroalgal assemblages from intertidal rock pools. Our model systems were synthetic assemblages of varying diversity and understory component and canopy species identity. We used assemblages invaded by the non-indigenous canopy forming alga Sargassum muticum and assemblages with the native canopy species Cystoseira tamariscifolia. We examined the effects of both climate change factors on several ecosystem functioning variables (i.e. photosynthetic efficiency, productivity, respiration and biomass) and how these effects could be shaped by the diversity and species identity of assemblages. CO₂ alone or in combination with temperature affected the performance of macroalgae at both individual and assemblage level. In particular, high CO₂ and high temperature (20°C) drastically reduced the biomass of macroalgal assemblages and affected their productivity and respiration rates. The identity of canopy species also played an important role in shaping assemblage responses, whereas species richness did not seem to affect such responses. Species belonging to the same functional effect group responded differently to the same environmental conditions. Data suggested that assemblages invaded with S. muticum might be more resistant in a future scenario of climate change. Thus, in a future scenario of increasing temperature and CO₂ concentration, macroalgal assemblages invaded with canopy-forming species sharing response traits similar to those of S. muticum could be favoured.

Prey quality has previously been shown to affect the growth and reproduction of predatory arthropods, however relatively little is known about the specific nutrients responsible for these effects. We tested if the macronutrient content (i.e. lipid and protein) of live prey affected mate attraction, reproductive behavior, egg production and nutrient reserves of adult female praying mantids, Pseudomantis albofimbriata. Females on a high-protein diet produced more than twice as many eggs as females on a high-lipid diet despite being fed the same overall biomass of prey. Furthermore, the lipid and protein composition of eggs and the female body was directly related to the diet that females were fed (i.e. high lipid content on the high-lipid diet). Even more striking was the effect of diet treatment on the number of males attracted to females – only one male was attracted to females on the high-lipid treatment and 56 males were attracted to females on the high-protein treatment. Although it is not unexpected that females with more eggs would attract more males, the extreme nature of this difference is certainly surprising because previous studies have shown that females with only a couple of eggs can attract multiple males. Hence, our results suggest that female pheromone production may be affected by the quality/nutritional composition of eggs rather than simply the number of eggs. We found no significant difference in any of the other behaviours measured during mating trials, including the frequency of sexual cannibalism. The positive effects of prey protein content on mate attraction and egg production suggest that praying mantids might be expected to choose more protein-biased prey in nature or, if prey choice is limited, to have higher reproductive output or population growth in communities dominated by protein-rich prey.

Amensalism may be common between non-trophically linked animals in natural ecosystems, where variation among species in body sizes and foraging modes may give rise to one-sided interference. However, species and ecosystem-level consequences of animal–animal amensalism are largely unknown. In a Tibetan alpine meadow, dominant herbivorous grasshoppers trigger a death feigning anti-predator response of co-occurring grassland caterpillars despite posing no consumptive threat. We hypothesized that: 1) grasshoppers reduce the performance of caterpillars while incurring no cost to themselves; and 2) this amensalism reduces top–down control of plant composition and biomass. We tested these hypotheses by factorial manipulation of both herbivores within replicate field enclosures. Grasshoppers significantly suppressed caterpillar feeding, growth rate, survival, reproductive effort and delayed metamorphosis. In contrast, grasshopper performance was unaffected by the caterpillars. Suppression of caterpillar feeding decreased overall herbivore suppression of plant biomass by 58% and shifted the functional composition of the plant community (i.e. increased sedge: forb ratio). These results suggest that consideration of non-trophic interactions such as amensalism will help predict the consequences of species losses for the structure and functioning of ecosystems.

Apparent competition between prey is hypothesized to occur more frequently in environments with low densities of preferred prey, where predators are forced to forage for multiple prey items. In the arctic tundra, numerical and functional responses of predators to preferred prey (lemmings) affect the predation pressure on alternative prey (goose eggs) and predators aggregate in areas of high alternative prey density. Therefore, we hypothesized that predation risk on incidental prey (shorebird eggs) would increase in patches of high goose nest density when lemmings were scarce. To test this hypothesis, we measured predation risk on artificial shorebird nests in quadrats varying in goose nest density on Bylot Island (Nunavut, Canada) across three summers with variable lemming abundance. Predation risk on artificial shorebird nests was positively related to goose nest density, and this relationship was strongest at low lemming abundance when predation risk increased by 600% as goose nest density increased from 0 to 12 nests ha⁻¹. Camera monitoring showed that activity of arctic foxes, the most important predator, increased with goose nest density. Our data support our incidental prey hypothesis; when preferred prey decrease in abundance, predator mediated apparent competition via aggregative response occurs between the alternative and incidental prey items.

