A multivariate approach reveals diversity of ontogenetic niche shifts across taxonomic and functional groups

1. Shifts in the fundamental and realised niche of individuals during their ontogeny are ubiquitous in nature, but we know little about what aspects of the niche change and how these changes vary across species within communities. However, this knowledge is essential to predict the dynamics of populations and communities and how they respond to environmental change.
2. Here I introduce a range of metrics to describe different aspects of shifts in the realised trophic niche of individuals based on stable isotopes. Applying this multi-variate approach to 2,272 individuals from 13 taxonomic and functional distinct species (Amphibia, Hemiptera, Coleoptera, Odonata) sampled in natural pond communities allowed me to: (1) describe and quantify the diversity of trophic niche shift patterns over ontogeny in multi-dimensional space, and (2) identify what aspects of ontogenetic shifts vary across taxa, and functional groups.
3. Results revealed that species can differ substantially in which aspects of the trophic niche change and how they change over ontogeny. Interestingly, patterns of ontogenetic niche shifts grouped in distinct taxonomic clusters in multi-variate space, including two distinct groups of predators (Hemiptera versus Odonata). Given the differences in traits (especially feeding mode) across groups, this suggests that differences in ontogenetic niche shifts across species could at least partially be explained by variation in traits and functional roles of species.
4. These results emphasise the importance of a multivariate approach to capture the large diversity of trophic niche shifts patterns possible in natural communities and suggest that differences in ontogenetic niche shifts follow general patterns.

     

High intraspecific variability in the functional niche of a predator is associated with ontogenetic shift and individual specialization

Investigations on the functional niche of organisms have primarily focused on differences among species and tended to neglect the potential effects of intraspecific variability despite the fact that its potential ecological and evolutionary importance is now widely recognized. In this study, we measured the distribution of functional traits in an entire population of largemouth bass (Micropterus salmoides) to quantify the magnitude of intraspecific variability in functional traits and niche (size, position, and overlap) between age classes. Stable isotope analyses (δ¹³C and δ¹⁵N) were also used to determine the association between individual trophic ecology and intraspecific functional trait variability. We observed that functional traits were highly variable within the population (mean coefficient variation: 15.62% ± 1.78% SE) and predominantly different between age classes. In addition, functional and trophic niche overlap between age classes was extremely low. Differences in functional niche between age classes were associated with strong changes in trophic niche occurring during ontogeny while, within age classes, differences among individuals were likely driven by trophic specialization. Each age class filled only a small portion of the total functional niche of the population and age classes occupied distinct portions in the functional space, indicating the existence of ontogenetic specialists with different functional roles within the population. The high amplitude of intraspecific variability in functional traits and differences in functional niche position among individuals reported here supports the recent claims for an individual‐based approach in functional ecology.     

Trait‐based community assembly of aquatic macrophytes along a water depth gradient in a freshwater lake

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Convergent changes in the trophic ecology of extremophile fish along replicated environmental gradients

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Between‐lake variation in the trophic ecology of an invasive crayfish

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Seasonal polyphenism of spotted‐wing Drosophila is affected by variation in local abiotic conditions within its invaded range,likely influencing survival and regional population dynamics

1. Overwintering Drosophila often display adaptive phenotypic differences beneficial for survival at low temperatures. However, it is unclear which morphological traits are the best estimators of abiotic conditions, how those traits are correlated with functional outcomes in cold tolerance, and whether there are regional differences in trait expression.
2. We used a combination of controlled laboratory assays, and collaborative field collections of invasive Drosophila suzukii in different areas of the United States, to study the factors affecting phenotype variability of this temperate fruit pest now found globally.
3. Laboratory studies demonstrated that winter morph (WM) trait expression is continuous within the developmental temperature niche of this species (10–25°C) and that wing length and abdominal melanization are the best predictors of the larval abiotic environment.
4. However, the duration and timing of cold exposure also produced significant variation in development time, morphology, and survival at cold temperatures. During a stress test assay conducted at ?5°C, although cold tolerance was greater among WM flies, long‐term exposure to cold temperatures as adults significantly improved summer morph (SM) survival, indicating that these traits are not controlled by a single mechanism.
5. Among wild D. suzukii populations, we found that regional variation in abiotic conditions differentially affects the expression of morphological traits, although further research is needed to determine whether these differences are genetic or environmental in origin and whether thermal susceptibility thresholds differ among populations within its invaded range.

