Although phenological matching between two and three trophic interactions has received some attention, it has largely been disregarded in explaining the lack of strong cascade dynamics in terrestrial systems. We studied the response of the specialist seed predator, Paroxyna plantaginis (Tephritidae) and associated generalist parasitoids (Chalcidoidea) to controlled fertilisation of individuals of naturally growing Tripolium vulgare (Asteraceae) on four island populations (Skeppsvik Archipelago, Sweden). We consistently found evidence of nutrient limitation: fertilised plants increased their biomass, produced more capitula (the oviposition units for tephritid flies), were more at risk of attack by the tephritids, and puparia were heavier in fertilised plants. During some parts of the season tephritids became more heavily parasitized, supporting the presence of cascade dynamics, however net parasitism over season decreased in response to nutrient addition. We found no evidence that capitulum size complicated parasitoid access to the tephritids, however the extended bud production prolonged the flowering season. Thus, tephritids utilized the surplus production of capitula throughout the entire season, while parasitoids did not expand their oviposition time window accordingly. Implications for top down regulation and cascade dynamics in the system are discussed.
This study evaluates how preference by a generalist slug herbivore Arion subfuscus changes inversely with seedling size across three levels of fertilization for three full‐sib families of willow seedlings. We analyzed seedlings for condensed tannin and protein concentration, and related these data to changes in palatability. In preference tests over time, leaf discs from more fertilized seedlings experienced an extended window of vulnerability compared to discs from less fertilized seedlings, which were also more tannin‐rich. In a whole seedling selection study, slugs readily attacked smaller seedlings (<5 cm) but rarely attacked taller seedlings (>10 cm). However, a general difference in risk of damage close to 50% existed when comparing shorter and taller individuals within each family and level of fertilizer. The decrease in palatability with height of the seedlings was positively correlated with an increase in condensed tannin concentration. We found no effect of seedling size on protein concentration. Akaiki index criterion model comparisons suggested that only main effects were important for explaining seedling choice by slugs as well as the ratio between proteins and condensed tannins. Seedling size, had the largest effect, followed by fertilizer level and family. Surprisingly, seedling size and fertilizer treatment had opposite effects on palatability to slugs. Size decreased probability of damage, whereas fertilization extended the window of susceptibility. Because the seedlings were even‐aged, differences in size are interpreted as differences in growth rate or vigor. The positive phenotypic correlation found between size and tannin production in the less preferred willow seedlings confirms that several plant defense traits may be selected for simultaneously, because fast growth may allow an early development of plant defenses. We discuss these results in the light of plant‐defense theories that predict a negative correlation between the allocation to growth and the production of secondary defense compounds.
Herbivory is one of the key drivers shaping plant community dynamics. Herbivores can strongly influence plant productivity directly through defoliation and the return of nutrients in the form of dung and urine, but also indirectly by reducing the abundance of neighbouring plants and inducing changes in soil processes. However, the relative importance of these processes is poorly understood. We, therefore, established a common garden experiment to study plant responses to defoliation, dung addition, moss cover, and the soil legacy of reindeer grazing. We used an arctic tundra grazed by reindeer as our study system, and Festuca ovina, a common grazing-tolerant grass species as the model species. The soil legacy of reindeer grazing had the strongest effect on plants, and resulted in higher growth in soils originating from previously heavily-grazed sites. Defoliation also had a strong effect and reduced shoot and root growth and nutrient uptake. Plants did not fully compensate for the tissue lost due to defoliation, even when nutrient availability was high. In contrast, defoliation enhanced plant nitrogen concentrations. Dung addition increased plant production, nitrogen concentrations and nutrient uptake, although the effect was fairly small. Mosses also had a positive effect on aboveground plant production as long as the plants were not defoliated. The presence of a thick moss layer reduced plant growth following defoliation. This study demonstrates that grasses, even though they suffer from defoliation, can tolerate high densities of herbivores when all aspects of herbivores on ecosystems are taken into account. Our results further show that the positive effect of herbivores on plant growth via changes in soil properties is essential for plants to cope with a high grazing pressure. The strong effect of the soil legacy of reindeer grazing reveals that herbivores can have long-lasting effects on plant productivity and ecosystem functioning after grazing has ceased.
Predator-induced morphological defences (PIMDs) are ubiquitous. Many PIMDs may be mediated by prey behaviour rather than directly cued by predators. A survey of 92 studies indicated 40 that quantified prey behaviour, all of which document positive associations between defence production and activity reduction. Thus, PIMDs are associated with changes in prey activity, which could have caused the morphological change. We propose two possible mechanisms: 1) decreased activity reduces feeding rate, resulting in lower growth and morphological change; and 2) activity reduction conserves energy, which is reallocated for growth, subsequently changing morphology. Resource availability also causes similar morphological change to predator presence, suggesting confounding effects of resources and predators with current methodology. Future studies should estimate food ingestion, assimilation efficiency, and growth rate in the presence and absence of predators, crossing predator presence with resource levels. Not all PIMDs will be behaviourally-mediated, but consideration of causal linkages between prey behaviour and PIMDs is warranted.
Empirical studies of the interaction between the anther smut fungus Microbotryum violaceum and its host plant Lychnis alpina were combined with modelling approaches to investigate how variation in the spatial distribution of host populations influences disease dynamics and variation in resistance. Patterns of disease incidence and prevalence were surveyed in three contrasting systems of natural L. alpina populations where there is substantial variation in spatial structure, ranging from large continuous populations through to small isolated patches. Disease incidence (fraction of populations where disease was present) was highest in the continuous situation, and lowest in the most isolated populations. The reverse was true for prevalence (fraction of individuals diseased). To better understand the long-term ecological and evolutionary consequences of differences in among population spatial structure, we developed a two-dimensional spatially explicit simulation model in which host-population spacing was modelled by varying the percentage of sites suitable for the host. The general patterns of disease incidence and prevalence generated in the simulations corresponded well with the patterns observed in natural populations of L. alpina and M. violaceum; i.e. the fraction of sites with disease increased while the average disease prevalence in diseased populations decreased when host populations became more connected. One likely explanation for the differences in disease incidence and prevalence seen in natural populations is that the evolution of host resistance varies as a function of the degree of fragmentation. This is supported by simulation results that were qualitatively similar to the survey data when resistance was allowed to vary, but not when hosts were assumed to be uniformly susceptible. In the former, the frequency of resistance increased markedly as host populations became more connected.
Droop's model was originally designed to describe the growth of unicellular phytoplankton species in chemostats but it is now commonly used for a variety of organisms in models of trophic interactions, ecosystem functioning, and evolution. Despite its ubiquitous use, Droop's model is still limited by several simplifying assumptions. For example, the assumption of equal theoretical maximum growth rates for all nutrients is commonly used to describe growth limited by multiple nutrients. This assumption, however, is both biologically unrealistic and potentially misleading. We propose the alternative hypothesis of equal realized maximum growth rates for all nutrients. We support our hypothesis with empirical and theoretical arguments and discuss how it may improve our understanding of the biology of growth, while avoiding some of the pitfalls of the previous assumption.