When searching for resources in heterogeneous environments, animals must rely on their abilities to detect the resources via their sensory systems. However, variation in the strength of the sensory cue may be mediated by the physical size of the resource patch. Patch detection of insects are often predicted by the scaling of sensory cues to patch size, where visual cues has been proposed to scale proportional to the diameter of the patch. The scaling properties of olfactory cues are, however, virtually unknown. Here, we investigated scaling rules for olfactory information in a gradient of numbers of odour sources, relevant to odour-mediated attraction under field conditions. We recorded moth antennal responses to sex pheromones downwind from pheromone patches and estimated the slope in the scaling relationship between the effective length of the odour plumes and the number of odour sources. These measurements showed that the effective plume length increased proportional to the square root of the number of odour sources. The scaling relationship, as estimated in the field experiment, was then evaluated against field data of the slope in the relationship between trap catch and release rate of chemical attractants for a wide range of insects. This meta-analysis revealed an average slope largely consistent with the estimated scaling relationship between the effective plume length and the number of odour sources. This study is the first to estimate the scaling properties of olfactory cues empirically and has implications for understanding and predicting the spatial distributions of insects searching by means of olfactory cues in heterogeneous environments.

After decades of intensive research, the relative importance of top–down and bottom–up control for structuring ecological communities is still a particularly disputed issue among ecologists. In our study, we determine the relative role of bottom–up and top–down forces in structuring the composition of 13 arthropod functional groups (FG) comprising different trophic consumer levels. Based on species-specific plant biomass and arthropod abundance data from 50 plots of a grassland biodiversity experiment, we quantified the proportions of bottom–up and top–down forces on consumer FG composition while taking into account direct and indirect effects of plant diversity, functional diversity, community biomass, soil properties and spatial arrangement of these plots. Variance partitioning using partial redundancy analysis explained 21–44% of total variation in arthropod functional group composition. Plant-mediated bottom–up forces accounted for the major part of the explainable variation within the composition of all FGs. Predator-mediated top–down forces, however, were much weaker, yet influenced the majority of consumer FGs. Plant functional group composition, notably legume composition, had the most important impact on virtually all consumer FGs. Compared to plant species richness and plant functional group richness, plant community biomass explained a much higher proportion of variation in consumer community composition.

The changes in plant–plant interactions along environmental gradients have been a focus of recent ecological research. It has been suggested that both above- and below-ground competition and their interplay vary along gradients, but few studies have investigated this idea, and in most cases, the role of facilitation has not been considered, despite its importance in high stress environments. Here we used two-layer ‘zone-of-influence’ models to simulate the effects of facilitation, size-asymmetry of competition, abiotic stress, resource availability and the balance of root–shoot growth on shoot and root interactions and their interplay along an environmental gradient. In the absence of facilitation, shoot and total competition became weaker, while root competition and the interplay between shoot and root competition were unchanged under increasing stress when root competition was completely symmetric. In contrast, shoot, root, total interactions and the interplay between shoot and root interactions were all negative, and they increased with increasing stress when root competition was size-symmetric. When facilitation was included in the models, net effects of shoot, root, total interactions and the interplay of root–shoot interactions were very different from those without facilitation, and many were positive under highly stressful conditions. The type of stress (non-resource or resource) did not significantly influence the simulation results. Our study provides an alternative interpretation of the interplay between above- and below-ground plant–plant interactions across an environmental gradient.