     

Trophic response to ecological conditions of habitats: Evidence from trophic variability of freshwater fish

1. To adapt to ecological and environmental conditions, species can change their ecological niche (e.g., interactions among species) and function (e.g., prey‐predation, diet competition, and habitat segregation) at the species and guild levels. Stable isotope analysis of bulk carbon and nitrogen of organisms has conventionally been used to evaluate such adaptabilities in the scenopoetic and bionomic views as the isotopic niche width.
2. Compound‐specific stable isotope analysis (CSIA) of nitrogen within amino acids provides trophic information without any disruption of scenopoetic views in the isotope ratios, unlike conventional bulk isotope analysis provides both information and therefore frequently hinders its usefulness for trophic information.
3. We performed CSIA of amino acids to understand the trophic variability of the pike gudgeon Pseudogobio esocinus and largemouth bass Micropterus salmoides as representative specialist and generalist fish species, respectively, from 16 ecologically variable habitats in the four major rivers of Korea.
4. There was little variation (1σ) in the trophic position (TP) among habitats for P. esocinus (± 0.2); however, there was considerably large variation for M. salmoides (± 0.6). The TP of M. salmoides was negatively correlated with the benthic invertebrate indices of the habitats, whereas the TP of P. esocinus showed no significant correlation with any indices. Thus, these two representative fish species have different trophic responses to ecological conditions, which is related to known differences in the trophic niche between specialists (i.e., small niche width) and generalists (i.e., large niche width).
5. Over the past four decades, the conventional bulk isotope analysis has not been capable of deconvoluting “scenopoetic” and “bionomic” information. However, in the present study, we demonstrated that the CSIA of amino acids could isolate trophic niches from the traditional ecological niche composed of trophic and habitat information and evaluated how biological and ecological indices influence the trophic response of specialists and generalists.

     

Beyond ecological opportunity: Prey diversity rather than abundance shapes predator niche variation

1. Ecological opportunity (i.e. the diversity of available resources) has a pivotal role in shaping niche variation and trophic specialisation of animals. However, ecological opportunity can be described with regard to both diversity and abundance of resources. The degree to which these two components contribute to niche variation remains unexplored.
2. To address this, we used an extensive dataset on fish diet and benthic invertebrate diversity and density from 73 sampling events in three Norwegian rivers in order to explore realised trophic niches and the response of dietary niche variation along gradients of resource diversity (potential trophic niches), resource density (as a proxy of resource abundance) and fish density (as a proxy of inter‐ and intra‐specific competition) in a freshwater top predator (the brown trout, Salmo trutta L.).
3. Linear models indicated that individual and population niche variation increased with increasing ecological opportunity in terms of prey diversity. However, no simple cause‐and‐effect associations between niche indices and prey abundance were found. Our multiple regression analyses indicated that the abundance of certain resources (e.g. Chironomidae) can interact with prey diversity to determine individual and population realised trophic niches. Niche variation (within‐individual component and inter‐individual diet variation) decreased with increasing inter‐ and intra‐specific competition.
4. This study extends prevailing trophic ecology theory by identifying diversity, rather than density, of available prey resources as a primary driver of niche variation in fish of temperate riverine systems with no extensive resource limitation. The study also shows that ecological opportunity may mask the direction of the effect (compression or expansion) of competition on niche variation when food resources are diverse.
5. Our study provides novel empirical insight to the driving forces behind niche variation and reveals that diversity, rather than density, of available prey resources may be a primary driver of niche variation in freshwater fish. Our study supports the view that a broader potential trophic niche promotes broader realised trophic niche variation by individuals, which leads to individual niche diversification by opening access to alternatives resources, resulting in a concomitant rise in the realised trophic niche width of the population.