Most animal pollination results from plant–insect interactions, but how we perceive these interactions may differ with the sampling method adopted. The two most common methods are observations of visits by pollinators to plants and observations of pollen loads carried by insects. Each method could favour the detection of different species and interactions, and pollen load observations typically reveal more interactions per individual insect than visit observations. Moreover, while observations concern plant and insect individuals, networks are frequently analysed at the level of species. Although networks constructed using visitation and pollen-load data have occasionally been compared in relatively specialised, bee-dominated systems, it is not known how sampling methodology will affect our perception of how species (and individuals within species) interact in a more generalist system. Here we use a Diptera-dominated high-Arctic plant–insect community to explore how sampling approach shapes several measures of species' interactions (focusing on specialisation), and what we can learn about how the interactions of individuals relate to those of species. We found that species degrees, interaction strengths, and species motif roles were significantly correlated across the two method-specific versions of the network. However, absolute differences in degrees and motif roles were greater than could be explained by the greater number of interactions per individual provided by the pollen-load data. Thus, despite the correlations between species roles in networks built using visitation and pollen-load data, we infer that these two perspectives yield fundamentally different summaries of the ways species fit into their communities. Further, individuals' roles generally predicted the species' overall role, but high variability among individuals means that species' roles cannot be used to predict those of particular individuals. These findings emphasize the importance of adopting a dual perspective on bipartite networks, as based on the different information inherent in insect visits and pollen loads.
There has been growing recent use of elevational gradients as tools for assessing effects of temperature changes on vegetation properties, because these gradients enable temperature effects to be considered over larger spatial and temporal scales than is possible through conventional experiments. While many studies have explored the direct effects of temperature, the indirect effects of temperature through its long-term influence on soil abiotic or biotic properties remain essentially unexplored. We performed two climate chamber experiments using soils from a subarctic elevational gradient in Abisko, Sweden to investigate the direct effects of temperature, and indirect effects of temperature via soil legacies, on growth of two grass species. The soils were collected from each of two vegetation types (heath, dominated by dwarf shrubs, and meadow, dominated by graminoids and herbs) at each of three elevations. We found that plants responded to both the direct effect of temperature and its indirect effect via soil legacies, and that direct and indirect effects were largely decoupled. Vegetation type was a major determinant of plant responses to both the direct and indirect effects of temperature; responses to soils from increasing elevation were stronger and showed a more linear decline for meadow than for heath soils. The influence of soil biota on plant growth was independent of elevation, with a positive influence across all elevations regardless of soil origin for meadow soils but not for heath soils. Taken together, this means that responses of plant growth to soil legacy effects of temperature across the elevational gradient were driven primarily by soil abiotic, and not biotic, factors. These findings emphasize that vegetation type is a strong determinant of how temperature variation across elevational gradients impacts on plant growth, and highlight the need for considering both direct and indirect effects of temperature on plant responses to future climate change.
I experimentally evaluated the impact of habitat structural complexity on the interactions between a generalist predator and a benthic macroinvertebrate prey assemblage in a freshwater pond. Benthivorous perch (Perca fluviatilis) were stocked over a range of natural densities (no fish, low, and high densities) into enclosures with or without dense submerged vegetation. The biomass of macroinvertebrate prey decreased over time in the presence of perch and was always higher in enclosures with vegetation present than in enclosures lacking vegetation. The increase in mass of perch was positively related to the abundance of macroinvertebrate prey and negatively related to perch density and the density of vegetation. In the treatments lacking vegetation, the proportion of zooplankton in the diet of perch increased, and the growth rate of perch decreased over time. In the vegetation treatments, the proportion of zooplankton in the diet was low throughout the experiment and the growth rate of perch was constant over time. As a consequence, initial increase in mass was considerably higher in the treatments lacking vegetation than in the vegetation treatments, whereas no such pattern was observed in the second half of the experiment. In the absence of vegetation, perch are apparently able to forage efficiently, but this may reduce the availability of macroinvertebrate prey to the extent that perch are forced to include less profitable zooplankton prey into their diet. In vegetated habitats, the foraging efficiency of perch is reduced, which possibly prevents over-exploitation of macroinvertebrate prey and consequently may allow for a moderate, but relatively constant, consumption of macroinvertebrates by perch. The density-dependence of growth rates in both vegetated and unvegetated habitats can only partly be explained by resource competition, which suggests the presence of an additional mechanism of density-dependence. In natural lake communities, efficient predation from benthivorous fish should keep the biomass of macroinvertebrate prey in structurally simple habitats below the high levels initially present in my experiment. In these communities, submerged vegetation may be an equally profitable habitat for juvenile perch as are open areas. Through its effects on the feeding efficiencies of juvenile perch and other benthivorous fish, submerged vegetation may affect individual growth rates and the size structure of perch populations, which may contribute to explain differences in fish community structure among lakes differing in submerged vegetation cover.
Omnivory (the consumption of resources from more than one trophic level) is widespread in nature and has the potential to produce a richness of indirect effects. Nevertheless, its effects on population dynamics have received very little attention. In its simplest case, omnivory involves a top consumer, an intermediate consumer, and a resource that is common to both consumers. Simple models predict that the intermediate consumer can only coexist with the top consumer if the former is more efficient in exploiting the common resource, which would imply a net positive effect of the top consumer on the equilibrium density of resources (compared to the situation where only the intermediate consumer is present). Among 22 experimental manipulations of omnivorous top consumers I found only 2 studies in which top consumers had significant positive effects on resources. This discrepancy between experimental results and model predictions is, at least partly, related to deviations of the experimental systems from model assumptions. However, considerations of relative body sizes of intermediate and top consumers suggest, that top consumers having negative net effects on the basic resource should be common in nature. I argue that in systems where intermediate consumers and basic resources are relatively similar in size, but both are much smaller than omnivorous top consumers (e.g. vertebrate omnivores feeding on benthos, soil invertebrates, terrestrial insects etc.), the direct negative effect of top consumers on basic resources should not be outweighed by indirect positive effects, and that other mechanisms (e.g. prey refuges) must be invoked to explain the persistence of intermediate consumers in many natural systems. I further argue that a better knowledge of the population dynamical consequences of omnivory and the role of relative consumer sizes is necessary to improve our understanding of the-trophic dynamics of different kinds of communities.
Metabolic theory proposes that individual growth is governed through the mass- and temperature-dependence of metabolism, and ecological stoichiometry posits that growth is maximized at consumer-specific optima of resource elemental composition. A given consumer's optimum, the threshold elemental ratio (TER), is proportional to the ratio of its maximum elemental gross growth efficiencies (GGEs). GGE is defined by the ratio of metabolism-dependent processes such that GGEs should be independent of body mass and temperature. Understanding the metabolic-dependencies of GGEs and TERs may open the path towards a theoretical framework integrating the flow of energy and chemical elements through ecosystems. However, the mass and temperature scaling of GGEs and TERs have not been broadly evaluated. Here, we use data from 95 published studies to evaluate these metabolic-dependencies for C, N and P from unicells to vertebrates. We show that maximum GGEs commonly decline as power functions of asymptotic body mass and exponential functions of temperature. The rates of change in maximum GGEs with mass and temperature are relatively slow, however, suggesting that metabolism may not causally influence maximum GGEs. We additionally derived the theoretical expectation that the TER for C:P should not vary with body mass and this was supported empirically. A strong linear relationship between carbon and nitrogen GGEs further suggests that variation in the TER for C:N should be due to variation in consumer C:N. In general we show that GGEs may scale with metabolic rate, but it is unclear if there is a causal link between metabolism and GGEs. Further integrating stoichiometry and metabolism will provide better understanding of the processes governing the flow of energy and elements from organisms to ecosystems.
