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Spatial and temporal variation in perceived predation risk is an important determinant of movement and foraging activity of animals. Foraging in this landscape of fear, individuals need to decide where and when to move, and what resources to choose. Foraging theory predicts the outcome of these decisions based on energetic trade-offs, but complex interactions between perceived predation risk and preferences of foragers for certain functional traits of their resources are rarely considered. Here, we studied the interactive effects of perceived predation risk on food trait preferences and foraging behavior in bank voles (Myodes glareolus) in experimental landscapes. Individuals (n = 19) were subjected for periods of 24 h to two extreme, risk-uniform landscapes (either risky or safe), containing 25 discrete food patches, filled with seeds of four plant species in even amounts. Seeds varied in functional traits: size, nutrients, and shape. We evaluated whether and how risk modifies forager preference for functional traits. We also investigated whether perceived risk and distance from shelter affected giving-up density (GUD), time in patches, and number of patch visits. In safe landscapes, individuals increased time spent in patches, lowered GUD and visited distant patches more often compared to risky landscapes. Individuals preferred bigger seeds independent of risk, but in the safe treatment they preferred fat-rich over carb-rich seeds. Thus, higher densities of resource levels remained in risky landscapes, while in safe landscapes resource density was lower and less diverse due to selective foraging. Our results suggest that the interaction of perceived risk and dietary preference adds an additional layer to the cascading effects of a landscape of fear which affects biodiversity at resource level.
Wild bee species are important pollinators in agricultural landscapes. However, population decline was reported over the last decades and is still ongoing. While agricultural intensification is a major driver of the rapid loss of pollinating species, transition zones between arable fields and forest or grassland patches, i.e., agricultural buffer zones, are frequently mentioned as suitable mitigation measures to support wild bee populations and other pollinator species. Despite the reported general positive effect, it remains unclear which amount of buffer zones is needed to ensure a sustainable and permanent impact for enhancing bee diversity and abundance. To address this question at a pollinator community level, we implemented a process-based, spatially explicit simulation model of functional bee diversity dynamics in an agricultural landscape. More specifically, we introduced a variable amount of agricultural buffer zones (ABZs) at the transition of arable to grassland, or arable to forest patches to analyze the impact on bee functional diversity and functional richness. We focused our study on solitary bees in a typical agricultural area in the Northeast of Germany. Our results showed positive effects with at least 25% of virtually implemented agricultural buffer zones. However, higher amounts of ABZs of at least 75% should be considered to ensure a sufficient increase in Shannon diversity and decrease in quasi-extinction risks. These high amounts of ABZs represent effective conservation measures to safeguard the stability of pollination services provided by solitary bee species. As the model structure can be easily adapted to other mobile species in agricultural landscapes, our community approach offers the chance to compare the effectiveness of conservation measures also for other pollinator communities in future.
Perceived predation risk varies in space and time. Foraging in this landscape of fear alters forager-resource interactions via cascading nonconsumptive effects. Estimating these indirect effects is difficult in natural systems. Here, we applied a novel measure to quantify the diversity at giving-up density that allows to test how spatial variation in perceived predation risk modifies the diversity of multispecies resources at local and regional spatial levels. Furthermore, we evaluated whether the nonconsumptive effects on resource species diversity can be explained by the preferences of foragers for specific functional traits and by the forager species richness. We exposed rodents of a natural community to artificial food patches, each containing an initial multispecies resource community of eight species (10 items each) mixed in sand. We sampled 35 landscapes, each containing seven patches in a spatial array, to disentangle effects at local (patch) and landscape levels. We used vegetation height as a proxy for perceived predation risk. After a period of three nights, we counted how many and which resource species were left in each patch to measure giving-up density and resource diversity at the local level (alpha diversity) and the regional level (gamma diversity and beta diversity). Furthermore, we used wildlife cameras to identify foragers and assess their species richness. With increasing vegetation height, i.e., decreasing perceived predation risk, giving-up density, and local alpha and regional gamma diversity decreased, and patches became less similar within a landscape (beta diversity increased). Foragers consumed more of the bigger and most caloric resources. The higher the forager species richness, the lower the giving-up density, and alpha and gamma diversity. Overall, spatial variation of perceived predation risk of foragers had measurable cascading effects on local and regional resource species biodiversity, independent of the forager species. Thus, nonconsumptive predation effects modify forager-resource interactions and might act as an equalizing mechanism for species coexistence.
