Relatedness and resource diversity interact to influence the intensity of competition

When resource competition occurs between close relatives, the negative effects of competition are potentially amplified. However, kin selection theory predicts that natural selection should promote the evolution of mechanisms that minimize the intensity of competition between kin. Experimental tests of these hypotheses are mixed, however. Moreover, there is little consensus regarding the generality of either outcome, suggesting that the conditions important in determining the effects of competition between kin are likely complex and not fully understood. We performed two experiments using spadefoot toad tadpoles (Spea multiplicata) to evaluate the hypothesis that individuals can minimize the negative effects of exploitative competition by using alternative resources when competing with close relatives. Supporting our hypothesis, we found that only when individuals had access to alternative resources were the negative effects of competition between siblings less than between unrelated competitors. We suggest that mechanisms to lessen kin competition may be more likely to evolve in environments where alternative resources are available, and that selection to minimize exploitative competition between kin may promote the evolution of resource polyphenism. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 689–695.     

Seasonal population and individual niche dynamics in a tetra fish in the Pantanal wetlands

In seasonal tropical regions, rainfall and/or temporary floods during the wet season generally increase the abundance and diversity of food resources to many consumers as compared to the dry season. Therefore, seasonality can affect intraspecific competition and ecological opportunity, which are two important ecological mechanisms underlying population and individual niche variations. Here, we took advantage of the strong seasonality in the Pantanal wetlands to investigate how within‐ and between‐individual diet variations relate to seasonal population niche dynamics of the tetra fish Astyanax lacustris. We quantified dietary niche using gut contents and stable isotopes. Tetras had higher gut fullness and better body condition in the wet season, suggesting that competition is more intense in the dry season. The population niche was broader in the wet season due to an increase in diet divergence between individuals, in spite of potential stronger competition in the dry season. We posit that low ecological opportunity in the dry season limits the diversifying effect of intraspecific competition, constraining population niche expansion. Our results add new insights on how seasonality affects population and individual diets, indicating that intraspecific competition and ecological opportunity interact to determine temporal niche variations in seasonal environments.     

Intraspecific trait variation alters the outcome of competition in freshwater ciliates

Trait variation among heterospecific and conspecific organisms may substantially affect community and food web dynamics. While the relevance of competition and feeding traits have been widely studied for different consumer species, studies on intraspecific differences are more scarce, partly owing to difficulties in distinguishing different clones of the same species. Here, we investigate how intraspecific trait variation affects the competition between the freshwater ciliates Euplotes octocarinatus and Coleps hirtus in a nitrogen‐limited chemostat system. The ciliates competed for the microalgae Cryptomonas sp. (Cry) and Navicula pelliculosa (Nav), and the bacteria present in the cultures over a period of 33 days. We used monoclonal Euplotes and three different Coleps clones (Col 1, Col 2, and Col 3) in the experiment that could be distinguished by a newly developed rDNA‐based molecular assay based on the internal transcribed spacer (ITS) regions. While Euplotes feeds on Cry and on bacteria, the Coleps clones cannot survive on bacteria alone but feed on both Cry and Nav with clone‐specific rates. Experimental treatments comprised two‐species mixtures of Euplotes and one or all of the three different Coleps clones, respectively. We found intraspecific variation in the traits “selectivity” and “maximum ingestion rate” for the different algae to significantly affect the competitive outcome between the two ciliate species. As Nav quickly escaped top‐down control and likely reached a state of low food quality, ciliate competition was strongly determined by the preference of different Coleps clones for Cry as opposed to feeding on Nav. In addition, the ability of Euplotes to use bacteria as an alternative food source strengthened its persistence once Cry was depleted. Hence, trait variation at both trophic levels codetermined the population dynamics and the outcome of species competition.     

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.

     

Nonlinear effect of density on trophic niche width and between‐individual variation in diet in a neotropical cichlid

Density‐dependent dynamics have considerable effects in many ecological processes and patterns that characterize natural populations. In the present study, we aim at evaluating the effect of density on the diet width and between‐individual variability in nine different pike cichlid Crenicichla lepidota populations dwelling in floodplain lagoons. Our results indicated that low‐density populations exhibit small diet breadth as well as small between‐individual variability in their diet. In addition, these populations were characterized by a nested diet pattern, where the diet of specialist individuals represented a subset of the food items consumed by generalist individuals. Populations with intermediate densities had a larger populational diet breadth while the individual diet breadth remained unchanged. This pattern was due to an increase in between‐individual variability in diet, which consequently decreases the diet overlap among individuals and thus lower diet nestedness. Finally, under high densities, the niche width at the population level decreased because of lower between‐individual variability and higher diet overlap. Together, these results showed that niche width exhibits a non‐linear function with density. At first, an increase in density increased the niche width because of greater between‐individual diet variability. However, after a threshold density value, the effect was reversed, and the niche width decreased because of a higher diet overlap among individuals.     

