Determining trophic niche width: an experimental test of the stable isotope approach

Determining the trophic niche width of an animal population and the relative degree to which a generalist population consists of dietary specialists are long‐standing problems of ecology. It has been proposed that the variance of stable isotope values in consumer tissues could be used to quantify trophic niche width of consumer populations. However, this promising idea has not yet been rigorously tested. By conducting controlled laboratory experiments using model consumer populations (Daphnia sp., Crustacea) with controlled diets, we investigated the effect of individual‐ and population‐level specialisation and generalism on consumer δ¹³C mean and variance values. While our experimental data follow general expectations, we extend current qualitative models to quantitative predictions of the dependence of isotopic variance on dietary correlation time, a measure for the typical time over which a consumer changes its diet. This quantitative approach allows us to pinpoint possible procedural pitfalls and critical sources of measurement uncertainty. Our results show that the stable isotope approach represents a powerful method for estimating trophic niche widths, especially when taking the quantitative concept of dietary correlation time into account.     

Seasonal variation in the trophic structure of a spatial prey subsidy linking aquatic and terrestrial food webs: adult aquatic insects

Research over the past decade has established spatial resource subsidies as important determinants of food web dynamics. However, most empirical studies have considered the role of subsidies only in terms of magnitude, ignoring an important property of subsidies that may affect their impact in recipient food webs: the trophic structure of the subsidy relative to in situ resources. This may be especially important when subsidies are composed of organisms, as opposed to nutrient subsidies, because the trophic position of subsidy organisms may differ from in situ prey. I explored the relative magnitude and trophic structure of a cross-habitat prey subsidy, adult aquatic insects, in terrestrial habitats along three streams in the south–central United States. Overall, adult aquatic insects contributed more than one-third of potential insect prey abundance and biomass to the terrestrial habitat. This contribution peaked along a permanent spring stream, reaching as high as 94% of abundance and 86% of biomass in winter. Trophic structure of adult aquatic and terrestrial insects differed. Nearly all adult aquatic insects were non-consumers as adults, whereas all but one taxon of terrestrial insects were consumers. Such a difference created a strong relationship between the relative contribution of the prey subsidy and the trophic structure of the prey assemblage: as the proportion of adult aquatic insects increased, the proportion of consumers in the prey assemblage declined. Specific effects varied seasonally and with distance from the stream as the taxonomic composition of the subsidy changed, but general patterns were consistent. These findings show that adult aquatic insect subsidies to riparian food webs not only elevate prey availability, but also alter the trophic structure of the entire winged insect prey assemblage.     

基于稳定同位素方法的闽江口常见鱼类群落营养结构的季节性变化

为了解闽江口常见鱼类群落的营养结构特征,于2015年冬季(1月)、春季(5月)、夏季(8月)、秋季(11月)采集闽江口底拖网渔业资源样品,利用稳定同位素分析闽江口常见鱼类群落营养结构特征及其季节性变化,并计算了基于δ¹³C-δ¹⁵N量化的7个营养结构的群落范围指标.结果表明: 闽江口常见鱼类群落除夏季和秋季的δ¹³C值之间不存在显著性差异,其余各季节之间δ¹³C值均差异显著;δ¹⁵N值春季与其余3个季节都表现出显著性差异,而其余3个季节δ¹⁵N值互相之间均无显著性差异.稳定同位素的量化指标表明: 闽江口常见鱼类群落的营养结构存在季节差异,春季的生态位总空间(TA)、核心生态位空间(SEAc)、鱼类基础食物来源(CR)、营养长度(NR)、鱼类营养多样性(CD)均为四季的最大值;从夏季至秋季再到冬季,生态位总空间(TA)、核心生态位空间(SEAc)、鱼类基础食物来源(CR)、营养长度(NR)、鱼类营养多样性(CD)和营养位置差异(MNND)均逐步上升.闽江口常见鱼类群落营养结构的季节差异可能与鱼类生殖洄游和群落内食源多样性的变化密切相关.     

The importance of terrestrial resource subsidies for shredders in open‐canopy streams revealed by stable isotope analysis

1. Allochthonous detritus is the major source of energy in forested streams, but less is known of the importance of terrestrial subsidies to open‐canopy streams. Here, we used stable isotope analysis to assess the importance of allochthonous versus autochthonous energy sources to invertebrate shredders in four open‐canopy streams in Sweden. Shredders and potential food sources were analysed at both open sites and those with deciduous trees in the riparian zone. 2. Mixing models showed that allochthonous coarse particulate organic matter was the most important energy source to shredders at both the open and wooded sites, suggesting that terrestrial subsidies may be an important process in open‐canopy streams, just as they are in forested streams. 3. However, shredders at open sites had a larger proportion of biofilm in their diet than at wooded sites, indicating an ability of shredders to adjust their diet with food availability. 4. We also used the carbon and nitrogen stable isotope signatures to assess the size of the feeding niche of shredders. Feeding on both allochthonous and autochthonous sources at open sites was reflected in a larger feeding niche than at wooded sites for one of the three species analysed. There was substantial overlap of the feeding niche among shredder species, indicating a high functional redundancy within this guild.     