In many plants, leaves that are young and/or old (senescent) are not green. One adaptive hypothesis proposed that leaf color change could be a warning signal reducing insect attack. If leaf coloration involves less herbivory, it remains unclear why leaves in many species are constantly green. To examine whether green leaves reduce herbivory by physical defense as an alternative to the supposed warning signal of red leaves, we conducted comparative analyses of leaf color and protective tissues of 76 woody species in spring. The protective features (trichomes, enhanced cuticle and multiple epidermis) and the distribution of red pigments within leaves were examined in both young and mature leaves. We observed that redness was more frequent in young leaves than in senescent leaves. Compared to 36 species with red young leaves, 40 species with green young leaves showed a significantly higher incidence of enhanced cuticle and trichomes in both phylogenetic and non-phylogenetic analyses. The phylogenetic analysis indicated that the multiple origins of mechanical protection were generally associated with loss of red coloration. Our finding of relatively poor mechanical protection in red young leaves provides additional evidence for the adaptive explanation of leaf color change.

Omega-3 (ω3) and -6 (ω6) polyunsaturated fatty acids (PUFA) are essential for all aquatic animals, but their dietary availability can be highly heterogeneous in space and time. The way consumers retain PUFA across such heterogeneous feeding conditions remains poorly understood. In a series of feeding experiments, we investigated how retention efficiencies (i.e. amount in consumer biomass/amount ingested) of PUFA and bulk carbon responded to heterogeneous PUFA intake in Daphnia magna. Heterogeneous PUFA intake was achieved by exposing D. magna to algal diets of different PUFA content and composition for specific time periods. The retention efficiency of carbon did not change among dietary treatments. At shorter exposure to PUFA-rich diet, retention efficiencies of most PUFA were 2–3 times higher than that of bulk carbon, clearly indicating PUFA bioaccumulation in D. magna. Increasing exposure to PUFA-rich diet caused exponential decrease of retention efficiencies for most PUFA. However, D. magna receiving more PUFA were richer in these compounds despite lower retention efficiency. Eicosapentaenoic (20:5ω3) and arachidonic acid (20:4ω6) and their precursors were always supplied in the same proportions (3.6:1), but the 20:5ω3/20:4ω6 ratio in D. magna (an important measure of nutritional quality for consumers) increased with exposure time to these PUFA from 2.2:1 to 3.8:1, thus eventually matching the diet. Our results suggest that D. magna is an efficient gatherer, accumulator, and repository of PUFA under low/fragmented dietary availability. However, at higher availabilities, PUFA are not always bioaccumulated in D. magna. Hence, the efficiency of PUFA transfer by daphnids in food webs may depend on temporal PUFA availability and its range of variation. Finally, we show that heterogeneity in PUFA intake may also affect higher trophic levels by influencing nutritionally critical PUFA ratios of zooplankton.

The decomposition rates of plant litter mixtures often deviate from the averaged rates of monocultures of their component litter species. The mechanisms behind these non-additive effects in decomposition of litter mixtures are lively debated. One plausible explanation for non-additive effects is given by the improved microenvironmental condition (IMC) theory. According to this theory, plant litter species, whose physical characteristics improve the microclimatic conditions for decomposers, will promote the decomposition of their co-occurring litter species. We tested the IMC theory in relation to leaf litter and soil moisture in two contrasting moisture conditions in a dry subarctic mountain birch forest with vascular plant leaf litters of poor and high quality. The non-additive effects in mass loss of litter mixtures increased when moisture conditions in litter and soil became more favourable for plant litter decomposition. The sign of this increase (antagonistic or synergistic) in non-additive effects was more predictable for litter mixtures of poor litter quality. Although the specific mechanisms underlying the IMC theory depended on the litter quality of the litter mixtures, a standardized water holding capacity (WHC) was the litter trait most closely related to the non-additive effects in mixtures of both poor and high quality litter types. Furthermore, we found that higher dissimilarity in WHC traits between the component litter species in a mixture increased synergistic effects in litter mixtures under limiting moisture conditions. However, under improved moisture conditions, increased antagonistic effects were observed. Thus, we found clear support for the IMC theory and showed that climatic conditions and leaf litter physical traits determine whether the non-additive effects in litter mixtures are antagonistic or synergistic. Our study emphasizes the need to include litter physical traits into predictive models of mixing effects on plant litter decomposition and in general suggests climate specificity into these models.