During 1991–95, mammalian predators (weasel, Mustela nivalis, stoat, M. erminea, mink, M. vison, and red fox, Vulpes vulpes) were excluded in late summers from a 2 ha piece of a north Norwegian mountain slope. The exclosure extended from an outpost of luxuriant sub-arctic birch forest to a typical arctic–alpine habitat complex, including productive willow scrublands, tundra heaths, dry ridges and snow-beds. The exclosure thus encompassed the entire range of habitat conditions encountered in a typical north Fennoscandian mountain and tundra landscapes. During 1991–95, the exclosure was predator-proof from late July to late September. In wintertime and in early summer, the exclosure was accessible to mammalian predators. Vole dynamics in the short-term exclosure were compared to dynamics in five reference areas with similar habitat conditions.
In 1991, when vole densities were rising in the area, neither collective vole densities nor densities of individual vole species differed significantly between the exclosure and the replicated controls. Spring densities of voles were never significantly different between the exclosure and the controls. With respect to autumnal densities of voles, however, the exclosure was a statistical outlier in the peak year 1992 and throughout the gradual decline phase of 1993–95. In the peak year, the difference in collective vole densities was modest (30%), but increased to two-fold during the first two decline years and was almost four-fold in the crash year of 1995. The strongest response was displayed by field voles (Microtus agrestis), hypothesized to be the pivotal prey species of weasels, especially by females and young individuals, i.e. by the functional categories especially sensitive to mammalian predation. These results are consistent with the hypothesis that predation plays a pivotal role for the regulation of herbivorous mammals in relatively productive arctic–alpine habitats.
Over a period of seven seasons (1995-2001 inclusively) four distinct epidemics of the rust pathogen Melampsoridium betulinum were recorded in stands of Betula pubescens. During those epidemics, host plants incurred varying levels of disease severity. Some individuals suffered high levels on all occasions; some low; while yet others were either intermediately affected or showed variable disease severity from season to season. Tests of pathotypic differences among 33 isolates of M. betulinum collected from six sites found a broad range of pathotypes ranging from highly virulent to highly avirulent. Similarly a random sample of 40 B. pubescens lines from one site showed a wide range of resistance phenotypes, although individuals that were either susceptible to all but one pathotype or resistant to all pathotypes were commonest. A strong relationship existed between the cumulative level of disease incurred by B. pubescens individuals in the field and the mean susceptibility of each host line as determined by their reaction to infection by each of the 33 different isolates of M. betulinum individually. Resistance in this B. pubescens population to M. betulinum is postulated to be based on a mixture of quantitative and qualitative traits, selection for which has resulted from an interplay of life history attributes of both host and pathogen.
Human impacts such as eutrophication, overexploitation and climate change currently threaten future global food and drinking water supplies. Consequently, it is important that we understand how anthropogenic resource (bottom–up) and consumer (top–down) manipulations affect aquatic food web structure and production. Future climate changes are predicted to increase the inputs of terrestrial dissolved organic carbon to lakes. These carbon subsidies can either increase or decrease total basal production in aquatic food webs, depending on bacterial competition with phytoplankton for nutrients. This study examines the effects of carbon subsidies (bottom–up) on a pelagic community exposed to different levels of top–down predation. We conducted a large scale mesocosm experiment in an oligotrophic clear water lake in northern Sweden, using a natural plankton community exposed to three levels of glucose addition (0, 420 and 2100 mg C l–1 total added glucose) and three levels of young-of-the-year perch Perca fluviatilis density (0, 0.56 and 2 individuals m–3). Bacterioplankton production doubled with glucose addition, but phytoplankton production was unaffected, in contrast to previous studies that have manipulated carbon, nutrients or light simultaneously. This suggests that carbon addition alone is not sufficient to reduce autotrophic production, at least in an oligotrophic lake dominated by mixotrophic phytoplankton. Larval perch grazing did not produce a classical trophic cascade, but substantially altered the species composition of crustacean zooplankton and ciliate trophic levels. Glucose addition increased the biomass of rotifers, thus potentially increasing energy transfer through the heterotrophic pathway, but only when fish were absent. This study illustrates that changes in community structure due to selective feeding by top-predators can determine the influence of bottom–up carbon subsidies.
How are resource consumption and growth rates of litter-consuming detritivores affected by imbalances between consumer and litter C:N:P ratios? To address this question, we offered leaf litter as food to three aquatic detritivore species, which represent a gradient of increasing body N: P ratios: a crustacean, a caddisfly and a stonefly. The detritivores were placed in microcosms and submerged in a natural stream. Four contrasting leaf species were offered, both singly and in two-species mixtures, to obtain different levels of stoichiometric imbalance between the resources and their consumers. The results suggest that detritivore growth was constrained by N rather than C or P, even though 1) the N: P ratios of the consumers' body tissue was relatively low and 2) microbial leaf conditioning during the experiment reduced the N:P imbalance between detritivores and leaf litter. This surprisingly consistent N limitation may be a consequence of cumulative N-demand arising from the production of N-rich chitin in the exoskeletons of all three consumer species, which is lost during regular moults, in addition to N-demand for silk production by the caddisfly. These N requirements are not commonly quantified in stoichiometric analyses of arthropod consumers. There was no evidence for compensatory feeding, but when offered mixed-species litter varying in C:N:P ratios, detritivores consumed more of the litter species showing the highest N:P and lowest C:N ratio, accelerating the mass loss of the preferred leaf species in the litter mixture. These results show that imbalances in consumer-resource stoichiometry can have contrasting effects on coupled processes, highlighting a challenge in developing a mechanistic understanding of the role of stoichiometry in regulating ecosystem processes such as leaf litter decomposition.
Several mechanisms for biological invasions have been proposed, yet to date there is no common framework that can broadly explain patterns of invasion success among ecosystems with different resource availabilities. Ecological stoichiometry (ES) is the study of the balance of energy and elements in ecological interactions. This framework uses a multi-nutrient approach to mass-balance models, linking the biochemical composition of organisms to their growth and reproduction, which consequently influences ecosystem structure and functioning. We proposed a conceptual model that integrates hypotheses of biological invasions within a framework structured by fundamental principles of ES. We then performed meta-analyses to compare the growth and production performances of native and invasive organisms under low- and high-nutrient conditions in terrestrial and aquatic ecosystems. Growth and production rates of invasive organisms (plants and invertebrates) under both low- and high-nutrient availability were generally larger than those of natives. Nevertheless, native plants outperformed invasives in aquatic ecosystems under low-nutrient conditions. We suggest several distinct stoichiometry-based mechanisms to explain invasion success in low- versus high-nutrient conditions; low-nutrient conditions: higher resource-use efficiency (RUE; C:nutrient ratios), threshold elemental ratios (TERs), and trait plasticity (e.g. ability of an organism to change its nutrient requirements in response to varying nutrient environmental supply); high-nutrient conditions: higher growth rates and reproductive output related to lower tissue C:nutrient ratios, and increased trait plasticity. Interactions of mechanisms may also yield synergistic effects, whereby nutrient enrichment and enemy release have a disproportionate effect on invasion success. To that end, ES provides a framework that can help explain how chemical elements and energy constrain key physiological and ecological processes, which can ultimately determine the success of invasive organisms.
