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.

     

Effects of foundation species genotypic diversity on subordinate species richness in an assembling community

Foundation (dominant or matrix) species play a key role in structuring plant communities, influencing processes from population to ecosystem scales. However, the effects of genotypic diversity of foundation species on these processes have not been thoroughly assessed in the context of assembling plant communities. We modified the classical filter model of community assembly to include genotypic diversity as part of the biotic filter. We hypothesized that the proportion of fit genotypes (i.e. competitively superior and dominant) affects niche space availability for subordinate species to establish with consequence for species diversity. To test this hypothesis, we used an individual‐based simulation model where a foundation species of varying genotypic diversity (number of genotypes and variability among genotypes) competes for space with subordinate species on a spatially heterogeneous lattice. Our model addresses a real and practical problem in restoration ecology: choosing the level of genetic diversity of re‐introduced foundation and subordinate species. Genotypic diversity of foundation species significantly affected equilibrium community diversity, measured as species richness, either positively or negatively, depending upon environmental heterogeneity. Increases in genotypic diversity gave the foundation species a wider niche breadth. Under conditions of high environmental heterogeneity, this wider niche breadth decreased niche space for other species, lowering species richness with increased genotypic diversity until the genotypes of the foundation species saturated the landscape. With a low level of environmental heterogeneity, increasing genotypic diversity caused the foundation species niche breadth to be overdispersed, resulting in a weak positive relationship with species richness. Under these conditions, some genotypes are maladapted to the environment lowering fitness of the foundation species. These effects of genotypic diversity were secondary to the larger effects of overall foundation species fitness and environmental heterogeneity. The novel aspect of incorporating genotype diversity in combination with environmental heterogeneity in community assembly models include predictions of either positive or negative relationships between species diversity and genotypic diversity depending on environmental heterogeneity, and the conditions under which these factors are potentially relevant. Mechanistically, differential niche availability is imposed by the foundation species.     

Evidence that niche specialization explains species-energy relationships in lake fish communities

1. Interspecific niche differences have long been identified as a major explanation for the occurrence of species-rich communities. However, much fieldwork studying variation in local species richness has focused upon physical habitat attributes or regional factors, such as the size of the regional species pool. 2. We applied indices of functional diversity and niche overlap to data on the species niche to examine the importance of interspecific niche differentiation for species richness in French lake fish communities. We combined this information with environmental data to test generalizations of the physiological tolerance and niche specialization hypotheses for species-energy relationships. 3. We found evidence for a largely non-saturating relationship (relative to random expectation) between species richness and functional evenness (evenness of spacing between species in niche space), while functional richness (volume of niche space occupied) peaked at moderate levels of species richness and niche overlap showed an initial decrease followed by saturation. This suggests that increased niche specialization may have allowed species to coexist in the most species-rich communities. 4. We tested for evidence that increased temperature, local habitat area, local habitat diversity and immigration affected species richness via increased niche specialization. Temperature explained by far the largest amount of variation in species richness, functional diversity and niche overlap. These results, combined with the largely non-saturating species richness-functional evenness relationship, suggest that increased temperature may have permitted increased species richness by allowing increased niche specialization. 5. These results emphasize the importance of niche differences for species coexistence in species-rich communities, and indicate that the conservation of functional diversity may be vital for the maintenance of species diversity in biological communities. Our approach may be applied readily to many types of community, and at any scale, thus providing a flexible means of testing niche-based hypotheses for species richness gradients.     

Spatial structure and niche patterns of a rodent community in the south Bukhara desert (Middle Asia)

The distribution of niches in resource space and the niche patterns of a 13 species community of Middle Asian desert rodents were studied by use of discriminant function analysis Nineteen quantitative parameters of the environment (soil structure and vegetation), measured at 600 sample plots within twenty four one ha grids, were considered The first three canonical axes of resource space account for 83% of the variance, the first axis represents a general landscape gradient from sand to clay soils, the second axis reflects a gradient of increasing productivity, and the third axis reflects a gradient of increasing protectability of the environment The distribution of niches in resource space is not even, there are two distinct spatial guilds consisting of psammophilous and sclerophilous species There is a negative correlation between niche position (distance from species centroid to the center of the resource space) and maximal population biomass At the same time there are no correlations between the niche breadth and the niche position due to the absence of species with really broad niches Rodent biomass increases along the productivity axis and reaches a peak in the middle part of the substrate axis Rodent species diversity increases along the substrate axis from sand to clay soils Changes of species diversity along productivity axis have humpshaped patterns Maximum species diversity was recorded at low level of productivity on the sandy soils and shifted to intermediate levels of productivity on sandy-loam and clay soils     