Foraging by consumers acts as a biotic filtering mechanism for biodiversity at the trophic level of resources. Variation in foraging behaviour has cascading effects on abundance, diversity, and functional trait composition of the community of resource species. Here we propose diversity at giving-up density (DivGUD), i.e. when foragers quit exploiting a patch, as a novel concept and simple measure quantifying cascading effects at multiple spatial scales. In experimental landscapes with an assemblage of plant seeds, patch residency of wild rodents decreased local alpha-DivGUD (via elevated mortality of species with large seeds) and regional gamma-DivGUD, while dissimilarity among patches in a landscape (beta-DivGUD) increased. By linking theories of adaptive foraging behaviour with community ecology, DivGUD allows to investigate cascading indirect predation effects, e.g. the ecology-of-fear framework, feedbacks between functional trait composition of resource species and consumer communities, and effects of inter-individual differences among foragers on the biodiversity of resource communities.
Biodiversity conservation and agricultural production have been largely framed as separate goals for landscapes in the discourse on land use. Although there is an increasing tendency to move away from this dichotomy in theory, the tendency is perpetuated by the spatially explicit approaches used in research and management practice. Transition zones (TZ) have previously been defined as areas where two adjacent fields or patches interact, and so they occur abundantly throughout agricultural landscapes. Biodiversity patterns in TZ have been extensively studied, but their relationship to yield patterns and social-ecological dimensions has been largely neglected. Focusing on European, temperate agricultural landscapes, we outline three areas of research and management that together demonstrate how TZ might be used to facilitate an integrated landscape approach: (i) plant and animal species' use and response to boundaries and the resulting effects on yield, for a deeper understanding of how landscape structure shapes quantity and quality of TZ; (ii) local knowledge on field or patch-level management and its interactions with biodiversity and yield in TZ, and (iii) conflict prevention and collaborative management across land-use boundaries.
This review provides a synthesis of current knowledge on the morphological and functional traits of testate amoebae, a polyphyletic group of protists commonly used as proxies of past hydrological changes in paleoecological investigations from peatland, lake sediment and soil archives. A trait-based approach to understanding testate amoebae ecology and paleoecology has gained in popularity in recent years, with research showing that morphological characteristics provide complementary information to the commonly used environmental inferences based on testate amoeba (morpho-)species data. We provide a broad overview of testate amoeba morphological and functional traits and trait-environment relationships in the context of ecology, evolution, genetics, biogeography, and paleoecology. As examples we report upon previous ecological and paleoecological studies that used trait-based approaches, and describe key testate amoebae traits that can be used to improve the interpretation of environmental studies. We also highlight knowledge gaps and speculate on potential future directions for the application of trait-based approaches in testate amoeba research.
Trait-based approaches have broadened our understanding of how the composition of ecological communities responds to environmental drivers. This research has mainly focussed on abiotic factors and competition determining the community trait distribution, while effects of trophic interactions on trait dynamics, if considered at all, have been studied for two trophic levels at maximum. However, natural food webs are typically at least tritrophic. This enables indirect interactions of traits and biomasses among multiple trophic levels leading to underexplored effects on food web dynamics. Here, we demonstrate the occurrence of mutual trait adjustment among three trophic levels in a natural plankton food web (Lake Constance) and in a corresponding mathematical model. We found highly recurrent seasonal biomass and trait dynamics, where herbivorous zooplankton increased its size, and thus its ability to counter phytoplankton defense, before phytoplankton defense actually increased. This is contrary to predictions from bitrophic systems where counter-defense of the consumer is a reaction to prey defense. In contrast, counter-defense of carnivores by size adjustment followed the defense of herbivores as expected. By combining observations and model simulations, we show how the reversed trait dynamics at the two lower trophic levels result from a "trophic biomass-trait cascade" driven by the carnivores. Trait adjustment between two trophic levels can therefore be altered by biomass or trait changes of adjacent trophic levels. Hence, analyses of only pairwise trait adjustment can be misleading in natural food webs, while multitrophic trait-based approaches capture indirect biomass-trait interactions among multiple trophic levels.