Trophic niche width and overlap of two sympatric gulls in the southwestern mediterranean

The diets of two potential competitor species, Audouin's Larus audouinii and yellow-legged gulls Larus cachinnans, were examined while they bred at the Chafarinas Islands during 1993, 1994, and 1995. Data were collected during two commercial fishing regimes: (1) trawling and purse seine fisheries, and (2) diurnal trawlers only. Since the food supply for the gulls in this area was heavily reliant on the activity of purse seine fisheries, these contrasting situations allowed us to analyze short-term effects, induced by daily changes in food supply, on niche width, dietary shift, and niche overlap between the two species. Overall, both species relied mainly on fish for food, especially Clupeiforms, in the case of Audouin's gull irrespective of the fishing situation, and in the case of the yellow-legged gull, only when purse seine fishing was in operation. When purse seine boats did not operate (food shortage), yellow-legged gulls broadened their niche, consuming equal amounts of all the feeding resources, and they showed a dietary shift toward a greater consumption of prey from refuse tips. In contrast, Audouin's gulls did not change their niche width, but showed a slight dietary shift away from the consumption of epipelagic fish, compensated by an increase in reliance on benthic-mesopelagic resources. Niche overlap was clearly higher on days when both fishing fleets operated, probably because a superabundant food resource facilitates high overlap without affecting coexistence between the two species. Since our study was developed on the basis of daily variations in food supply, and competition effects are to be expected on a longer-term basis, these changes can be seen as the outcome of the coexistence of two species in stable competitive equilibrium. Received: 4 October 1996 / Accepted: 20 May 1997     

Stable isotope analyses reveal dense trophic species packing and clear niche differentiation in a malagasy primate community

Understanding the mechanisms maintaining local species richness is a major topic in tropical ecology. In ecological communities of Madagascar, primates represent a major part of mammalian diversity and, thus, are a suitable taxon to study these mechanisms. Previous research suggested that ecological niche differentiation facilitates the coexistence of lemurs. However, detailed data on all species making up diverse local primate assemblages is rarely available, hampering community‐wide tests of niche differentiation among Malagasy mammals. Here, we took an indirect approach and used stable isotopes as long‐term indicators of individuals' diets to answer the question of whether trophic patterns and food‐related mechanisms stabilize coexistence in a species‐rich lemur community. We analyzed stable carbon and nitrogen isotopes in hair collected from eight syntopic lemurs in Kirindy Forest. We found that lemur species were well separated into trophic niches and ranged over two trophic levels. Furthermore, species were densely packed in isotopic space suggesting that past competitive interactions between species are a major structuring force of this dry forest lemur community. Results of other comparative studies on primates and our findings underline that—in contrast to communities worldwide—the structure and composition of lemur communities follow predictions of ecological niche theory. Patterns of competitive interactions might be more clearly revealed in Malagasy primate communities than elsewhere because lemurs represent a large fraction of ecologically interacting species in these communities. The pronounced trophic niche differentiation among lemurs is most likely due to intense competition in the past as is characteristic for adaptive radiations. Am J Phys Anthropol 153:249–259, 2014. © 2013 Wiley Periodicals, Inc.     

Niche width and niche overlap: a method based on type-2 fuzzy sets

Complicated ecosystems and the high non-linearity of evolution has made biology more adaptable to a variety of environments. The relationship between life and environment demands a dynamic definition of niche and its measurement. In this paper we propose a model of niche with dynamic character based on the “broad band” effect in type-2 fuzzy sets. The niche in this definition is an interval in each ecological dimension which is dynamic in character and depends on the actual environment. We also give formulas for niche width and niche overlap. We compute the niche width and overlap for plants and animals and then compare these results with previous results. The results for niche width in this paper reflect the diversity of resources used by species or communities. The results for niche overlap demonstrate overlap under different environmental conditions. The results are, moreover, intervals, which could provide more information. The model in this paper could therefore be used to describe the state of every resource comprehensively, reflecting the interaction between species and environment.     

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.     