Leaving uncut refuges during meadow harvesting increases the functional diversity of Orthoptera

In the past, insect species richness was high in Central European seminatural grasslands, which were characterized by low‐intensity land use. Currently, however, the hay in most of these grasslands is mechanically harvested, which negatively impacts insect biodiversity. One way to reduce this negative effect is to leave unmown patches as refuges. In the current research we evaluated the short‐term effects of leaving an unmown patch on the taxonomic and functional diversity of the Orthoptera assemblage in a meadow. We found that orthopteran species richness and abundance were significantly reduced by mowing, whether or not a patch was left uncut. In contrast, functional evenness, indicating distribution of species abundances in a niche space, was reduced by mowing only if the plot lacked an uncut refuge. Functional richness, indicating the amount of niche space occupied by species, was elevated if the plot had an uncut refuge. Larger species were negatively affected by mowing, while habitat specialists, mobile species and soil‐ovipositing species benefitted from it. We infer that the presence of an uncut patch increased the diversity of habitats available to orthopterans and maintained even distribution of species among niche space. In summary, leaving an unmown refuge in grasslands could increase the functional diversity of orthopterans, even if it does not preserve taxonomic diversity.     

Functional redundancy in ecology and conservation

Multiple studies have shown that biodiversity loss can impair ecosystem processes, providing a sound basis for the general application of a precautionary approach to managing biodiversity. However, mechanistic details of species loss effects and the generality of impacts across ecosystem types are poorly understood. The functional niche is a useful conceptual tool for understanding redundancy, where the functional niche is defined as the area occupied by a species in an n-dimensional functional space. Experiments to assess redundancy based on a single functional attribute are biased towards finding redundancy, because species are more likely to have non-overlapping functional niches in a multi-dimensional functional space. The effect of species loss in any particular ecosystem will depend on i) the range of function and diversity of species within a functional group, ii) the relative partitioning of variance in functional space between and within functional groups, and iii) the potential for functional compensation (degree of functional niche overlap) of the species within a functional group. Future research on functional impairment with species loss should focus on identifying which species, functional groups, and ecosystems are most vulnerable to functional impairment from species loss, so that these can be prioritized for management activities directed at maintaining ecosystem function. This will require a better understanding of how the organization of diversity into discrete functional groups differs between different communities and ecosystems.     

Intraspecific niche flexibility facilitates species coexistence in a competitive community with a fluctuating environment

We analyzed the role of niche usage flexibility (i.e. niche width) in promoting species coexistence in competitive communities in a one-dimensional niche space. We included two types of stochasticity, namely, a random sampling effect of community founding and environmental fluctuation. Fluctuation was further divided into two categories: niche-independent fluctuation (synchronized over the niche space) and niche-dependent fluctuation (variable among individual niche positions). In the analysis, two types of genetic and inheritance systems of individual niche position were considered, i.e. sexual reproduction with multiple loci and asexual reproduction with phenotypic plasticity. We found that niche usage flexibility promoted species diversity only under restricted situations when the environment was constant, but it generally promoted diversity when the environment fluctuated. In particular, under niche-independent fluctuation, niche usage flexibility significantly enhanced species diversity. In contrast, the analysis also predicted that when niche flexibility was constant, species diversity decreased with increasing environmental correlation between neighboring niches.     

Contrasting Effects of Intraspecific Trait Variation on Trait-Based Niches and Performance of Legumes in Plant Mixtures