Intraspecific genetic variation in Paramecium revealed by mitochondrial cytochrome C oxidase I sequences

Studies of intraspecific genetic diversity of ciliates, such as population genetics and biogeography, are particularly hampered by the lack of suitable DNA markers. For example, sequences of the non-coding ribosomal internal transcribed spacer (ITS) regions are often too conserved for intraspecific analyses. We have therefore identified primers for the mitochondrial cytochrome c oxidase I (COI) gene and applied them for intraspecific investigations in Paramecium caudatum and Paramecium multimicronucleatum. Furthermore, we obtained sequences of the ITS regions from the same strains and carried out comparative sequence analyses of both data sets. The mitochondrial sequences revealed substantially higher variation in both Paramecium species, with intraspecific divergences up to 7% in P. caudatum and 9.5% in P. multimicronucleatum. Moreover, an initial survey of the population structure discovered different mitochondrial haplotypes of P. caudatum in one pond, thereby demonstrating the potential of this genetic marker for population genetic analyses. Our primers successfully amplified the COI gene of other Paramecium. This is the first report of intraspecific variation in free-living protozoans based on mitochondrial sequence data. Our results show that the high variation in mitochondrial DNA makes it a suitable marker for intraspecific and population genetic studies.     

Intraspecific trait variation across elevation predicts a widespread tree species' climate niche and range limits

Global change is widely altering environmental conditions which makes accurately predicting species range limits across natural landscapes critical for conservation and management decisions. If climate pressures along elevation gradients influence the distribution of phenotypic and genetic variation of plant functional traits, then such trait variation may be informative of the selective mechanisms and adaptations that help define climatic niche limits. Using extensive field surveys along 16 elevation transects and a large common garden experiment, we tested whether functional trait variation could predict the climatic niche of a widespread tree species (Populus angustifolia) with a double quantile regression approach. We show that intraspecific variation in plant size, growth, and leaf morphology corresponds with the species' total climate range and certain climatic limits related to temperature and moisture extremes. Moreover, we find evidence of genetic clines and phenotypic plasticity at environmental boundaries, which we use to create geographic predictions of trait variation and maximum values due to climatic constraints across the western US. Overall, our findings show the utility of double quantile regressions for connecting species distributions and climate gradients through trait‐based mechanisms. We highlight how new approaches like ours that incorporate genetic variation in functional traits and their response to climate gradients will lead to a better understanding of plant distributions as well as identifying populations anticipated to be maladapted to future environments.     

Intraspecific competition reduces niche width in experimental populations

Intraspecific competition is believed to drive niche expansion, because otherwise suboptimal resources can provide a refuge from competition for preferred resources. Competitive niche expansion is well supported by empirical observations, experiments, and theory, and is often invoked to explain phenotypic diversification within populations, some forms of speciation, and adaptive radiation. However, some foraging models predict the opposite outcome, and it therefore remains unclear whether competition will promote or inhibit niche expansion. We conducted experiments to test whether competition changes the fitness landscape to favor niche expansion, and if competition indeed drives niche expansion as expected. Using Tribolium castaneum flour beetles fed either wheat (their ancestral resource), corn (a novel resource) or mixtures of both resources, we show that fitness is maximized on a mixed diet. Next, we show that at higher population density, the optimal diet shifts toward greater use of corn, favoring niche expansion. In stark contrast, when beetles were given a choice of resources, we found that competition caused niche contraction onto the ancestral resource. This presents a puzzling mismatch between how competition alters the fitness landscape, versus competition's effects on resource use. We discuss several explanations for this mismatch, highlighting potential reasons why optimality models might be misleading.     

The roles of morphological traits,resource variation and resource partitioning associated with the dietary niche expansion in the fish‐eating bat Myotis pilosus

Niche expansion and shifts are involved in the response and adaptation to environmental changes. However, it is unclear how niche breadth evolves and changes toward higher‐quality resources. Myotis pilosus is both an insectivore and a piscivore. We examined the dietary composition and seasonality in M. pilosus and the closely related Myotis fimbriatus using next‐generation DNA sequencing. We tested whether resource variation or resource partitioning help explain the dietary expansion from insects to fish in M. pilosus. While diet composition and diversity varied significantly between summer and autumn, the proportion of fish‐eating individuals did not significantly change between seasons in M. pilosus. Dietary overlap between M. pilosus and M. fimbriatus during the same seasons was much higher than within individual species across seasons. We recorded a larger body size, hind foot length, and body mass in M. pilosus than in M. fimbriatus and other insectivorous trawling bats from China. Similar morphological differences were found between worldwide fishing bats and nonfishing trawling bats. Our results suggest that variation in insect availability or interspecific competition may not play important roles in the dietary expansion from insects to fish in M. pilosus. Myotis pilosus has morphological advantages that may help it use fish as a diet component. The morphological advantage promoting dietary niche evolution toward higher quality resources may be more important than variation in the original resource and the effects of interspecific competition.     