Because species interactions are often context-dependent, abiotic factors such as temperature and biotic factors such as food quality may alter species interactions with potential consequences to ecosystem structure and function. For example, altered predator–prey interactions may influence the dynamics of trophic cascades, affecting net primary production. In a three-year field experiment, we manipulated a plant–grasshopper–spider food chain in mesic tallgrass prairie to investigate the effects of temperature and food quality on grasshopper performance, and to understand the direct and indirect tritrophic interactions that contribute to trophic cascades. Because spiders are active at cooler temperatures than grasshoppers in our system, we hypothesized that predator effects would be strongest in cooled treatments, and weakest in warmed treatments. Grasshopper spider interactions were highly context-dependent and varied significantly with food quality, temperature treatment and year. Spiders most often reduced grasshopper survival in the cooled and ambient temperature treatments, but had little to no effect on grasshopper survival in the warmed treatments, as hypothesized. In some years, plants compensated for grasshopper herbivory and trophic cascades were not observed despite significant effects of predators on grasshopper survival. However, in the year they were observed, trophic cascades only occurred in cooled treatments where predator effects on grasshoppers were strongest. Predicting ecosystem responses to climate change will require an understanding of how temperature influences species interactions. Our results demonstrate that changes in daily temperature regimes can alter predator–prey interactions among arthropods with consequences for ecosystem processes such as primary production and the relative importance of top–down and bottom–up processes.

Most species introductions are not expected to result in invasion, and species that are invasive in one area are frequently not invasive in others. However, cases of introduced organisms that failed to invade are reported in many instances as anecdotes or are simply ignored. In this analysis, we aimed to find common characteristics between non-invasive populations of known invasive species and evaluated how the study of failed invasions can contribute to research on biological invasions. We found intraspecific variation in invasion success and several recurring explanations for why non-native species fail to invade; these included low propagule pressure, abiotic resistance, biotic resistance, genetic constraints and mutualist release. Furthermore, we identified key research topics where ignoring failed invasions could produce misleading results; these include studies on historical factors associated with invasions, distribution models of invasive species, the effect of species traits on invasiveness, genetic effects, biotic resistance and habitat invasibility. In conclusion, we found failed invasions can provide fundamental information on the relative importance of factors determining invasions and might be a key component of several research topics. Therefore, our analysis suggests that more specific and detailed studies on invasion failures are necessary.

Habitat-forming invasive species have complex impacts on native communities. Positive above ground and negative below ground impacts are reported, suggesting that habitat-forming invasive species may affect community components differently. Furthermore, such effects may vary depending on the density of the invader. We determined the responses of community components to different densities of the invasive green alga Caulerpa taxifolia in southeastern Australia. Initially we investigated differences in soft-sediment faunal communities (above and below ground) across a biomass gradient at two invaded sites. Caulerpa taxifolia biomass was positively associated with the composition and abundance of the epifaunal community, but negatively correlated with the abundance of infauna. To examine the response of common community members in more detail, we caged two species of mollusk (the infaunal bivalve, Anadara trapezia and the epifaunal gastropod, Batillaria australis) across the same biomass gradient to determine lethal and sublethal effects of C. taxifolia biomass on individuals. Survivorship of A. trapezia was low when C. taxifolia was above 300 g m⁻². Negative sublethal effects were also density-dependent with A. trapezia tissue weight being lowest above this same C. taxifolia biomass. The proportion of B. australis surviving was unaffected by C. taxifolia biomass. However, the total number of live B. australis recovered in cages increased as C. taxifolia biomass increased, providing further evidence of positive density dependent effects (in line with the survey data) of C. taxifolia on epifauna. Finally, we removed C. taxifolia from plots of differing C. taxifolia biomass and followed community change for 5 months. Community change following C. taxifolia removal was also density dependent as recovery 5 months post-removal depended on the initial biomass of C. taxifolia, suggesting a lag in the recovery of communities due to residual environmental effects post-removal (i.e. hysteresis). We have shown that the effects of a habitat-forming invasive species are biomass dependent and also affect community components differently, suggesting that, globally, the impact of these types of invaders may be context dependent.