     

Unraveling the relative contribution of inter‐ and intrapopulation functional variability in wild populations of a tadpole species

Functional traits are increasingly recognized as an integrative approach by ecologists to quantify a key facet of biodiversity. And these traits are primarily expressed as species means in previous studies, based on the assumption that the effects of intraspecific variability can be overridden by interspecific variability when studying functional ecology at the community level. However, given that intraspecific variability could also have important effects on community dynamics and ecosystem functioning, empirical studies are needed to investigate the importance of intraspecific variability in functional traits. In this study, 256 Scutiger boulengeri tadpole individuals from four different populations are used to quantify the functional difference between populations within a species, and the relative contribution of inter‐ and intrapopulation variability in functional traits. Our results demonstrate that these four populations differ significantly in functional attributes (i.e., functional position, functional richness, and low functional overlap), indicating that individuals from different populations within a species should be explicitly accounted for in functional studies. We also find similar relative contribution of inter‐ (~56%) and intrapopulation (~44%) variation to the total variability between individuals, providing evidence that individuals within populations should also be incorporated in functional studies. Overall, our results support the recent claims that intraspecific variability cannot be ignored, as well as the general idea of “individual level” research in functional ecology.     

Trait coordination and environmental filters shape functional trait distributions of forest understory herbs

1. Understanding the drivers of trait selection is critical for resolving community assembly processes. Here, we test the importance of environmental filtering and trait covariance for structuring the functional traits of understory herbaceous communities distributed along a natural environmental resource gradient that varied in soil moisture, temperature, and nitrogen availability, produced by different topographic positions in the southern Appalachian Mountains.
2. To uncover potential differences in community‐level trait responses to the resource gradient, we quantified the averages and variances of both abundance‐weighted and unweighted values for six functional traits (vegetative height, leaf area, specific leaf area, leaf dry matter content, leaf nitrogen, and leaf δ¹³C) using 15 individuals of each of the 108 species of understory herbs found at two sites in the southern Appalachians of western North Carolina, USA.
3. Environmental variables were better predictors of weighted than unweighted community‐level average trait values for all but height and leaf N, indicating strong environmental filtering of plant abundance. Community‐level variance patterns also showed increased convergence of abundance‐weighted traits as resource limitation became more severe.
4. Functional trait covariance patterns based on weighted averages were uniform across the gradient, whereas coordination based on unweighted averages was inconsistent and varied with environmental context. In line with these results, structural equation modeling revealed that unweighted community‐average traits responded directly to local environmental variation, whereas weighted community‐average traits responded indirectly to local environmental variation through trait coordination.
5. Our finding that trait coordination is more important for explaining the distribution of weighted than unweighted average trait values along the gradient indicates that environmental filtering acts on multiple traits simultaneously, with abundant species possessing more favorable combinations of traits for maximizing fitness in a given environment.

     

Intraspecific and interspecific trait variability in tadpole meta‐communities from the Brazilian Atlantic rainforest

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Trophic complexity of small fish in nearshore food webs

Small nearshore fishes are an important part of lacustrine and functional diversity and link pelagic and benthic habitats by serving as prey for larger nearshore and offshore fishes. However, the trophic complexity of these small nearshore fishes is often unrecognized and detailed studies of their role in food webs are lacking. Here, we examined niche space patterns of small nearshore fish species using Bayesian analyses of carbon and nitrogen stable isotope data in nine freshwater lakes that are among the largest lakes in Minnesota. We found considerable variability in niche areas within species and high variability in niche overlap across species. At the assemblage level, niche overlap (average diet overlap of all species pairs at a lake) decreased as whole-lake species richness increased, possibly indicating a greater degree of resource specialization in more speciose lakes. Overall fish niche space was weakly but significantly related to niche space of their invertebrate prey. Although nearshore benthic resources contributed to fish diets in all lakes, all fish species also had non-negligible and variable contributions from pelagic zooplankton. This inter- and intraspecific variability in trophic niche space likely contributes to the multi-level trophic complexity, functional diversity, and potentially food web resilience to ecosystem changes.