Microbial generalist or specialist: Intraspecific variation and dormancy potential matter

Microbial generalists and specialists coexist in the soil environment while having distinctive impacts on microbial community dynamics. In microbial ecology, the underlying mechanisms as to why a species is a generalist or a specialist remain ambiguous. Herein, we collected soils across a national scale and identified bacterial generalists and specialists according to niche breadth at the species level (OTU level), and the single-nucleotide differences in each species were measured to investigate intraspecific variation (at zero-radius OTU level). Compared with that of the specialists, the intraspecific variation of the generalists was much higher, which ensured their wider niche breadth and lower variability. The higher asynchrony and different niche preferences of conspecific individuals and the higher dormancy potential within the generalists further contributed to their stability in varying environments. Besides, generalists were less controlled by environmental filtering, which was indicated by the stronger signature of stochastic processes in their assembly, and had higher diversification and transition rates that allowed them to adapt to environmental changes to a greater extent than specialists. Overall, this study provides a new comprehensive understanding of the rules of assembly and the evolutionary roles of bacterial generalists and specialists. It also highlights the importance of intraspecific variation and the dormancy potential in the stability of species.     

Climatic‐niche evolution with key morphological innovations across clades within Scutiger boulengeri (Anura: Megophryidae)

The studies of climatic‐niche shifts over evolutionary time accompanied by key morphological innovations have attracted the interest of many researchers recently. We applied ecological niche models (ENMs), ordination method (environment principal component analyses; PCA‐env), combined phylogenetic comparative methods (PCMs), and phylogenetic generalized least squares (PGLS) regression methods to analyze the realized niche dynamics and correspondingly key morphological innovations across clades within Scutiger boulengeri throughout their distributions in Qinghai–Tibet Plateau (QTP) margins of China. Our results show there are six clades in S. boulengeri and obvious niche divergences caused by niche expansion in three clades. Moreover, in our system, niche expansion is more popular than niche unfilling into novel environmental conditions. Annual mean temperature, annual precipitation, and precipitation of driest month may contribute to such a shift. In addition, we identified several key climatic factors and morphological traits that tend to be associated with niche expansion in S. boulengeri clades correspondingly. We found phenotypic plasticity [i.e., length of lower arm and hand (LAHL), hind‐limb length (HLL), and foot length (FL)] and evolutionary changes [i.e., snout–vent length (SVL)] may together contribute to niche expansion toward adapting novel niche, which provides us a potential pattern of how a colonizing toad might seed a novel habitat to begin the process of speciation and finally adaptive radiation. For these reasons, persistent phylogeographic divisions and accompanying divergences in niche occupancy and morphological adaption suggest that for future studies, distinct genetic structure and morphological changes corresponding to each genetic clade should be included in modeling niche evolution dynamics, but not just constructed at the species level.     

Resource-based models of competitive interactions. I. Intraspecific competition in Ratibida columnifera ( Asteraceae)

We constructed and tested a series of regression models of intraspecific competition in Ratibida columnifera (Asteraceae), based on the growth and water use of individual plants. Models were constructed from a set of plants ("model") grown without competition under three watering regimes. Each model was then tested on another set of plants ("test") grown, singly or in pairs, under two watering regimes, one of them different than those of the "model" plants. Both sets of plants were grown simultaneously. Models that used only estimates of plant dry mass (ISON [interval by interval size only], and SON [size only]) were outperformed (i.e., the difference between predicted values of final dry mass and their true values were larger on average) by models that incorporated direct measures of water uptake and usage (ISAW [interval by interval size and water], WON [water only], SAW [size and water]). Harvest biomass predictions given by these three last models deviated from values of true biomass by an average of only 6.1%.     