Beta diversity and nestedness are central concepts of ecology and biogeography and evaluation of their relationships is in the focus of contemporary ecological and conservation research. Beta diversity patterns are originated from two distinct processes: the replacement (or turnover) of species and the loss (or gain) of species leading to richness differences. Nested distributional patterns are generally thought to have a component deriving from beta diversity which is independent of replacement processes. Quantification of these phenomena is often made by calculating a measure of beta diversity, and the resulting value being subsequently partitioned into a contribution by species replacement plus a fraction shared by beta diversity and nestedness. Three methods have been recently proposed for such partitioning, all of them based on pairwise comparisons of sites. In this paper, the performance of these methods was evaluated on theoretical grounds and tested by a simulation study in which different gradients of dissimilarity, with known degrees of species replacement and species loss, were created. Performance was also tested using empirical data addressing land-use induced changes in endemic arthropod communities of the Terceira Island in the Azores. We found that the partitioning of β_cc (dissimilarity in terms of the Jaccard index) into two additive fractions, β_-3 (dissimilarity due to species replacement) plus β_rich (dissimilarity due to richness differences) reflects the species replacement and species loss processes across the simulated gradients in an ecologically and mathematically meaningful way, whilst the other two methods lack mathematical consistency and prove conceptually self-contradictory. Moreover, the first method identified a selective local extinction process for endemic arthropods, triggered by land-use changes, while the latter two methods overweighted the replacement component and led to false conclusions. Their basic flaw derives from the fact that the proposed replacement and nestedness components (deemed to account for species loss) are not scaled in the same way as the measure that accounts for the total dissimilarity (Sørensen and Jaccard indices). We therefore recommend the use of β_cc=β_-3+β_rich, since its components are scaled in the same units and their responses are proportional to the replacement and the gain/loss of species.

Intraspecific variation in body size is common in animals and plants. Body size affects trophic interactions like foraging ability and vulnerability to predation, which in turn affect individual fitness as well as population stability and extinction risk. Experimental and theoretical work has shown that the size distribution of individuals within cohorts is strongly influenced by intraspecific competition for resources, often leading to skewed frequency distributions. However, little is known about the effects of environmental factors such as climate and eutrophication on the cohort size-structure of natural populations. We use a long-term time series of scientific monitoring of a freshwater fish (European perch Perca fluviatilis) to investigate the effects of density dependence, predation, nutrient availability, climate and the timing of spawning on the cohort size distributions. We find that the mean length of the fish is best predicted by the extrinsic factors phosphorus concentration and summer temperature, and the densities of the different age-classes, whereas the skewness of the length distribution is best predicted by phosphorus concentration, summer temperature, abundance of small fish, and the timing of spawning. Higher nutrient levels, temperatures and densities of small fish increase food availability and thus reduce competition, which is reflected in increased mean length and decreased skewness. The timing of spawning affects skewness presumably through changes in the initial size variation of the cohort and the length of the first growth season. Our results indicate that higher temperatures increase the mean length and decrease skewness due to the concurrent eutrophication of the lake. The study thereby highlights the potential impact of human-induced environmental change on the size structure of fish populations. More studies are needed to understand better the complex mechanisms through which these factors alter the intensity of intraspecific competition in fish communities.

Estimation of extinction thresholds arising from Allee effects (Allee thresholds) and related probabilities of population extinction is notoriously difficult. One way is to analyze adequately parameterized population models. Traditionally, a point estimate is substituted for the Allee effect strength in such models. However, each point estimate entails an underlying uncertainty. We explore how accounting for this uncertainty affects the probability of population extinction, and show that this probability decreases sigmoidally with increasing population density, even in the absence of any stochasticity. Deviations from when only a point estimate of the Allee effect strength is used can be significant, unless stochasticity is added and the stochastic noise intensity is high. Significant deviations from when only a point estimate is used also occur when the Allee threshold and the environmental carrying capacity of the species are close enough one to another. We also show that the impact of the uncertainty in the Allee effect strength estimate increases as the Allee effect strength itself increases and decreases as the species recovery potential increases. This is not a good news, since we would like to preferentially and efficiently manage slowly recovering populations prone to strong Allee effects. Still, there is a way to come up with relatively good Allee threshold estimates. Besides an obvious option of collecting as many data as possible, the impact of the uncertainty can be mitigated by diversifying Allee effect experiments such that we put more emphasis on larger size groups. This is somewhat surprising, given that frequent complaints on the (im)possibility of detecting Allee effects concern difficulties in locating, observing and experimenting on rare populations. Our results extend current theory surrounding Allee effects and have broad ramifications for applied ecology.