Global warming has created a need for studies along climatic gradients to assess the effects of temperature on ecological processes. Altitudinal and latitudinal gradients are often used as such, usually in combination with air temperature data from the closest weather station recorded at 1.52 m above the ground. However, many ecological processes occur in, at, or right above the soil surface. To evaluate how representative the commonly used weather station data are for the microclimate relevant for soil surface biota, we compared weather station temperatures for an altitudinal (500900 m a.s.l.) and a latitudinal gradient (4968 degrees N) with data obtained by temperature sensors placed right below the soil surface at five sites along these gradients. The mean annual temperatures obtained from weather stations and adjusted using a lapse rate of -5.5 degrees C km-1 were between 3.8 degrees C lower and 1.6 degrees C higher than those recorded by the temperature sensors at the soil surface, depending on the position along the gradients. The monthly mean temperatures were up to 10 degrees C warmer or 5 degrees C colder at the soil surface. The within-site variation in accumulated temperature was as high as would be expected from a 300 m change in altitude or from a 4 degrees change in latitude or a climate change scenario corresponding to warming of 1.63.8 degrees C. Thus, these differences introduced by the decoupling are significant from a climate change perspective, and the results demonstrate the need for incorporating microclimatic variation when conducting studies along altitudinal or latitudinal gradients. We emphasize the need for using relevant temperature data in climate impact studies and further call for more studies describing the soil surface microclimate, which is crucial for much of the biota.
Some communities are susceptible to invasions and some are not. Why? Elton suggested in 1958 that the ability of the community to withstand invading species - its biotic resistance - depends on the number of resident species. Later contributors have emphasized the habitat's ability to support species, as well as the contribution of individual species to the resistance. In this study we use information from 184 introductions of Arctic char into Swedish lakes to study both abiotic and biotic aspects of the resident community's ability to resist introductions. We find that the best model included the proportion of forest cover and the proportion of agricultural land cover in the watershed in combination with the presence versus absence of northern pike. Thus, the most important biotic factor to explain the outcome of introductions of Arctic char is the presence of northern pike, a large piscivore. This means that one single species explains the outcome of the introductions better than does the species richness or the saturation level of the community.
Plant phenotypes often differ in their resistance to natural enemies, but the mechanism for this has seldom been identified. The aim of this study was to determine if the spatial patterns of phenotype use of a highly specialized insect herbivore (the galling sawfly Pontania triandrae) in a natural willow population can be related to phenotypic variation in plant secondary chemistry. Furthermore, we tested if traits that confer resistance to one type of natural enemy, i.e. the galling sawfly, also confer resistance to others, in our case a leaf beetle Gonioctena linnaeana and the rust fungus Melampsora amygdalinae. We identified 18 phenotypes with high and 18 phenotypes with low gall density in our field population and determined gall densities, the degree of leaf damage and rust infection on each phenotype and collected leaves for chemical analyses. The concentration of phenolics was higher in phenotypes with high density of galls suggesting that this galling sawfly may use phenolics as oviposition cues. Rust infection showed the opposite pattern, with lower levels on clones with high concentration of phenolics, while leaf damage by G. linnaeana did not differ between clone types. This indicates that these important natural enemies may assert divergent selection on willow phenotypes and that this might provide a mechanism for maintaining phenotypic variation within willow populations.
Variation in growth rates among individuals leading to the formation of broad size distributions is commonly observed in animal cohorts. Here we use laboratory derived size-scaling relationships to identify mechanisms driving changes in size distribution patterns within cohorts during early ontogeny. We introduced young-of-the-year perch (Perca fluviatilis) cohorts with different variation in body size distributions in pond enclosures. We kept the exploitative competitive environment constant by adjusting the number of introduced fish such that metabolic requirements were constant between different treatments. Based on modelling results we theoretically derived relative growth rates of differently sized fish when only taken exploitative competitive interactions into account. In agreement with predictions we found that initial variation in body size was negatively correlated with subsequent changes in body size variation in the pond experiment. Corresponding results were obtained in a field study covering 13 studied young-of-the-year perch cohorts in a small lake. Besides having a lower maximum growth capacity, initially large fish also suffered more from resource limitation in our experiment. The results suggest that exploitation competition is a major factor behind growth patterns in young fish cohorts, generally leading to size convergence. To explain the commonly observed pattern of size divergence in animal cohorts, including fish, we suggest that differential timing of diet shifts or mechanisms not related to exploitative interactions must be taken into account. For diet shifts to lead to size divergence we suggest that individuals with an initial size advantage need access to an exclusive prey which has a high growth potential. This, in turn, allows initially larger individuals to surf on a wave of growing prey while individuals only capable to feed on a depressed initial resource experience low growth rates.
Th e degree to which growth in early life stages of animals is regulated via density-dependent feedbacks through preyresources is much debated. Here we have studied the infl uence of size- and density-dependent mechanisms as well as sizeselectivepredation pressure by cannibalistic perch Perca fl uviatilis on growth patterns of young-of-the-year (YOY) perchcovering several lakes and years. We found no infl uence of initial size or temperature on early body size development ofperch. In contrast, there was a negative relationship between reproductive output and the length of YOY perch at fi ve weeksof age. However, rather than an eff ect of density-dependent growth mediated via depressed resources the relationship wasdriven by positive size-selective cannibalism removing large individuals. Hence, given a positive correlation between thedensity of victims and predation pressure by cannibals, size-dependent interactions between cannibals and their victimsmay wrongly be interpreted as patterns of density-dependent growth in the victim cohort. Overall, our results support theview that density-dependent resource-limitation in early life stages is rare. Still, patterns of density-dependent growth mayemerge, but from variation in size-selective predation pressure rather than density as such. Th is illustrates the importanceof taking overall population demography and predatory interactions into account when studying growth patterns amongrecruiting individuals.