大别山五针松种内和种间竞争强度

通过对安徽省岳西县大别山五针松群落内的53株对象木及2079株竞争木的调查,运用Hegyi单木竞争模型分析了大别山五针松的种内和种间竞争强度。结果表明,大别山五针松的种内和种间竞争强度分别为17.11%和82.89%,说明竞争主要来自种间。大别山五针松的伴生种较多,种内与主要伴生种间的竞争关系为短柄枹大别山五针松种内黄山松满山红茅栗金缕梅紫茎四照花灯台树鹅耳枥。竞争强度随对象木胸径的增大而减小,当对象木的胸径小于25cm时,所受到的竞争强度较大;当胸径在25cm以上时,竞争强度变化很小,二者符合幂函数关系(CI=AD-B),所得的预测模型能很好的预测大别山五针松种内和种间竞争强度。     

Seasonality and interspecific competition shape individual niche variation in co‐occurring tetra fish in Neotropical streams

The drivers of intraspecific niche variation and its effects on species interactions are still unclear, especially in species‐rich Neotropical environments. Here, we investigated how ecological opportunity and interspecific competition affect the degree of individual trophic specialization and the population niche breadth in tetra fish. We studied the four ecologically similar species (Psalidodon aff. gymnodontus, P. aff. paranae, P. bifasciatus, and Bryconamericus ikaa) in subtropical headwater streams (three sites with two co‐occurring species and three sites with only one species). We sampled fish in two contrasting seasons (winter/dry and summer/wet) and quantified their trophic niches using gut content analysis. Psalidodon bifasciatus was the only species distributed over all the sampled streams. We observed seasonal differences in population trophic niche breadth of P. bifasciatus just when this species co‐occurred with P. aff. gymnodontus. These findings confirm the complex nature of the effects of interspecific competition, depending, for instance, on the identity of the competitor. The degree of individual specialization of P. bifasciatus was higher in the winter, and it was not influenced by the presence of another species. Conversely, the other two Psalidodon species studied presented greater individual specialization in the summer, when fish consumed a higher proportion of allochthonous items (terrestrial insects and seeds), and there were no effects only for B. ikaa. Herein, our results suggest that seasonality in food‐resource availability is a major driver of niche variation and it has the potential to play an important role in how these similar tetra species interact and coexist. Abstract in Portuguese is available with online material.     

天然东北红豆杉(Taxus cuspidata)种内和种间竞争

竞争是植物种内和种间关系的主要形式之一。通过对黑龙江穆棱东北红豆杉自然保护区的95株东北红豆杉对象木及980株竞争木的调查,运用Hegyi的单木竞争指数计算分析了东北红豆杉的种内和种间竞争强度。东北红豆杉的种内竞争强度不大,占总竞争的4%。竞争压力更多的来自于种间竞争,占总竞争的96%。与东北红豆杉竞争激烈的树种主要是冷杉、紫椴、色木槭和红松等地带性植被的优势种。随着东北红豆杉胸径的增大,所受到的竞争压力逐渐减小,胸径在20cm以前所受到的竞争压力最大,竞争强度与对象木胸径符合幂函数（CI=AD^-B）关系。     

Tolerance niche expansion and potential distribution prediction during Asian openbill bird range expansion

It is prevalent to use ecological niche models in the analysis of species expansion and niche changes. However, it is difficult to estimate the niche when alien species fail to establish in exotic areas. Here, we applied the tolerance niche concept, which means that niche of species can live and grow but preclude a species from establishing self‐sustaining populations, in such fail‐to‐establish events. Taking the rapidly expanded bird, Asian openbill (Anastomus oscitans), as a model species, we investigated niche dynamics and its potential effects on the population by Niche A and ecospat, predicted potential distribution by biomod2. Results showed that niche expansion has occurred in two non‐native populations caused by the tolerance of colder and wetter environments, and potential distribution mainly concentrated on equatorial islands. Our study suggested that the expanded niche belongs to tolerance niche concept according to the populations'' dynamics and GPS tracking evidence. It is essential to consider source populations when we analyze the alien species. We recommended more consideration to the application of tolerance niche in alien species research, and there is still a need for standard measurement frameworks for analyzing the tolerance niche.     

Intraspecific trait variation across scales: implications for understanding global change responses

Recognition of the importance of intraspecific variation in ecological processes has been growing, but empirical studies and models of global change have only begun to address this issue in detail. This review discusses sources and patterns of intraspecific trait variation and their consequences for understanding how ecological processes and patterns will respond to global change. We examine how current ecological models and theories incorporate intraspecific variation, review existing data sources that could help parameterize models that account for intraspecific variation in global change predictions, and discuss new data that may be needed. We provide guidelines on when it is most important to consider intraspecific variation, such as when trait variation is heritable or when nonlinear relationships are involved. We also highlight benefits and limitations of different model types and argue that many common modeling approaches such as matrix population models or global dynamic vegetation models can allow a stronger consideration of intraspecific trait variation if the necessary data are available. We recommend that existing data need to be made more accessible, though in some cases, new experiments are needed to disentangle causes of variation.     