Niche differentiation, assumed to be a key mechanism of species coexistence, requires that species differ in their functional traits. So far it remains unclear to which extent trait plasticity leads to niche shifts of species at higher plant diversity, thereby increasing or decreasing niche overlap between species. To analyse this question it is convenient to measure niches indirectly via the variation in resource-uptake traits rather than directly via the resources used. We provisionally call these indirectly measured niches trait-based niches. We studied shoot- and leaf-morphological characteristics in seven legume species in monoculture and multi-species mixture in experimental grassland. Legume species varied in the extent of trait variation in response to plant diversity. Trait plasticity led to significant shifts in species niches in multiple dimensions. Single-species niches in several traits associated with height growth and filling of canopy space were expanded, while other niche dimensions were compressed or did not change with plant diversity. Niche separation among legumes decreased in dimensions related to height growth and space filling, but increased in dimensions related to leaf size and morphology. The total extent of occupied niche space was larger in mixture than in the combined monocultures for dimensions related to leaf morphology and smaller for dimensions related to whole-plant architecture. Taller growth, greater space filling and greater plasticity in shoot height were positively, while larger values and greater plasticity in specific leaf area were negatively related with increased performance of species in mixture. Our study shows that trait variation in response to plant diversity shifts species niches along trait axes. Plastically increased niche differentiation is restricted to niche dimensions that are apparently not related to size-dependent differences between species, but functional equivalence (convergence in height growth) rather than complementarity (divergence in traits associated with light acquisition) explains increased performance of legumes in mixture.     

Species richness and niche space for temperate and tropical folivores

We measured structural and chemical traits of the leaves of native, broad-leaved trees in two temperate localities [southern Ontario, Canada (34 species), and Missouri (36 species)] and one tropical locality [central Panama (samples of 21 and 23 species)] to test the hypothesis that the greater diversity of tree species and herbivore species in the tropics is associated with greater resource niche space for herbivores. Variables were leaf toughness, water content, dry mass per unit area, several structural and nutritional carbohydrates, common mineral elements, including nitrogen and phosphorus, and several defensive compounds, including tannins and alkaloids. The four samples were almost fully separable by discriminant analysis on the basis of these leaf traits. Variance in log-transformed trait values among species was lowest in the most northern sample, but did not differ significantly between Missouri and Panama. Niche space, estimated as the square root of the total variance in the log-transformed variables within each locality, varied approximately as Panama = 1, Missouri = 0.8, Ontario = 0.5. Although niche space decreases towards higher latitudes, the change does not match the ca. sixfold decrease in tree species richness or the ca. fourfold decrease in Lepidopteran species richness over the latitude range of our samples. Accordingly, tropical folivore diversity is associated with greater resource niche overlap, greater niche specialization, and/or more completely filled niches, or with variation in niche dimensions not measured in this study.     

Environmental marginality and geographic range limits: a case study with Arabidopsis lyrata ssp. lyrata

Understanding the factors that govern the distribution of species is a central goal of evolutionary ecology. It is commonly assumed that geographic range limits reflect ecological niche limits and that species experience increasingly marginal conditions towards the edge of their ranges. Using spatial data and ecological niche models we tested these hypotheses in Arabidopsis lyrata. Specifically, we asked whether range limits coincide with predicted niche limits in this system and whether the suitability of sites declines towards the edge of the species’ range in North America. We further explored patterns of environmental change towards the edge of the range and asked whether genome‐wide patterns of genetic diversity decline with increasing peripherality and environmental marginality. Our results suggest that latitudinal range limits coincide with niche limits. Populations experienced increasingly marginal environments towards these limits – though patterns of environmental change were more complex than most theoretical models for range limits assume. Genomic diversity declined towards the edge of the species’ range and with increasing distance from the estimated centre of the species’ niche in environmental space, but not with the suitability of sites based on niche model predictions. Thus while latitudinal range limits in this system are broadly associated with niche limits, the link between environmental conditions and genetic diversity (and thus the adaptive potential of populations) is less clear.     

Niches versus neutrality: uncovering the drivers of diversity in a species-rich community

Ecological models suggest that high diversity can be generated by purely niche-based, purely neutral or by a mixture of niche-based and neutral ecological processes. Here, we compare the degree to which four contrasting hypotheses for coexistence, ranging from niche-based to neutral, explain species richness along a body mass niche axis. We derive predictions from these hypotheses and confront them with species body-mass patterns in a highly sampled marine phytoplankton community. We find that these patterns are consistent only with a mechanism that combines niche and neutral processes, such as the emergent neutrality mechanism. In this work, we provide the first empirical evidence that a niche-neutral model can explain niche space occupancy pattern in a natural species-rich community. We suggest this class of model may be a useful hypothesis for the generation and maintenance of species diversity in other size-structured communities.     

Is regional species diversity bounded or unbounded?