Fifty years later: trophic ecology and niche overlap of a native and non-indigenous fish species in the western basin of Lake Erie

Since the introduction of white perch (Morone americana) into Lake Erie over 50 years ago, the population size of native yellow perch (Perca flavescens) has decreased up to 79 % and significant changes to the ecosystem have occurred. We examined long-term population estimates and used stable isotopes of carbon (δ ¹³C) and nitrogen (δ ¹⁵N) paired with stomach content analysis to quantify the trophic ecology and niche overlap of adult yellow perch and white perch in the western basin of Lake Erie. We found that changes in yellow perch abundance since 1979 appeared to be better correlated with changes in fishery exploitation rates than with food competition effects from white perch. At the time of this study, yellow perch were found to have higher δ ¹³C values, indicating greater utilization of benthic food resources than white perch, and white perch occupied higher trophic positions based on δ ¹⁵N. The diets of both species varied spatially and seasonally based on stable isotopes and stomach contents, likely driven by changes in prey abundance. Comparison of niche widths using stable isotope population metrics and Schoener diet similarity index suggested a low to moderate degree of niche overlap between species. Isotopic niches of white perch were generally larger than those of yellow perch demonstrating broader resource utilization by this non-indigenous species. We submit that isotopic niche overlap comparisons are more appropriate for studies seeking to understand interactions among populations over course temporal scales, while diet overlap indices, such as the Schoener index provide a means to study fine-scale interactions such as ontogenetic and seasonal diet shifts.     

Climatic niche evolution in turtles is characterized by phylogenetic conservatism for both aquatic and terrestrial species

Understanding how the climatic niche of species evolved has been a topic of high interest in current theoretical and applied macroecological studies. However, little is known regarding how species traits might influence climatic niche evolution. Here, we evaluated patterns of climatic niche evolution in turtles (tortoises and freshwater turtles) and whether species habitat (terrestrial or aquatic) influences these patterns. We used phylogenetic, climatic and distribution data for 261 species to estimate their climatic niches. Then, we compared whether niche overlap between sister species was higher than between random species pairs and evaluated whether niche optima and rates varied between aquatic and terrestrial species. Sister species had higher values of niche overlap than random species pairs, suggesting phylogenetic climatic niche conservatism in turtles. The climatic niche evolution of the group followed an Ornstein–Uhlenbeck model with different optimum values for aquatic and terrestrial species, but we did not find consistent evidence of differences in their rates of climatic niche evolution. We conclude that phylogenetic climatic niche conservatism occurs among turtle species. Furthermore, terrestrial and aquatic species occupy different climatic niches but these seem to have evolved at similar evolutionary rates, reinforcing the importance of habitat in understanding species climatic niches and their evolution.     

Predator and detritivore niche width helps to explain biocomplexity of experimental detritus-based food webs in four aquatic and terrestrial ecosystems

In the study of food webs, the existence and explanation of recurring patterns, such as the scale invariance of linkage density, predator–prey ratios and mean chain length, constitute long-standing issues. Our study focused on litter-associated food webs and explored the influence of detritivore and predator niche width (as δ¹³C range) on web topological structure. To compare patterns within and between aquatic and terrestrial ecosystems and take account of intra-habitat variability, we constructed 42 macroinvertebrate patch-scale webs in four different habitats (lake, lagoon, beech forest and cornfield), using an experimental approach with litterbags. The results suggest that although web differences exist between ecosystems, patterns are more similar within than between aquatic and terrestrial web types. In accordance with optimal foraging theory, we found that the niche width of predators and prey increased with the number of predators and prey taxa as a proportion of total taxa in the community. The tendency was more marked in terrestrial ecosystems and can be explained by a lower per capita food level than in aquatic ecosystems, particularly evident for predators. In accordance with these results, the number of links increased with the number of species but with a significantly sharper regression slope for terrestrial ecosystems. As a consequence, linkage density, which was found to be directly correlated to niche width, increased with the total number of species in terrestrial webs, whereas it did not change significantly in aquatic ones, where connectance scaled negatively with the total number of species. In both types of ecosystem, web robustness to rare species removal increased with connectance and the niche width of predators. In conclusion, although limited to litter-associated macroinvertebrate assemblages, this study highlights structural differences and similarities between aquatic and terrestrial detrital webs, providing field evidence of the central role of niche width in determining the structure of detritus-based food webs and posing foraging optimisation constraints as a general mechanistic explanation of food web complexity differences within and between ecosystem types.     

Seasonal modifications of longitudinal distribution patterns within a stream: Interspecific interactions in the niche overlap zones of two Ephemera mayflies