Plant responses to climatic extremes: within‐species variation equals among‐species variation

Within‐species and among‐species differences in growth responses to a changing climate have been well documented, yet the relative magnitude of within‐species vs. among‐species variation has remained largely unexplored. This missing comparison impedes our ability to make general predictions of biodiversity change and to project future species distributions using models. We present a direct comparison of among‐ versus within‐species variation in response to three of the main stresses anticipated with climate change: drought, warming, and frost. Two earlier experiments had experimentally induced (i) summer drought and (ii) spring frost for four common European grass species and their ecotypes from across Europe. To supplement existing data, a third experiment was carried out, to compare variation among species from different functional groups to within‐species variation. Here, we simulated (iii) winter warming plus frost for four grasses, two nonleguminous, and two leguminous forbs, in addition to eleven European ecotypes of the widespread grass Arrhenatherum elatius. For each experiment, we measured: (i) C/N ratio and biomass, (ii) chlorophyll content and biomass, and (iii) plant greenness, root ¹⁵N uptake, and live and dead tissue mass. Using coefficients of variation (CVs) for each experiment and response parameter, a total of 156 within‐ vs. among‐species comparisons were conducted, comparing within‐species variation in each of four species with among‐species variation for each seed origin (five countries). Of the six significant differences, within‐species CVs were higher than among‐species CVs in four cases. Partitioning of variance within each treatment in two of the three experiments showed that within‐species variability (ecotypes) could explain an additional 9% of response variation after accounting for the among‐species variation. Our observation that within‐species variation was generally as high as among‐species variation emphasizes the importance of including both within‐ and among‐species variability in ecological theory (e.g., the insurance hypothesis) and for practical applications (e.g., biodiversity conservation).     

濒危植物峨眉含笑的种内、种间竞争

研究植物种内、种间竞争关系是探究植物濒危原因的重要方式之一, 根据竞争来源和竞争预测模型可以制定具有针对性的保护策略。本文以雅安周公山峨眉含笑(Michelia wilsonii)野生种群为研究对象, 使用逐步扩大范围法确定峨眉含笑的竞争范围半径, 运用Hegyi单木竞争模型计算竞争指数(competition index, CI), 分析其种内、种间竞争关系。结果表明: 峨眉含笑的最适竞争范围半径是10 m, 能较好地反映其种内竞争强度; 峨眉含笑的竞争压力主要来自种内, 种内竞争指数(348.72, 62.52%)远大于种间竞争指数(209.03, 37.48%); 小树、中树阶段个体的竞争强度较大, 平均竞争指数(3.97、3.14)远高于总体平均竞争指数(2.68); 内有21种竞争木, 其中杉木(Cunninghamia lanceolata)、华中樱桃(Prunus conradinae)、细刺枸骨(Ilex hylonoma)是峨眉含笑的主要竞争树种; 胸径与竞争指数间服从指数函数关系(CI = 3.8907e^-0.048x, R² = 0.1087, P < 0.01), 随着对象木胸径的增大, 竞争指数不断降低, 当胸径达到30 cm后, 竞争强度基本稳定。综上, 小树、中树阶段的峨眉含笑个体受到极强的种内竞争, 初入老树阶段的个体受到较强的种间竞争。为更好地保护峨眉含笑的天然种群, 降低竞争对种群更替的影响, 在林分管理中需要促进小树、中树的个体更新, 减轻植株间的竞争消耗, 加速峨眉含笑的生长和维持种群稳定。     

Ecological release exposes genetically based niche variation

The evolutionary trajectories of ecological niches have profound impacts on community, population and speciation dynamics, yet the underlying causes of niche lability vs. stasis are poorly understood. Here, we conducted a field experiment to quantify the effects of competition and, conversely, competitive release on the microevolutionary processes driving microhabitat niche evolution in an annual plant population restricted to California vernal pool wetlands. Removing competitors generated a strong increase in mean fitness, the exposure of genetically based niche variation and directional selection for niche evolution in the experimental population. In contrast, genetic variation in the microhabitat niche and directional selection for niche evolution were not detected in individuals growing with competitors. These results indicate that ecological opportunity (here, the removal of competitors) can trigger the immediate expression of latent, heritable niche variation that is necessary for rapid evolutionary responses; conversely, competitors may restrict niche evolution, contributing to niche conservatism in saturated communities.