Cannibalistic interactions generally depend on the size relationship between cannibals and victims. In many populations, alarge enough size variation to allow for cannibalism may not only develop among age-cohorts but also within cohorts. Westudied the implications of variation in hatching period length and initial cohort size for the emergence of cannibalism andbimodal size distributions within animal cohorts using a physiologically structured population model. We found that thedevelopment of size bimodality was critically dependent on hatching period length, victim density and the presence of afeedback via shared resources. Cannibals only gained enough energy from cannibalism to accelerate in growth when victimdensity was high relative to cannibal density at the onset of cannibalism. Furthermore, we found that the opportunity forearly hatchers to initially feed on an unexploited resource increases the likelihood both for cannibalism to occur and sizebimodality to develop. Once cannibals accelerated in growth relative to victims size bimodality, reduced victim numbersand relaxed resource competition resulted. Th us, in addition to that cannibals profi ted from cannibalism through energyextraction, their potential victims also benefi ted as the resource recovered due to cannibal thinning. To ensure recruitmentsuccess, it can be critical that a few individuals can accelerate in growth and reach a size large enough to escape sizedependentpredation and winter starvation. Hence, within-cohort cannibalism may be a potentially important mechanismto explain recruitment variation especially for cannibalistic species in temperate climates with strong seasonality. However,the scope for size bimodality to develop as a result of cannibalism may be limited by low victim densities and size andfood-dependent growth rates.
We set up two alternative hypotheses on how environmental variables could foster nestedness; one of “nested habitats” and another of “nested habitat quality”. The former hypothesis refers to situations where the nestedness of species depends on a nestedness of discrete habitats. The latter considers situations where all species in an assemblage increase in abundance along the same environmental gradient, but differ in specialisation or tolerance. We tested whether litter-dwelling land snails (terrestrial gastropods) in boreal riparian forest exhibited a nested community structure, whether such a pattern was related to differences in environmental variables among sites, and which of the two hypotheses that best could account for the found pattern. We sampled litter from 100 m2 plots in 29 mature riparian forest sites along small streams in the boreal zone of Sweden. The number of snail species varied between 3 and 14 per site. Ranking the species-by-site matrix by PCA scores of the first ordination axis revealed a similarly significant nested pattern as when the matrix was sorted by number of species, showing that the species composition in this meta-community can be properly described as nested. Several environmental variables, most notably pH index, were correlated with the first PCA axis. All but two species had positive eigenvectors in the PCA ordination and the abundance increased considerably along the gradient for most of the species implying that the hypothesis of “nested habitats” was rejected in favour of the “nested habitat quality” hypothesis. Analyses of nestedness have seldom been performed on equal sized plots, and our study shows the importance of understanding that variation in environmental variables among sites can result in nested communities. The conservation implications are different depending on which of our two hypotheses is supported; a conservation focus on species “hotspots” is more appropriate if the communities are nested because of “nested habitat quality”.
Recent studies have shown that biotic interactions can shape species’ distributions, but empirical data on multiple biotic interactions are scarce. Therefore, we examined effects of plant-plant and plant-herbivore interactions on plant survival, growth and reproduction at different altitudes. For these purposes we conducted a factorial neighbor removal and large herbivore exclusion experiment with six transplant species (three tall forbs with their main distribution at low altitudes and three small forbs with their main distribution at high altitudes) on Låktačohkka Mountain, northern Sweden, replicated at two altitudes (ca. 600 and 900 m a.s.l.) and consequently a 2.1 °C difference in summer air temperatures. Overall transplant survival was 93%. Two out of three tall forbs grew better at low than at high altitudes, while no significant differences in growth between altitudes were found for any of the three small forbs. Since the main difference in abiotic conditions between the altitudes was most likely in temperature (as the sites were topographically and edaphically matched as closely as possible), this result indicates that climatic warming could induce upward migration of tall low-altitude forbs. Negative plant-plant interactions prevailed at both altitudes, and we found indications that competition may set the lower altitudinal limits of some small tundra forbs. Thus, increased competition in response to climate warming may potentially shift the lower margins of high-altitude forbs’ distributions upward. Large mammalian grazers reduced the growth of tall forbs and enhanced the flowering of small forbs, and grazers could thus at least partly counteract the anticipated warming-induced distribution shifts.
This study investigates the allocation of allochthonous organic carbon (AlloOC) to pelagic respiration and biomass production in unproductive lakes. Metabolic process rates and stable isotopic composition (delta C-13) of crustacean zooplankton and respired CO2 were measured in the epilimnion of 13 forest lakes in northern Sweden. The delta C-13 of zooplankton was low (-31.2 to -38.0 parts per thousand) compared to that of respired CO2 (-28.4 to -30.6 parts per thousand), implying that the relative importance of AlloOC was lower for zooplankton (ca 40%) than for respiration (ca 80%). Combining delta C-13 and carbon flux data revealed that a large amount of metabolized AlloOC was lost in respiration, compared to the amount transferred to zooplankton (< 3%). Thus, despite large respiratory losses, AlloOC was still important for zooplankton growth, implying a high supply of AlloOC in comparison to phytoplankton generated organic carbon in the lakes.
We used stable carbon (δ13C) and nitrogen (δ15N) isotopes to assess the importance of benthic algae for the zooplankton individual growth in winter in a shallow, clear subarctic lake. The δ13C values of calanoid (Eudiaptomus graciloides) and cyclopoid (Cyclops scutifer) zooplankton in autumn suggest a food resource of pelagic origin during the ice-free period. The zooplankton δ13C values were high in spring compared to autumn. E. graciloides did not grow over winter and the change in δ13C was attributed to a decrease in lipid content during the winter. In contrast, the increase in δ13C values of C. scutifer over the winter was explained by their growth on organic carbon generated by benthic algae. The δ15N of the C. scutifer food resource during winter was low compared to δ15N of the benthic community, suggesting that organic matter generated by benthic algae was mainly channelled to zooplankton via 15N-depleted heterotrophic bacteria. The results demonstrate that benthic algae can sustain zooplankton metabolic demands and growth during long winters, which, in turn, may promote zooplankton growth on pelagic resources during the summer. Such multi-chain omnivory challenges the view of zooplankton as mainly dependent on internal primary production and stresses the importance of benthic resources for the productivity of plankton food webs in shallow lakes.
Despite long-standing research, the processes that drive community assembly remain poorly understood. We censused macroinvertebrate communities and measured flood disturbance in 17 Scandinavian mountain streams to assess the hypothesis that communities are shaped by stochastic processes under stable conditions, and increasingly by deterministic processes as disturbance becomes more severe. Each study stream was categorized as being stable (n=5), intermediate (n=7), or disturbed (n=5) depending on the severity of scouring floods. Following spring floods, the number of potential colonisers decreased with increasing disturbance, suggesting that disturbance filtered out species unable to cope with the stress involved. Communities at stable sites had the highest beta diversity, indicating that stochastic processes of community assembly were most important under the least disturbed conditions. In partial contrast with our predictions, the lowest beta diversity occurred between intermediate (not disturbed) sites, suggesting that increasing disturbance first enhances determinism but then rekindles stochasticity at severity levels beyond intermediate. Macroinvertebrate communities were shaped by deterministic processes, which recruit potential regional colonists depending on niche differences and disturbance conditions and by stochastic processes, which distribute the selected species randomly among individual localities. Although often considered opposing, stochastic and deterministic processes interact hierarchically, with relative strength modified by disturbance.