The shape of a defense-growth trade-off governs seasonal trait dynamics in natural phytoplankton
(2020)
Theory predicts that trade-offs, quantifying costs of functional trait adjustments, crucially affect community trait adaptation to altered environmental conditions, but empirical verification is scarce. We evaluated trait dynamics (antipredator defense, maximum growth rate, and phosphate affinity) of a lake phytoplankton community in a seasonally changing environment, using literature trait data and 21 years of species-resolved high-frequency biomass measurements. The trait data indicated a concave defense-growth trade-off, promoting fast-growing species with intermediate defense. With seasonally increasing grazing pressure, the community shifted toward higher defense levels at the cost of lower growth rates along the trade-off curve, while phosphate affinity explained some deviations from it. We discuss how low fitness differences of species, inferred from model simulations, in concert with stabilizing mechanisms, e.g., arising from further trait dimensions, may lead to the observed phytoplankton diversity. In conclusion, quantifying trade-offs is key for predictions of community trait adaptation and biodiversity under environmental change.
Trait-based approaches to investigate (short- and long-term) phytoplankton dynamics and community assembly have become increasingly popular in freshwater and marine science. Although the nature of the pelagic habitat and the main phytoplankton taxa and ecology are relatively similar in both marine and freshwater systems, the lines of research have evolved, at least in part, separately. We compare and contrast the approaches adopted in marine and freshwater ecosystems with respect to phytoplankton functional traits. We note differences in study goals relating to functional trait use that assess community assembly and those that relate to ecosystem processes and biogeochemical cycling that affect the type of characteristics assigned as traits to phytoplankton taxa. Specific phytoplankton traits relevant for ecological function are examined in relation to
herbivory, amplitude of environmental change and spatial and temporal scales of study. Major differences are identified, including the shorter time scale for regular environmental change in freshwater ecosystems compared to that in the open oceans as well as the
type of sampling done by researchers based on site-accessibility. Overall, we encourage researchers to better motivate why they apply trait-based analyses to their studies and to make use of process-driven approaches, which are more common in marine studies. We further propose fully comparative trait studies conducted along the habitat gradient spanning freshwater to brackish to marine systems, or along geographic gradients. Such studies will benefit from the combined strength of both fields.
Trade-offs between functional traits are ubiquitous in nature and can promote species coexistence depending on their shape. Classic theory predicts that convex trade-offs facilitate coexistence of specialized species with extreme trait values (extreme species) while concave trade-offs promote species with intermediate trait values (intermediate species). We show here that this prediction becomes insufficient when the traits translate non-linearly into fitness which frequently occurs in nature, e.g., an increasing length of spines reduces grazing losses only up to a certain threshold resulting in a saturating or sigmoid trait-fitness function. We present a novel, general approach to evaluate the effect of different trade-off shapes on species coexistence. We compare the trade-off curve to the invasion boundary of an intermediate species invading the two extreme species. At this boundary, the invasion fitness is zero. Thus, it separates trait combinations where invasion is or is not possible. The invasion boundary is calculated based on measurable trait-fitness relationships. If at least one of these relationships is not linear, the invasion boundary becomes non-linear, implying that convex and concave trade-offs not necessarily lead to different coexistence patterns. Therefore, we suggest a new ecological classification of trade-offs into extreme-favoring and intermediate-favoring which differs from a purely mathematical description of their shape. We apply our approach to a well-established model of an empirical predator-prey system with competing prey types facing a trade-off between edibility and half-saturation constant for nutrient uptake. We show that the survival of the intermediate prey depends on the convexity of the trade-off. Overall, our approach provides a general tool to make a priori predictions on the outcome of competition among species facing a common trade-off in dependence of the shape of the trade-off and the shape of the trait-fitness relationships.