     

Functional structure and diversity of invertebrate communities in a glacierised catchment of the tropical Andes

1. In many mountainous areas, glaciers feed streams characterised by harsh environmental conditions, such as low water temperature, high turbidity, low channel stability, and high temporal variability in flow. Additionally, in many glacierised catchments, the mixture of streams arising from different water sources (glacier melt, groundwater, rainfall) generates high levels of environmental heterogeneity, which enhance species turnover rates and increase regional diversity.
2. Studies from mainly temperate regions have revealed some consistent patterns: a predominance of traits adaptive to harsh environmental conditions and reduced functional diversity with increased glaciality, both strongly related to environmental filtering. Here, we investigated variation in functional structure and diversity between macroinvertebrate communities from 15 stream sites, with different water sources (five glacier-fed, five groundwater-fed, and five mixed source) and level of glacier influence, in a glacierised catchment in the Ecuadorian Andes.
3. Our results revealed functional differences between communities inhabiting the different stream types. As found in temperate regions, high levels of glaciality were associated with an increase of small-sized taxa that do not swim but are temporarily attached to or burrow in the substrate, have a flying-adult stage, and feed by collecting–gathering. Similarly, we found a general decrease in functional diversity at sites with higher glacier influence. A null modelling approach suggested that in some of our glacier-fed sites, environmental filtering may be the main driver of community assembly, whereas other mechanisms, mainly regional (such as dispersal), but also local (such as intraspecific competition), may gain importance as glacier influence decreases.
4. Assemblage composition in streams in tropical glacierised catchments may be driven by both local and regional processes that generate a gradient of decreasing functional diversity with stronger glacier influence. However, lack of knowledge of relevant traits for taxa in tropical glacierised streams currently poses a substantial obstacle to predicting changes likely to arise from global warming and glacier melt in this region.

     

Oribatid mites reveal that competition for resources and trophic structure combine to regulate the assembly of diverse soil animal communities

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Intrapopulation variability in a functional trait: Susceptibility of Daphnia to limitation by dietary fatty acids

1. Functional traits are measurable characteristics of an organism that have an impact on its fitness. Variation in functional traits between and among species has been suggested to represent the basis for competition and selection, thus allowing for evolution in natural populations.
2. In freshwater ecosystems, the availability of essential polyunsaturated fatty acids (PUFAs), in particular ω3‐ and ω6‐PUFAs, determines the food quality of phytoplankton for the herbivorous zooplankton Daphnia, an unselective filter feeder. The content of such essential PUFAs in the phytoplankton is thus a functional phytoplankton trait affecting the trophic transfer efficiency and dynamics at the pelagic plant–herbivore interface.
3. In turn, the susceptibility of consumers to become limited by the availability of essential PUFAs is a fitness‐determining trait of Daphnia genotypes, and variability of this herbivore trait may thus affect the daphnids’ intrapopulation competition. To estimate the intrapopulation variation in susceptibility, we isolated clonal lines of Daphnia longispina from a natural population and compared the strength of their limitation by dietary PUFA availability via standardised laboratory growth assays. We used a liposome supplementation technique to enrich a PUFA‐poor green alga with essential ω3‐ and ω6‐PUFAs and determined juvenile somatic growth rate of different D. longispina genotypes as a fitness proxy.
4. As expected, D. longispina genotypes that coexisted in a natural population differed markedly in their specific patterns of susceptibility to dietary PUFA availability. On average, the D. longispina population was more strongly susceptible to limitations in the availability of the ω6‐PUFA arachidonic acid (20:4ω6) than to limitations in the availability of ω3‐PUFAs α‐linolenic acid (18:3ω3) and eicosapentaenoic acid (20:5ω3).
5. The ability to cope with PUFA limitation is thus a crucial trait that can probably affect intraspecific competition and Daphnia population structure. Therefore, we suggest that such intrapopulation variation in susceptibility to absence of dietary PUFAs might be one of the driving forces of natural selection and local adaptation among freshwater zooplankton.