Two conflicting hypotheses have been proposed to explain large‐scale species diversity patterns and dynamics. The unbounded hypothesis proposes that regional diversity depends only on time and diversification rate and increases without limit. The bounded hypothesis proposes that ecological constraints place upper limits on regional diversity and that diversity is usually close to its limit. Recent evidence from the fossil record, phylogenetic analysis, biogeography, and phenotypic disparity during lineage diversification suggests that diversity is constrained by ecological processes but that it is rarely asymptotic. Niche space is often unfilled or can be more finely subdivided and still permit coexistence, and new niche space is often created before ecological limits are reached. Damped increases in diversity over time are the prevalent pattern, suggesting the need for a new ‘damped increase hypothesis'. The damped increase hypothesis predicts that diversity generally increases through time but that its rate of increase is often slowed by ecological constraints. However, slowing due to niche limitation must be distinguished from other possible mechanisms creating similar patterns. These include sampling artifacts, the inability to detect extinctions or declines in clade diversity with some methods, the distorting effects of correlated speciation‐extinction dynamics, the likelihood that opportunities for allopatric speciation will vary in space and time, and the role of undetected natural enemies in reducing host ranges and thus slowing speciation rates. The taxonomic scope of regional diversity studies must be broadened to include all ecologically similar species so that ecological constraints may be accurately inferred. The damped increase hypothesis suggests that information on evolutionary processes such as time‐for‐speciation and intrinsic diversification rates as well as ecological factors will be required to explain why regional diversity varies among times, places and taxa.     

Temporal variation in spatial organization of a rodent community in the southwestern Kyzylkum desert (Middle Asia)

The distribution of niches in resource space and the niche patterns of a 14-species community of Middle Asian desert rodents were studied during two years - at low and high rodent density - using discriminant function analysis Nineteen quantitative environmental parameters (soil structure and vegetation), measured in 550 plots within 22 1 -ha grids, were considered The first three canonical axes of resource space account for 72% of the variance The first two axes represent complex environmental gradients the first axis represents a general landscape gradient from sand to clay soils, the second axis reflects a gradient of in creasing productivity The third axis reflects with in-habitat environmental variation All community parameters, as well as parameters of individual species niches, were unstable between years At the same time, different parameters vary in different extent Position of niche centroids along macro-habitat axes, as. well as macrohabitat niche breadth, were relatively stable between years, but these parameters for microhabitat axis and values of niche overlap were much more variable A strong correlation between changes m relative between-habitat niche breadth and differences in average niche overlap with relative changes in species abundances indicate density dependence of these parameters Changes in niche overlap is a consequence of between-year differences in guild patterns Guild structure was pronounced at high density when the level of niche overlap was intermediate At low density, when the level of niche overlap decreased, guild structure was incon-spicous Different levels of diversity differed in their sensitivity to density changes α-diversity was relatively constant as a result of between-year stability of niche centroid positions However level of ß-diversity varied significantly between years reflecting changes in the level of niche overlap, because a decrease in niche overlap leads to an increase in the rate of species turnover     

Adaptive radiation and the evolution of nectarivory in a large songbird clade

The accumulation of exceptional ecological diversity within a lineage is a key feature of adaptive radiation resulting from diversification associated with the subdivision of previously underutilized resources. The invasion of unoccupied niche space is predicted to be a key determinant of adaptive diversification, and this process may be particularly important if the diversity of competing lineages within the area, in which the radiation unfolds, is already high. Here, we test whether the evolution of nectarivory resulted in significantly higher rates of morphological evolution, more extensive morphological disparity, and a heightened build‐up of sympatric species diversity in a large adaptive radiation of passerine birds (the honeyeaters, about 190 species) that have diversified extensively throughout continental and insular settings. We find that a large increase in rates of body size evolution and general expansion in morphological space followed an ancestral shift to nectarivory, enabling the build‐up of large numbers of co‐occurring species that vary greatly in size, compared to related and co‐distributed nonnectarivorous clades. These results strongly support the idea that evolutionary shifts into novel areas of niche space play a key role in promoting adaptive radiation in the presence of likely competing lineages.     