1. During the last couple of decades, invasive species have become a worldwide problem in many freshwater systems. Besides higher plants and animals, microbes, in particular the potentially toxic cyanobacterium Cylindrospermopsis raciborskii, has attracted increasing attention, due to its spread towards temperate zones of the northern and southern hemisphere. A number of advantageous functional traits and a high intraspecific plasticity have been suggested to explain its invasion success. 2. The aim of this study was to examine intraspecific functional trait variability in 12 different isolates of C.raciborskii originating from different lakes in an invaded region in Northeast Germany. We measured growth rate, C:N:P ratios, chlorophyll-a content and the abundance of heterocysts under nutrient-replete and phosphorus-limited conditions. Moreover, the isolate-specific morphology and grazing losses by an herbivorous rotifer, as a top-down force, were studied. 3. DNA fingerprinting revealed that all isolates were genetically different. C.raciborskii exhibited a large variability in all measured traits among isolates. The C:P, N:P and Chl-a:C ratios differed by a factor of two or more. The trait variability among isolates was higher under nutrient-replete conditions, except for the C:P ratio, which varied most during phosphorus limitation. The susceptibility to grazing, calculated as maximum ingestion rates of the rotifer Brachionus calyciflorus on C.raciborskii, varied most among isolates, but was not related to any of the measured physiological or morphological traits, i.e. no trade-off was found. 4. Ecological and genetic clustering did not match, indicating that the genetic relationship based on DNA fingerprinting did not cover ecological differences. 5. Our results show a high trait variability within locally occurring and partly co-occurring C.raciborskii isolates. No overall trade-offs between the measured functional traits were found. This demonstrates the ecological relevance of linking multiple traits, e.g. competitive and consumptive. Furthermore, this study emphasises the importance of analysing more than one strain of a species, as different strains show different trait values potentially relevant for their invasibility and the field of general trait-based ecology.
Trait-based approaches have become increasingly successful in community ecology. They assume that the distribution of functional traits within communities responds in a predictable way to alterations in environmental forcing and that strong forcing may accelerate such trait changes. We used high frequency measurements of phytoplankton to test these assumptions. We analyzed the seasonal and long-term dynamics of the community trait mean within a multi-dimensional trait space under alternating multifactorial environmental conditions. The community trait mean exhibited a distinct recurrent annual pattern that reflected minor changes in climate, herbivory and nutrients. Independent of early spring conditions, the community trait mean was repeatedly driven into a narrow confined area in the trait space under pronounced herbivory during the clear water phase. The speed of movement was highest at the onset and the relaxation of such strong unidirectional forcing. Thus, our data support the conceptual framework of trait-based ecology that alterations in environmental conditions are systematically tracked by adjustments in the dominant functional trait values and that the speed of trait changes depends on the kind and intensity of the selection pressure. Our approach provides a sensitive tool to detect small functional differences in the community related to subtle differences in forcing.
The importance of intraspecific trait variability for community dynamics and ecosystem functioning has been underappreciated. There are theoretical reasons for predicting that species that differ in intraspecific trait variability will also differ in their effects on ecosystem functioning, particularly in variable environments. We discuss whether species with greater trait variability are likely to exhibit greater temporal stability in their population dynamics, and under which conditions this might lead to stability in ecosystem functioning. Resolving this requires us to consider several questions. First, are species with high levels of variation for one trait equally variable in others? In particular, is variability in response and effects traits typically correlated? Second, what is the relative contribution of local adaptation and phenotypic plasticity to trait variability? If local adaptation dominates, then stability in function requires one of two conditions: (i) individuals of appropriate phenotypes present in the environment at high enough frequencies to allow for populations to respond rapidly to the changing environment, and (ii) high levels of dispersal and gene flow. While we currently lack sufficient information on the causes and distribution of variability in functional traits, filling in these key data gaps should increase our ability to predict how changing biodiversity will alter ecosystem functioning.