     

Effects of water clarity and other environmental factors on trophic niches of two sympatric piscivores

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Behavioural plasticity and trophic niche shift: How wintering geese respond to habitat alteration

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Testing for functional significance of traits: Effect of the light environment in tropical tree saplings

1. Functional traits have been examined to explain the growth rates of forest communities in different sites. However, weak or nonexistent relations are often found, especially due to the following methodological aspects: 1) lack of an environmental context (e.g., light, water, or nutrient supply), 2) use of nonfunctional traits, 3) an approach that does not contemplate phenotypic integration, and 4) neglect of intraspecific variation.
2. Here we measured relative growth rates, crown, and leaf traits in saplings of six tropical tree species growing in two light environments (Gap and Understory) to test whether contrasting light environments modulates trait–trait and trait–growth relationships. Moreover, we tested whether models that integrate traits of different dimensions of the plant (crown and leaf) improve the strength of trait–growth relations.
3. Light availability changed both trait–trait and trait–growth relationships. Overall, in Understory, crown traits (crown length and total leaf area) have a stronger effect on growth rates, while physiological traits related to nutrient acquisition (nitrogen concentration), photochemical efficiency (chlorophyll pigments and chlorophyll a fluorescence), and biochemical efficiency (potassium use efficiency) are strong in Gap. Models including multiple traits explained growth rates better in Gap (up to 62%) and Understory (up to 47%), but just in Gap the best model comprises traits that are representative of different dimensions of the plant.
4. Synthesis. We advanced the knowledge behind the light effects on tree sapling by posit that trait–trait and trait–growth relationships vary across light environments. Therefore, light availability is a key environmental factor to be considered when choosing the set of traits to be measured in functional approach studies using tropical tree saplings. In compliance with the phenotype integration hypothesis, functional traits are better predictors of growth rates when grouped in a set of traits of different dimensions of the plant that represent different functional mechanisms.

     

Some like it deep: Intraspecific niche segregation in ruffe (Gymnocephalus cernua)

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Different groups of ground-dwelling spiders share similar trophic niches in temperate forests

1. Generalistic interactions between predator and prey may vary with ecosystem type, predator traits, and prey traits, but the interplay of these factors has not been assessed in ground food webs.
2. We investigated trophic interactions of ground-dwelling spiders across eight forests in European Russia associated with body size, hunting strategy, microhabitat specialization, potential prey type, potential prey population density, and forest type (coniferous vs. broadleaved). We analyzed 128 individual spiders, including juveniles, all identified to the family level with two complementary methods: molecular gut content analysis, and stable isotope analysis of carbon and nitrogen.
3. The results suggest that feeding frequency of spiders is affected by predator body size and by selection of certain prey type. Stable isotope analysis showed similar trophic niches among spider families, varying moderately with forest type. Larger spiders had higher Δ¹³C values than smaller ones, but similar Δ¹⁵N values, suggesting that different size classes of spiders belong to different food chains. Results based on stable isotope and molecular gut content analyses were weakly linked, indicating them targeting different trophic niche dimensions.
4. At least for the group-level interactions, family identity and hunting strategy of predator has little predictive power while predator body size and prey traits affected trophic niche dimensions calling for future studies in this direction. Large spiders feed more and rely on different basal resources than small spiders, suggesting that including small species and juveniles provides a more comprehensive picture of food web organization.