Trypsin inhibitors have been suggested to constitute an inducible defense in the sedge Carex bigelowii, and some former studies suggest that this might be a cause for the cyclic population dynamics in many alpine and arctic small mammals, for example lemmings (Lemmus lemmus). We investigated this further by using a method of simulated grazing (clipping) at different intensities, in three different habitats with varying resource availability, with different harvest times (hours after clipping), and two different stages of ramets (reproductive/vegetative) in a study from the Swedish mountain range. Our results do not indicate that C. bigelowii has an inducible defense constituted by an increase in trypsin inhibitor activity (TIA), but rather that the amount of soluble plant proteins (SPP) is lowered in wounded plants. The responses were somewhat different in the three habitats, with ramets growing in the marsh showing the highest ratio of TIA to SPP, due to low amounts of SPP. We did not find any significant effects of harvest time, or of the stage of the ramet that could support the hypothesis of an inducible defense. To conclude, we could not find any evidence for an inducible defense consisting of trypsin inhibitors in Carex bigelowii ramets, but we did find variations in the amount of SPP that may have nutritional consequences for herbivores.
The impact of food web complexity in open and closed food webs was explored experimentally under controlled laboratory conditions. We used an aquatic model community consisting of two forms of primary producers, phytoplankton (Scenedesmus obtusiuscusculus) and periphyton (Nitzschia perminuta) and two types of consumers, Daphnia pulex feeding on phytoplankton and Chydorous sphaericus feeding on both periphyton and phytoplankton. Three different food webs all having the phytoplankton and periphyton, but having either one of the consumers or both were set up. These food webs were studied in an open and closed treatment. In the open treatment, phytoplankton was continuously flowing through the aquaria whereas in the closed system all the phytoplankton was delivered at the start of the experiment.
D. pulex had a positive effect on the density of C. sphaericus in both the open and closed treatments. In the open treatment C. sphaericus increased to very high numbers and was able to depress phytoplankton and thereby negatively affect D. pulex. Our study shows that the explicit handling of the population dynamics of both grazers allowed us to show how a compensatory increase in one primary producer due to increased grazing on another primary producer creates a negative feedback between consumers: C. sphaericus increased and negatively affected D. pulex.
Herbivores play a key role in the carbon (C) cycle of arctic ecosystems, but these effects are currently poorly represented within models predicting land-atmosphere interactions under future climate change. Although some studies have examined the influence of various individual species of herbivores on tundra C sequestration, few studies have directly compared the effects of different herbivore assemblages. We measured peak growing season instantaneous ecosystem carbon dioxide (CO2) exchange (photosynthesis, respiration and net ecosystem exchange) on replicated plots in arctic tundra which, for 14 years, have excluded different portions of the herbivore population (grazed controls, large mammals excluded, both small and large mammals excluded). Herbivory suppressed photosynthetic CO2 uptake, but caused little change in ecosystem respiration. Despite evidence that small mammals consume a greater portion of plant biomass in these ecosystems, the effect of excluding only large herbivores was indistinguishable from that of excluding both large and small mammals. The herbivory-induced decline in photosynthesis was not entirely attributable to a decline in leaf area but also likely reflects shifts in plant community composition and/or species physiology. One shrub species - Betula nana - accounted for only around 13% of total aboveground vascular plant biomass but played a central role in controlling ecosystem CO2 uptake and release, and was suppressed by herbivory. We conclude that herbivores can have large effects on ecosystem C cycling due to shifts in plant aboveground biomass and community composition. An improved understanding of the mechanisms underlying the distinct ecosystem impacts of different herbivore groups will help to more accurately predict the net impacts of diverse herbivore communities on arctic C fluxes.
Herbivores are generally faced with a plethora of resources which differ in quality. Therefore, they should be able to select foods which most closely match their metabolic needs. Here, we tested the hypothesis that copepods of the species Acartia tonsa select prey cells based on quality differences within prey species. We assessed age-specific variation in feeding behaviour and evaluated the potential consequences of such variation for nutrient cycles. Nauplii (young) stages characterized by a low nitrogen to phosphorus (N:P) ratio in their body tissue selected for phosphorus-rich food, while older copepodite stages with higher body N: P selected for nitrogen-rich food. Further, the analysis of a 35-year data set in the southern North Sea revealed a positive correlation between the abundance of nauplii and the ratio of dissolved inorganic N:P, thus suggesting that P-availability for primary producers declines with the population densities of nauplii. Our findings demonstrate that a combination of stage-specific selective feeding and body stoichiometry has the potential to affect cycling of limiting nutrients when consumer populations change in composition.
There has been a long‐standing debate on what creates stability in food webs. One major finding is that weak interactions can mute the destabilizing potential of strong interactions. Considering that stage structure is common in nature, that existing studies on stability that include population stage structure point in different directions, and the recent theoretical developments in the area of stage structure, there is a need to address the effects of population stage structure in this context. Using simple food web modules, with stage structure in an intermediate consumer, we here begin to theoretically investigate the effects of stage structure on food web stability. We found a general correspondence to previous results such that strong interactions had destabilizing effects and weak interactions that result in decreased energy flux had stabilizing effects. However, we also found a number of novel results connected to stage structure. Interestingly, weak interactions can be destabilizing when they excite other interactions. We also found that cohort cycles and predator–prey cycles did not respond in the same way to increasing interactions strength. We found that the combined effects of two predators feeding on the same prey can strongly destabilize a system. Consistent with previous studies, we also found that stage‐specific feeding can create a refuge effect that leads to a lack of strong destabilization at high interaction strength. Overall, stage structure had both stabilizing and destabilizing aspects. Some effects could be explained by our current understanding of energetic processes; others need additional consideration. Additional aspects such as shunting of energy between stages, control of biomass fluxes, and interactions between lags and energy flux, should be considered.
The index of relative competition intensity (RCI) has serious built-in biases, due to its asymptotic behavior when competition intensity is high and its tendency to obtain very low values when plants with neighbors intact perform better than neighbor removal plants. These biases have been partially corrected in the index of relative neighbor effect (RNE), but the existence of fixed upper and lower bounds (−1≤RNE≤+1) still creates problems and biases in communities where the average intensity of competition or facilitation is high and plant performance pronouncedly varies in space. The third commonly used index, the logarithm of response ratio (lnRR), is mathematically and statistically sound, but when computed from pair-wise comparisons between neighbor removal and control plants, this index reflects the geometric mean of the treatment effect. Moreover, linear patterns in lnRR reflect exponential patterns in the intensity of competition. As the interest of ecologists usually focuses on arithmetic means, we propose a corrected index of relative competition intensity, CRCI=arc sin (RNE). This index is fairly linear within the observed ranges of competition and facilitation, and for the range of competition intensities where RNE behaves reasonably, the two indices obtain almost identical values.
We compared the performance of the four indices, using both imagined and real data, the latter from systems where the responses of plants to neighbor removal ranged from weak to moderate, so that RNE and CRCI were expected to behave similarly. The indices were computed both from pooled data for each community and as averages of pair-wise comparisons. lnRR and CRCI were found to behave in a consistent and bias-free manner, yielding similar results regardless of method of computation. This was, by and large, the case with RNE, too, but as the values of indices grew, the values from pair-wise comparisons became increasingly smaller than values computed from pooled data. RCI yielded grossly aberrant results in computations based on pair-wise comparisons. Therefore, the further use of RCI is unadvisable and studies where RCI has been derived from pair-wise comparisons should be excluded from meta-analyses.