In most biodiversity studies, taxonomic diversity is the measure for the multiplicity of species and is often considered to represent functional diversity. However, trends in taxonomic diversity and functional diversity may differ, for example, when many functionally similar but taxonomically different species co-occur in a community. The differences between these diversity measures are of particular interest in diversity research for understanding diversity patterns and their underlying mechanisms. We analysed a temporally highly resolved 20-year time series of lake phytoplankton to determine whether taxonomic diversity and functional diversity exhibit similar or contrasting seasonal patterns. We also calculated the functional mean of the community in n-dimensional trait space for each sampling day to gain further insights into the seasonal dynamics of the functional properties of the community. We found an overall weak positive relationship between taxonomic diversity and functional diversity with a distinct seasonal pattern. The two diversity measures showed synchronous behaviour from early spring to mid-summer and a more complex and diverging relationship from autumn to late winter. The functional mean of the community exhibited a recurrent annual pattern with the most prominent changes before and after the clear-water phase. From late autumn to winter, the functional mean of the community and functional diversity were relatively constant while taxonomic diversity declined, suggesting competitive exclusion during this period. A further decline in taxonomic diversity concomitant with increasing functional diversity in late winter to early spring is seen as a result of niche diversification together with competitive exclusion. Under these conditions, several different sets of traits are suitable to thrive, but within one set of functional traits only one, or very few, morphotypes can persist. Taxonomic diversity alone is a weak descriptor of trait diversity in phytoplankton. However, the combined analysis of taxonomic diversity and functional diversity, along with the functional mean of the community, allows for deeper insights into temporal patterns of community assembly and niche diversification.
A unified understanding of the relationship between disturbance and biodiversity is needed to predict biotic responses to global change. Recent advances have identified the need to deconstruct traditional models of disturbance into intensity and frequency to reconcile empirical studies that appear to generate contradictory associations between species diversity and disturbance. We integrate results from theoretical simulation modelling, field-based surveys of 5176 vegetation plots from 48 transects across 6 sites, and experimental pot-based manipulations of flooding to identify how disturbance drives species diversity within ephemeral wetlands in South Island, New Zealand. We find empirical, hump-shaped and positive relationships between species diversity and both disturbance intensity and frequency, mirroring patterns from a simulation model in which species differed in their demographic responses to disturbance. More generally, our simulations show that the relationships between diversity and disturbance shift from positive to hump-shaped to negative as species that are favored at low disturbance because of their resistance strategies, defined by low mortality and recruitment, decline within communities relative to resilient species. Resilient species with higher mortality and recruitment rates are instead favored as disturbance intensity and frequency intensify. Our theoretical findings suggest that sites must also have a third group of unique species with intermediate resilience and resistance. Analyses of community composition along our disturbance gradients support this prediction, emphasizing that shifts in community-level resistance and resilience drive empirical associations between diversity and disturbance. Overall, terrestrial plants may be unable to resist intense and frequent flooding, even with specialized traits. Only fast-growing species with high regeneration from seed may respond once flooding subsides and dominate community composition in these situations, especially on nutrient-rich soils. However, different strategies can co-occur at intermediate disturbance, ultimately increasing species richness. As disturbances become more pervasive globally, our results suggest that differences in the niches of species, rather than demographic stochasticity, drive biodiversity patterns. These niche-based processes may especially prevail, without accompanying losses in species richness, where sites are initially dominated by resistant taxa or life history strategies that balance resistance and resilience.