The analysis of animal movement is a large and continuously growing field of research. Detailed knowledge about movement strategies is of crucial importance for understanding eco-evolutionary dynamics at all scales – from individuals to (meta-)populations. This and the availability of detailed movement and dispersal data motivated Nathan and colleagues to published their much appreciated call to base movement ecology on a more thorough mechanistic basis. So far, most movement models are based on random walks. However, even if a random walk might describe real movement patterns acceptably well, there is no reason to assume that animals move randomly. Therefore, mechanistic models of foraging strategies should be based on information use and memory in order to increase our understanding of the processes that lead to animal movement decisions.

Plant individuals rely on pollinator services for their reproduction and often have to share these services with co-occurring neighbours, creating complex indirect plant–plant interactions. Many current theoretical models focus on the effect of floral resources’ density on the net outcome of these indirect plant–plant interactions, often neglecting the identity of plant species in the communities and especially the species’ spatial pattern. To fill this gap, we created a spatially explicit model whose goal was to study the interplay between relative densities and spatial distribution patterns of two plant species differing in their attractiveness for pollinators. Since theory predicts that pollinator behaviour strongly governs the outcome of pollination, we allowed the pollinators to systematically change their plant preferences based on their foraging experience. Thus the interplay between density and spatial patterns of plants was tested over a continuum of behaviours from specialists to generalists. Our most striking finding was that reproductive success of the less attractive species was affected in an opposite way by spatial patterns depending on whether the species had relatively low or high densities. Namely, when the less attractive species was highly abundant, its survival was higher when aggregated in large monospecific patches than when uniformly distributed. On the other hand, when the attractive species was more abundant, the less attractive species survived better when uniformly distributed. These results were consistent as long as the scale of the plant spatial aggregation was similar to or larger than the pollinators’ detection range. Our results suggest that aggregated plant spatial patterns manipulate pollinator behaviour by trapping them within monospecific patches. This effect was sufficiently strong to enhance the survival of a competitively inferior species and hence to act in a way similar to the more familiar niche or temporal separation among plant species.

Genetic diversity has emerged as an important source of variation in the ecological properties of populations, but there are few studies of genetic diversity effects on colonisation processes. This relative scarcity of studies is surprising given the influence of colonisation on species coexistence, invasion, and population persistence. Here, we manipulated relatedness in experimental populations of colonising larvae in four sessile marine invertebrates. We then examined the influence of coloniser relatedness on the number, spatial arrangement and phenotype of colonisers following permanent settlement. Overall, relatedness influenced colonisation in all four species, but the effects of relatedness on colonisation differed among species. The variable responses of species to manipulations of relatedness likely reflect differences in intensity of inter- and intra-specific competition among adults, as well as the differential consequences of larval behaviours for each species. Relatedness appears to play an underappreciated role in the colonisation process, and we recommend that future studies of genetic diversity effects consider not only adult stages – the focus of most work to date – but also the importance of genetic diversity in early life history stages.

Network theory in ecology has been central to understanding species co-occurrence patterns, specialization and community stability. However, network theory has traditionally focused on the ‘higher’ trophic level where exploitation of network ‘partners’ (i.e. individual interactions in response to resource availability) have remained underappreciated. In this study we tested how clumping and host availability influenced mistletoe–host interactions in a semi-arid woodland, central Australia. We used a hierarchical approach that evaluated individual interactions by modifying the traditional randomization technique to simulate clumping and host exploitation. Using published literature we then compared our results with mistletoes from other genera. We found that mistletoes clump on fewer trees than predicted, even though interaction strength was no different from random expectations, and we found no evidence that common trees were heavily infected as predicted by the host availability hypothesis. The rate of host exploitation (measured as the proportion of trees infected) in semi-arid Australia is similar to that for mistletoe genera in other parts of the world. We hypothesize that specific host trees act as a focal point for infection that facilitates the spread and overall population size of mistletoes. Overall our results indicate that resources, such as the number of trees in a mistletoe network, are less important than clumping of individual plants. We suggest that exploitation of available resources may play a similar role in other networks that extend beyond antagonistic relationships such as parasite or herbivore interactions.