The effect of tree species diversity on fine-root production in a young temperate forest

The phenomenon of overyielding in species-diverse plant communities is mainly attributed to complementary resource use. Vertical niche differentiation belowground might be one potential mechanism for such complementarity. However, most studies that have analysed the diversity/productivity relationship and belowground niche differentiation have done so for fully occupied sites, not very young tree communities that are in the process of occupying belowground space. Here we used a 5–6 year old forest diversity experiment to analyse how fine-root (<2 mm) production in ingrowth cores (0–30 cm) was influenced by tree species identity, as well as the species diversity and richness of tree neighbourhoods. Fine-root production during the first growing season after the installation of ingrowth cores increased slightly with tree species diversity, and four-species combinations produced on average 94.8% more fine-root biomass than monocultures. During the second growing season, fine-root mortality increased with tree species diversity, indicating an increased fine-root turnover in species-rich communities. The initial overyielding was attributable to the response to mixing by the dominant species, Pseudotsuga menziesii and Picea abies, which produced more fine roots in mixtures than could be expected from monocultures. In species-rich neighbourhoods, P. abies allocated more fine roots to the upper soil layer (0–15 cm), whereas P. menziesii produced more fine roots in the deeper layer (15–30 cm) than in species-poor neighbourhoods. Our results indicate that, although there may be no lasting overyielding in the fine-root production of species-diverse tree communities, increasing species diversity can lead to substantial changes in the production, vertical distribution, and turnover of fine roots of individual species.     

Functional traits reveal the expansion and packing of ecological niche space underlying an elevational diversity gradient in passerine birds

Variation in species richness across environmental gradients may be associated with an expanded volume or increased packing of ecological niche space. However, the relative importance of these alternative scenarios remains unknown, largely because standardized information on functional traits and their ecological relevance is lacking for major diversity gradients. Here, we combine data on morphological and ecological traits for 523 species of passerine birds distributed across an Andes-to-Amazon elevation gradient. We show that morphological traits capture substantial variation in species dietary (75%) and foraging niches (60%) when multiple independent trait dimensions are considered. Having established these relationships, we show that the 14-fold increase in species richness towards the lowlands is associated with both an increased volume and density of functional trait space. However, we find that increases in volume contribute little to changes in richness, with most (78%) lowland species occurring within the range of trait space occupied at high elevations. Taken together, our results suggest that high species richness is mainly associated with a denser occupation of functional trait space, implying an increased specialization or overlap of ecological niches, and supporting the view that niche packing is the dominant trend underlying gradients of increasing biodiversity towards the lowland tropics.     

Dietary ecospace and the diversity of euprimates during the Early and Middle Eocene

This study examined adapoid and omomyoid euprimate dietary and body size diversity from the Eocene of North America and Europe. Estimates of body weights and shearing quotients calculated from lower molars were plotted on a coordinate graph as a representation of dietary niche space (dietary ecospace) occupied by extinct species. By computing the areas, average intertaxon distances, and average distances from the centroid of the resulting polygons, comparisons of Eocene euprimate dietary and body size diversity were made. Results indicate that euprimate dietary niche space expanded significantly in North America from the Early to Middle Eocene, and at all times during the Early and Middle Eocene, the niche space occupied by North American euprimates exceeded that of corresponding European euprimates. These results confirm that fossil euprimate diversity, as measured by diet and body size, significantly differed across biogeographic areas. There are many possible explanations as to why North American euprimates were significantly more diverse in terms of diet and body size than their European counterparts. The explanation advocated here as most responsible for the increased diversity during the Early and Middle Eocene relates to the existence and increased sampling of more ecologically diverse environments, such as basin margins in the western interior of North America. These diverse environments could have promoted biological processes that led to the generation of increased diversity in North America compared to the isolated island refugia of Western Europe during this time.     

Diversity increases biomass production for trematode parasites in snails

Increasing species diversity typically increases biomass in experimental assemblages. But there is uncertainty concerning the mechanisms of diversity effects and whether experimental findings are relevant to ecological process in nature. Hosts for parasites provide natural, discrete replicates of parasite assemblages. We considered how diversity affects standing-stock biomass for a highly interactive parasite guild: trematode parasitic castrators in snails. In 185 naturally occurring habitat replicates (individual hosts), diverse parasite assemblages had greater biomass than single-species assemblages, including those of their most productive species. Additionally, positive diversity effects strengthened as species segregated along a secondary niche axis (space). The most subordinate species--also the most productive when alone--altered the general positive effect, and was associated with negative diversity effects on biomass. These findings, on a previously unstudied consumer class, extend previous research to illustrate that functional diversity and species identity may generally both explain how diversity influences biomass production in natural assemblages of competing species.