Silvicultural practices lead to changes in forest composition and structure and may impact species diversity from the overall regional species pool to stand-level species occurrence. We explored to what extent fine-scale occupancy patterns in differently managed forest stands are driven by environment and ecological traits in three regions in Germany using a multi-species hierarchical model. We tested for the possible impact of environmental variables and ecological traits on occupancy dynamics in a joint modelling exercise while taking possible variation in coefficient estimates over years and plots into account. Bird species richness differed across regions and years, and trends in species richness across years were different in the three regions. On the species level, forest management affected occupancy of species in all regions, but only 35% of the total assemblage-level variation in occurrence probability was explained by either forest type and successional stage and <?1% by forest edge. On the assemblage level, bird occurrence decreased with body mass in all regions. Species with smaller breeding ranges had lower occurrence probabilities in one region, while later spring arrival decreased occurrence probabilities in the two other regions. Spatial variation in the effect size of trait covariates such as species phylogeny and breeding strata showed that variation in patch occupancy due to fine-scale differences in forest management is, to some extent, predictable from ecological traits. Our results show that environmental factors and ecological traits jointly predict variation in bird occupancy patterns and their response to forest management. Observations at the fine scale of forest stands, at which conservation efforts can be arranged along with forest management practices in heterogeneous environments, have been shown to provide meaningful insights despite the difficulties involved in monitoring mobile organisms such as birds at the plot level.
1. Worldwide, the floristic composition of temperate forests bears the imprint of past land use for decades to centuries as forests regrow on agricultural land. Many species, however, display significant interregional variation in their ability to (re)colonize post-agricultural forests. This variation in colonization across regions and the underlying factors remain largely unexplored.
2. We compiled data on 90 species and 812 species x study combinations from 18 studies across Europe that determined species' distribution patterns in ancient (i.e. continuously forested since the first available land use maps) and post-agricultural forests. The recovery rate (RR) of species in each landscape was quantified as the log-response ratio of the percentage occurrence in post-agricultural over ancient forest and related to the species-specific life-history traits and local (soil characteristics and light availability) and regional factors (landscape properties as habitat availability, time available for colonization, and climate).
3. For the herb species, we demonstrate a strong (interactive) effect of species' life-history traits and forest habitat availability on the RR of post-agricultural forest. In graminoids, however, none of the investigated variables were significantly related to the RR.
4. The better colonizing species that mainly belonged to the short-lived herbs group showed the largest interregional variability. Their recovery significantly increased with the amount of forest habitat within the landscape, whereas, surprisingly, the time available for colonization, climate, soil characteristics and light availability had no effect.
5. Synthesis. By analysing 18 independent studies across Europe, we clearly showed for the first time on a continental scale that the recovery of short-lived forest herbs increased with the forest habitat availability in the landscape. Small perennial forest herbs, however, were generally unsuccessful in colonizing post-agricultural forest even in relatively densely forested landscapes. Hence, our results stress the need to avoid ancient forest clearance to preserve the typical woodland flora.
1. Improving the mechanistic basis of biodiversity-ecosystem function relationships requires a better understanding of how functional traits drive the dynamics of populations. For example, environmental disturbances or grazing may increase synchronization of functionally similar species, whereas functionally different species may show independent dynamics, because of different responses to the environment. Competition for resources, on the other hand, may yield a wide range of dynamic patterns among competitors and lead functionally similar and different species to display synchronized to compensatory dynamics. The mixed effect of these forces will influence the temporal fluctuations of populations and, thus, the variability of aggregate community properties.
2. To search for a relationship between functional and dynamics similarity, we studied the relationship between functional trait similarity and temporal dynamics similarity for 36 morphotypes of phytoplankton using long-term high-frequency measurements.
3. Our results show that functionally similar morphotypes exhibit dynamics that are more synchronized than those of functionally dissimilar ones. Functionally dissimilar morphotypes predominantly display independent temporal dynamics. This pattern is especially strong when short time-scales are considered.
4. Negative correlations are present among both functionally similar and dissimilar phytoplankton morphotypes, but are rarer and weaker than positive ones over all temporal scales.
5. Synthesis. We demonstrate that diversity in functional traits decreases community variability and ecosystem-level properties by decoupling the dynamics of individual morphotypes.