In arctic-alpine areas tall herb vegetation is restricted to sites with high productivity. At higher elevation, low prostate forbs and narrow-leaved graminoids dominate the vegetation in sites with a protecting snow cover during winter. In this study, I test whether herbivory or abiotic factors prevent tall forbs from growing at higher altitudes. Vegetation blocks from a tall herb meadow were transplanted to herbivore exclosures and open plots in a low-productive snowbed and a productive tall herb meadow. The tall forbs performed equally well in the exclosures on the low-productive snowbed as in the tall herb meadow, but decreased in the open plots on the low-productive snowbed. Thus, even if abiotic factors are ultimately causing many of the vegetation patterns observed in arctic-alpine plant communities, herbivory appears to be the main proximate factor responsible for the decreasing abundance of tall forbs along gradients of decreasing productivity in arctic-alpine areas.
The effects of long-term (I I yr) exclusion of vertebrate herbivores on competition intensity and plant community structure ere studied using manipulative field experiment,; in two arctic-alpine plant communities with contrasting productivity : an unproductive snowbed and a considerably more productive tall herb meadow. In the snowbed. the exclusion of herbivorous mammals resulted in a significant increase in the biomasses of vascular plants and cryptogams, whereas no corresponding response Lis observed on the tall herb meadow. The intensity of competition. measured with a neighbour removal experiment, did not differ significantly between three of the four habitat x treatment combinations - snowbed exposures. meadow exclosures and open meadow plots but as significantly loader on open snowbed plots, Our results thud, suggest that the low competition intensity in the unproductive snow bed is caused by herbivorous mammals, which tend to depress plant biomass in relatively unproductive habitats When herbivorous mammals have been excluded for a sufficiently long time to allots the build-up of plant biomass even in Unproductive habitat,. between-habitat differences in competition intensity disappear.
Positive interactions between plants typically occur where the presence of a species ameliorates the abiotic environment for another. However, there is also a potential for resource competition to act at the same time, which creates a situation where the net outcome is a balance between positive and negative interactions. We present data from a nine-year study in two extreme high alpine habitats that was designed to test whether the effects of established Ranunculus glacialis individuals on germination and growth of Oxyria, digyna are primarily positive or negative at the altitudinal limit of vascular plants. We show net effects ranging from neutral to negative, but no positive effects were detected. We also argue that close associations between plants in these harsh environments may both ameliorate and deteriorate the abiotic environment, and that experimental manipulations are necessary to tell the difference.
Heavy grazing and trampling by reindeer increase nutrient cycling and primary production in areas where grasslands hake replaced shrub and moss tundra. One way in which herbivores can affect nutrient cycling is through changing the litter decomposition processes. We studied the effect of herbivory on litter decomposition rate by reciprocal transplantation of litter between lightly grazed and heavily grazed areas. using the litterbag technique, We used litter from two of the most common species on the lightly grazed side, Betula nana and Empetrum nigrum, and two of the most common species on the heavily grazed side. Carex bigelowii and Deschampsia flexuosa, We found that herbivory improved litter quality by favouring species with easily decomposed litter. However. herbivore also improved litter quality by increasing the nitrogen content and lowering the C/N ratio of each species. Decomposition rates even correlated with the abundance of the plant category in question, Shrub litter decomposed faster in the lightly grazed area where shrubs were common, and graminoid litter decomposed faster in the heavily grazed area where graminoids were common. These results indicate that the decomposer micro-organisms are adapted to the most common litter types. This studs shoals that detailed information about the effect of herbivore on litter quality is important to understand differences between the short-term and long-term effects of herbivory on nutrient cycling and primary production.
Browsing and defoliation often increase the densities of insect herbivores on woody plants. Densities of herbivorous invertebrates were estimated in a long-term grazing manipulation experiment. More then 30-yr-old fences allow us to compare densities of invertebrate herbivores on Salix lanata in areas heavily grazed and areas lightly grazed by reindeer. The number of gall-forming insects (Pontania glabrifons) and gall-forming mitts were higher on the heavily grazed shrubs than on lightly grazed shrubs. In contrast to most short-term studies, the heavily grazed S. lanata had shorter current annual shoots. No difference in leaf size, leaf nitrogen content, or C:N ratio between grazing intensities were detected. However, the enhanced natural delta N-15 value indicates that heavily grazed shrubs get a higher proportion of their N directly from reindeer faeces. Lear weight per unit area and relative fluctuating asymmetry of leaf shape increased in heavily grazed S. lanata. Enhanced relative fluctuating asymmetry might indicate higher susceptibility to herbivores. Long-term grazing seems to increase the density of invertebrate herbivory in the same way as short-term grazing, even if the plant responses differ substantially.
An important process for the persistence of populations subjected to habitat loss and fragmentation is the dispersal of individuals between habitat patches. Dispersal involves emigration from a habitat patch, movement between patches through the surrounding landscape, and immigration into a new suitable habitat patch. Both landscape and physical condition of the disperser are known to influence dispersal ability, although disentangling these effects can often be difficult in the wild. In one of the first studies of its kind, we used an invertebrate model system to investigate how dispersal success is affected by the interaction between the habitat condition, as determined by food availability, and life history characteristics (which are also influenced by food availability). Dispersal of juvenile and adult mites (male and female) from either high food or low food natal patches were tested separately in connected three patch systems where the intervening habitat patches were suitable (food supplied) or unsuitable (no food supplied). We found that dispersal success was reduced when low food habitat patches were coupled to colonising patches via unsuitable intervening patches. Larger body size was shown to be a good predictor of dispersal success, particularly when the intervening landscape is unsuitable. Our results suggest that there is an interaction between habitat fragmentation and habitat suitability in determining dispersal success: if patches degrade in suitability and this affects the ability to disperse successfully then the effective connectance across landscapes may be lowered. Understanding these consequences will be important in informing our understanding of how species, and the communities in which they are embedded, may potentially respond to habitat fragmentation.
Density dependent interactions between higher and lower trophic levels were studied in two consecutive whole lake experiments in a highly productive lake. In the first experiment, the zooplanktivorous fish species, roach (Rutilus rutilus) (primary carnivore), was reduced to 27% of its previous population size (53%) of biomass) by a selective rotenone treatment. In the second experiment, zander (Stizostedion lucioperca) (secondary carnivore), were stocked in the lake resulting in a reduction in the population size of roach to 51% of its previous population size (31% of biomass). In both experiments, seasonal average biomass of Daphnia cucullata) increased in the two years with the lowest roach biomasses. but no shift to larger zooplankton forms took place. In the years with increased Daphnia biomass, phytoplankton biomass and transparency showed an increased seasonal variation (measured as coefficient of variation). In contrast to manipulations in which planktivorous fish were totally removed, no effect on seasonal average phytoplankton biomass was observed. Due to the high reproductive capacity of roach, the system returned to previous conditions within a year or two after the perturbations. Under unperturbed conditions, the lake showed few indications of instability as suggested by the paradox of enrichment hypothesis. This lack of instability can. among other things, be related to high zooplanktivore predation pressure present in highly productive lakes preventing overexploitation of primary producers by grazers. In addition, the omnivorous feeding characteristics of roach (feeding on both zooplankton and algae/detritus) and interactions between the open water and the detritus nutrient pool are likely to increase the stability of the system. The two perturbation experiments plus a previous natural experiment (winter fish kill), which all were pulsed perturbations, provide no evidence for the presence of alternative stable states in highly productive lakes. Possibly, a sustained perturbation over several years may shift highly productive lakes from a dominance of phytoplankton production to a dominance of macrophyte production leading to an alternative state.