Using a 30 day time series of aphid Aphis helianthi and coccinellid counts on 107 mapped racemes of Yucca glauca, we demonstrate progressive, predation-induced self-organization of aphid colonies on individual racemes into extremely low and extremely high population sizes. This was driven by a two-attractor structure of density dependence that developed only in the presence of coccinellid predators. Foraging movements of the coccinellids among plants produced a power law relationship (average power = 0.142) between aphid and coccinellid numbers. This resulted in increased predation pressure on mid-size colonies and decreased predation pressure on small and large populations. A field-parameterized mathematical model predicts a two-attractor structure in broad agreement with our observations. The overall system was integrated by the influence of the largest aphid populations, which determined the total number of coccinellids present, and thus the predation pressure throughout the system. Our study provides clear evidence of predator-driven self-organization of prey populations in a patchy environment.

Pollination systems are recognized as critical for the maintenance of biodiversity in terrestrial ecosystems. Therefore, the understanding of mechanisms that promote the integrity of those mutualistic assemblages is an important issue for the conservation of biodiversity and ecosystem function. In this study we present a new population dynamics model for plant–pollinator interactions that is based on the consumer–resource approach and incorporates a few essential features of pollination ecology. The model was used to project the temporal dynamics of three empirical pollination network, in order to analyze how adaptive foraging of pollinators (AF) shapes the outcome of community dynamics in terms of biodiversity and network robustness to species loss. We found that the incorporation of AF into the dynamics of the pollination networks increased the persistence and diversity of its constituent species, and reduced secondary extinctions of both plants and animals. These findings were best explained by the following underlying processes: 1) AF increased the amount of floral resources extracted by specialist pollinators, and 2) AF raised the visitation rates received by specialist plants. We propose that the main mechanism by which AF enhanced those processes is (trophic) niche partitioning among animals, which in turn generates (pollen vector) niche partitioning among plants. Our results suggest that pollination networks can maintain their stability and diversity by the adaptive foraging of generalist pollinators.

The number of animals in a population is conventionally estimated by capture–recapture without modelling the spatial relationships between animals and detectors. Problems arise with non-spatial estimators when individuals differ in their exposure to traps or the target population is poorly defined. Spatially explicit capture–recapture (SECR) methods devised recently to estimate population density largely avoid these problems. Some applications require estimates of population size rather than density, and population size in a defined area may be obtained as a derived parameter from SECR models. While this use of SECR has potential benefits over conventional capture–recapture, including reduced bias, it is unfamiliar to field biologists and no study has examined the precision and robustness of the estimates. We used simulation to compare the performance of SECR and conventional estimators of population size with respect to bias and confidence interval coverage for several spatial scenarios. Three possible estimators for the sampling variance of realised population size all performed well. The precision of SECR estimates was nearly the same as that of the null-model conventional population estimator. SECR estimates of population size were nearly unbiased (relative bias 0–10%) in all scenarios, including surveys in randomly generated patchy landscapes. Confidence interval coverage was near the nominal level. We used SECR to estimate the population of a species of skink Oligosoma infrapunctatum from pitfall trapping. The estimated number in the area bounded by the outermost traps differed little between a homogeneous density model and models with a quadratic trend in density or a habitat effect on density, despite evidence that the latter models fitted better. Extrapolation of trend models to a larger plot may be misleading. To avoid extrapolation, a large region of interest should be sampled throughout, either with one continuous trapping grid or with clusters of traps dispersed widely according to a probability-based and spatially representative sampling design.

Two explanations exist for the evolutionary origin of grouping in primary consumers: reduction of individual predation risk and resource-mediated aggregation. While several studies have assessed relationships between aggregation and predation risk, few studies have examined the circumstances under which resource-mediated aggregation can lead to stable group formation. Using a model, we examined if forage preference alone can generate stable aggregation, and what were the circumstances of its emergence and stability. The model was a spatially explicit grazing model using empirically derived parameters to simulate large ruminant foraging in a meadow. Simulation results indicated that aggregation can spontaneously arise if grazers exhibit preference for forage of higher nutritional quality, usually associated with intermediate stages of forage growth. In this case, foragers could establish and maintain ‘islands’ of high quality forage as a result of revisiting continuous paths of previously grazed patches. However, aggregation was an intermittent phenomenon and occurred only within a narrow range of parameters. If grazer density was low compared to the amount of forage, the grazers’ foraging paths intersected too rarely to form contiguous islands of high forage quality; if their density was too high, the entire available area was uniformly utilized and foraging movements resembled unbounded random walks. We conclude that it is difficult to conceive of the evolution of grouping without the involvement of predators, since the relationship between grazer and forage abundance is ultimately co-regulated by predator abundance, and because in modern grazers, predator avoidance and foraging behavior seem to be functionally inseparable. Future research should consider the reinforcing effects of predator avoidance as well as foraging behavior on consumer aggregation.