Steep climatic gradients may select for clinal adaptation in plant functional traits with implications for interspecific interactions and response to future climate change. Terpenes are common in Mediterranean environments and mediate plant interactions with both the abiotic and biotic environment, including herbivores. Clines in traits such as terpenes have received much attention because they are linked to plant fitness and experience strong selection from the abiotic and biotic environment. In this study, we tested for intraspecific variation in Artemisia californica terpene chemistry in a common garden of plants sourced from populations spanning a large precipitation gradient (6 latitude) and grown in treatments of high and low precipitation. We found genetic variation in terpene richness, diversity, concentration and composition among A. californica populations spanning this species' range. Of these traits, terpene composition and monoterpene concentration varied clinally with respect to source site latitude. Regarding terpene composition, pairwise dissimilarity among populations increased in parallel with geographic distance between source sites. At the same time, monoterpene concentration decreased monotonically from plants of southern origin (source sites with high temperature, aridity, and precipitation variability) to plants of northern origin. Our precipitation manipulation suggests that phenotypic selection by precipitation may underlie this clinal variation in monoterpene concentration, and that monoterpene concentration and other aspects of terpene chemistry are not phenotypically plastic. In summary, this study provides novel evidence for a genetically based latitudinal cline in plant secondary chemistry and suggests that adaptation to a key aspect of the abiotic environment may contribute to this intraspecific variation. Accordingly, changes in terpene chemistry under projected future climates will likely occur solely through the relatively slow process of adaptation, with important consequences for plant interactions with the abiotic environment and a diverse community of associates.
Herbivores drive shifts in plant species composition by interacting with vegetation through defoliation, trampling and nutrient addition: urine and faeces. As herbivore effects on vegetation accumulate over time, they might spillover to other trophic levels, but how and when this happens is poorly understood. Since it is methodologically demanding to measure biodiversity across spatial gradients, an alternative approach is to assess it through biodiversity indices of vascular plants. We employed the Index of biodiversity relevance developed for Swedish flora which provides an estimated number of organisms associated with a plant species, allowing the quantification of trophic community size. Values from this index were coupled with vegetation data from a network of 96 fenced and paired grazed plots across Fennoscandia. We analysed the role herbivory has on plant richness and diversity, and on the number of organisms that interact with the vegetation according to the index values. We also explored how herbivores influence the competitive effects of tall shrubs on other plants since the dominance of a vegetation type links directly to biodiversity. Plant diversity had no clear response to grazing. Overall vegetation and the vegetation subgroups herbs and non-fruit shrubs had higher biodiversity index values in fenced plots, indicating a higher number of plant–host interactions. Herb cover was negatively related to shrubs in both treatments but with a faster decline in the absence of herbivores. This study highlights the importance of maintaining herbivore populations in the Arctic to conserve the vegetation structure and biodiversity of the tundra. This method of coupling biodiversity indexes with vegetation data provides complementary information to the plant diversity, especially when methodological or time constraints prevent complete field inventories.
We studied the effect of productivity on competition intensity and the relationship between competition intensity and community species richness, using a removal experiment with the perennial plant Solidago virgaurea. The experiment was conducted in 16 different communities from two geographically distant areas (western Estonia and northern Norway). The results were compared with the results of previous experiments with Anthoxanthum odoratum from the same areas. Removal of neighbors had a positive effect on the biomass of both Solidago and Anthoxanthum, and this response was stronger in communities with higher productivity. Thus, the corrected index of relative competition intensity, CRCI, increased with increasing community productivity. Species richness was negatively correlated with CRCI in Estonia but not in Norway and not in the case of the pooled material. The results suggest that competitive exclusion operates at least in these communities which species pool is large.
Our results indicate that the relationship between competition intensity and productivity is non-linear. In our data, competition prevails in communities where living plant biomass exceeds 200 g m−2, whereas in less productive communities, competition remains undetected and direct plant–plant relationships might at times be even mutualistic. Moreover, we found that the relationship between competition intensity and productivity is strongly dependent on regional differences and is intimately connected to a concordant variation in the intensity of grazing. The least productive communities both in Estonia and in Norway are characterized by intensive grazing, which reduces importance of competition. Hence, the contrasting results corroborates the predictions of the hypothesis of exploitation ecosystems, predicting that trophic dynamics account for the relationship between competition intensity and primary productivity.
Food webs can respond in surprising and complex ways to temporary alterations in their species composition. When such a perturbation is reversed, food webs have been shown to either return to the pre-perturbation community state or remain in the food web configuration that established during the perturbation. Here we report findings from a replicated whole-lake experiment investigating food web responses to a perturbation and its consecutive reversal. We could identify three distinct community states in the food web that corresponded to the periods before, during and after the perturbation. Most importantly, we demonstrate the establishment of a distinct post-perturbation food web configuration that differed from both the pre- and during-perturbation communities in phytoplankton biomass and micro- and mesozooplankton species composition. We suggest that the pre- and post-perturbation food web configurations may represent two alternative stable community states. We provide explanations for how each of the contrasting communities may be maintained through altered species interactions. These findings add to the discussion of how natural food webs react to environmental change and imply that the range of potential ecosystem dynamics in response to perturbations can be wider and more complex than is often recognized.
A large number of studies have presented empirical arguments for the existence of alternative stable states (ASS) in a wide range of ecological systems. However, most of these studies have used non-manipulative, indirect methods, which findings remain open for alternative explanations. Here, we review the direct evidence for ASS resulting from manipulation experiments. We distinguish four conclusive experimental approaches which test for predictions made by the hysteresis effect: (1) discontinuity in the response to an environmental driving parameter, (2) lack of recovery potential after a perturbation, (3) divergence due to different initial conditions and (4) random divergence. Based on an extensive literature search we found 35 corresponding experiments. We assessed the ecological stability of the reported contrasting states using the minimum turnover of individuals in terms of life span and classified the studies according to 4 categories: (1) experimental system, (2) habitat type, (3) involved organisms and (4) theoretical framework. 13 experiments have directly demonstrated the existence of alternative stable states while 8 showed the absence of ASS in other cases. 14 experiments did not fulfil the requirements of a conclusive test, mostly because they applied a too short time scale. We found a bias towards laboratory experiments compared to field experiments in demonstrating bistability. There was no clear pattern of the distribution of ASS over categories. The absence of ASS in 38% of the tested systems indicates that ASS are just one possibility of how ecological systems can behave. The relevance of the concept of ASS for natural systems is discussed, in particular under consideration of the observed laboratory bias, perturbation frequency and variable environments. It is argued, that even for a permanently transient system, alternative attractors may still be of relevance.