Recent studies have described the architecture of plant–animal mutualistic networks, but little is known on how such networks disassemble as a consequence of global change. This is a relevant question because 1) species interactions seem to be very susceptible to habitat loss, and 2) the loss of a critical fraction of interactions can abruptly change the topology of the entire network with potential consequences for its functioning. Here we develop a spatially explicit metacommunity model based on the structure of 30 real mutualistic networks. We find that there is a critical value of habitat destruction beyond which interactions are lost very fast. Second, there is a homogeneous distribution of the number of interactions per patch when the habitat is pristine, while this becomes very skewed at the brink of extinction. This increase in skewness is discussed in the context of potential indicators of network collapse.

Fruit–frugivore interactions are crucial for the dynamics and regeneration of most forested ecosystems. Still, we lack an understanding of the potential variation in the sign and strength of such interactions in relation to variations in the spatial and temporal ecological context. Here, we evaluated spatial (three sites) and temporal (two fruiting seasons) local variation in the sign (seed predation versus dispersal) and strength (frequency and quantity) of the interactions among six frugivorous mammals and a community of Mediterranean fleshy-fruited shrubs. We examined mammal faecal samples and quantified frequency of seed occurrence, number of seeds per faecal sample, seed species diversity and quality of seed treatment (i.e. percentage of undamaged seeds). The frequency of seed occurrence and number of seeds per faecal sample strongly varied among dispersers, sites, seasons and fruit species. For instance, fox Vulpes vulpes faeces showed between 6 and 40 times more seeds than wild boar Sus scrofa faeces in seasons or sites in which Rubus and Juniperus seeds were dominant. However, in seasons or sites dominated by Corema seeds, wild boar faeces contained up to seven times more seeds than fox faeces. Mammalian carnivores (fox and badger, Meles meles) treated seeds gently, acting mostly as dispersers, whereas deer (Cervus elaphus and Dama dama) acted mainly as seed predators. Interestingly, rabbit Oryctolagus cuniculus acted as either mostly seed disperser or seed predator depending on the plant species. Our results indicated that the sign of fruit–frugivore interactions depended mainly on the identity of the partners. For a particular fruit–frugivore pair, however, our surrogate of interaction strength largely varied with the spatio-temporal context (year and habitat), leading to a low specificity across the seed–frugivore network. The high spatio-temporal variability of seed dispersal (in quantity, quality and seed diversity) by different frugivores would confer resilience against unpredictable environmental conditions, such as those typical of Mediterranean ecosystems.

Community assembly is a dynamic progression that reflects the interaction of several processes functioning at multiple scales. Understanding how these processes work in communities at different successional stages is important for identifying when regional or local processes are more important for community assembly, and for developing effective preservation and restoration strategies. We examined community assembly using a chronosequence of sub-alpine meadows in Qinghai-Tibetan Plateau that range from ‘natural’ (never farmed), to those that have been protected from agricultural exploitation for 1 to 10 years. We tested for shifts in species and traits among meadows and also for changes in environmental and spatial correlates of species distributions within meadows. We found that species richness increased and species composition returned to natural conditions within ten years of protection. These changes coincided with shifts in species traits; abundant species had high seed mass and specific leaf area in late-successional meadows, whereas the opposite occurred in early-successional meadows. Despite these shifts among meadows of different ages, spatial distributions of species within meadows did not change – when associated with abiotic variables, these spatial patterns reflected changes in soil pH and nitrogen. There was also no consistent change in the relative importance of environmental and spatial correlates of species distributions within meadows. These trends indicate that local processes of community assembly are similar within meadows even when species in those meadows differ. We conclude that successional change is a large-scale process that alters the species pool and resulting suite of traits that are present within meadows. As a result, regional planning that incorporates successional age should be the focus for the conservation of diversity in this area. In contrast, local processes work within the constraints of the species pool set by successional age, producing consistent patterns within meadows of different ages.