Niche differentiation between closely related species leads to differentiation of their habitats. Segregation based on slight differences in environmental factors, that is niche differentiation on the microhabitat scale, allows more species to inhabit a certain geographic space. Therefore, such fine scale niche differentiation is an important factor in the support of species diversity. In addition, niche differentiation on the microhabitat scale and/or the differentiation of breeding seasons can be considered typical mechanisms that facilitate multispecies’ co‐existence. In this study, sister species (Commonly, Ephemera japonica inhabit at upstream region and Ephemera strigata inhabit at middle stream region), which often coexist in the upper to middle reaches of river systems of the Japanese Islands, were targeted and the following aspects were investigated. First, differences in habitat preference and interspecific differences in flow distribution patterns on a geographically fine scale were tracked in detail. Subsequently, the temporal transitions of their distribution patterns were investigated in detail and seasonal changes were investigated. Finally, we thoroughly investigated the disappearance of nymphs of each species from the river due to emergence affected the distribution of each species (by conducting daily emergence surveys). Combining results of these multiple studies also suggested that there may be spatiotemporal interspecial interaction between these two species within/around their overlapping regions. Traditionally, the longitudinal distribution pattern of these two Ephemera mayflies has been thought to be established based on a difference in habitat preferences, but this study revealed that the interspecific interaction between the two species also plays an important role. This study provides new insights into species diversity and distribution pattern formation in river‐dwelling species.     

Habitat overlap/niche segregation in two Umbilicaria lichens: a possible mechanism

Summary Umbilicaria muhlenbergii and Umbilicaria papulosa are two similar lichens which show extensive habitat overlap. Current theory predicts that for those resources that are limited, and of critical importance to both species, segregation is demand for or use of those resources ought to be present. Controlled experiments showed that the upper and lower cortical surfaces of the two species functioned in very different ways so that U. muhlenbergii is wetted most rapidly by runoff and mist-like precipitation whereas U. papulosa is wetted most rapidly by large raindrops. An analysis of the meteorological conditions in the natural habitat for the two lichens shows that the four months providing significant productivity in U. papulosa are the same months with a very high frequency of thunderstorms (which produce large drops). Conversely, the months which produce the greatest production in U. muhlenbergii are spread out over most of the year. When water is supplied after periods of dryness, the two species absorb it at different times and from different sources. This difference supports current ideas about niche segregation in plants but represents the first time it has been demonstrated for lichens.     

Patterns of resource use and isotopic niche overlap among three species of sharks occurring within a protected subtropical estuary

Predation is one of the most fundamental and unifying concepts in ecology, and we are beginning to obtain a more complete understanding of how predators drive community structure and ecosystem function through their impacts on prey. We know considerably less about how predators affect each other through intraguild interactions, which is surprising considering predators often occur simultaneously and may compete for resources while avoiding being killed themselves. In the present study, we examined aspects of inter- and intra-specific resource use among three species of large-bodied predatory sharks (blacktip, bull, lemon) co-occurring within a subtropical, protected bay in the southeastern USA. Specifically, we inferred relative trophic position, isotopic niche overlap, and patterns of resource use of sharks using stable isotope analysis of carbon-13 and nitrogen-15 from blood and fin cartilage samples. We also combined these approaches with estimates of abundance and occurrence from empirical shark surveys to consider whether these species may exhibit resource partitioning in space and time. We found that all three species overlapped in space, and there was some isotopic niche overlap between the species. We also found evidence of temporal isotopic niche stability, suggesting that co-occurring shark species may compete for available prey resources, but individuals of those species may have similar patterns of resource use over time. We discuss our findings as they relate to the ecologies of the species in question and how sound conservation and management of ecosystems can allow for predator diversity, sympatry, and stable use of resources at the top of the food chain.     

Spatial variation in the strength of mutualism between a jumping spider and a terrestrial bromeliad: Evidence from the stable isotope N

Psecas chapoda, a neotropical jumping spider strictly associated with the terrestrial bromeliad Bromelia balansae in cerrados and semi-deciduous forests in South America, effectively contributes to plant nutrition and growth. In this study, our goal was to investigate if spider density caused spatial variations in the strength of this spider–plant mutualism. We found a positive significant relationship between spider density and δ¹⁵N values for bromeliad leaves in different forest fragments. Open grassland Bromeliads were associated with spiders and had higher δ¹⁵N values compared to forest bromeliads. Although forest bromeliads had no association with spiders their total N concentrations were higher. These results suggest that bromeliad nutrition is likely more litter-based in forests and more spider-based in open grasslands. This study is one of the few to show nutrient provisioning and conditionality in a spider–plant system.     

Trophic niche width,offspring condition and immunity in a raptor species

Strategies developed by organisms to maximize foraging efficiency have a strong influence on fitness. The way in which the range of food resources is exploited has served to classify species, populations and individuals from more specialist (narrow trophic niche) to more generalist (broad trophic niche). Recent studies have provided evidence that many of the considered generalist species/populations are actually composed of different specialist individuals (individual specialization). Even the existence of generalism as an adaptive strategy has been questioned. In this study, we investigated the relationship between trophic niche width, individual quality and offspring viability in a population of common kestrel Falco tinnunculus during 4 years. We showed that the diet of kestrels varied significantly among years and that individuals of better quality fed their offspring with a higher diversity of prey species and a higher amount of food. Moreover, body condition and immune response of nestlings were positively correlated with diversity of prey delivered by parents. Our study suggests that generalism has the potential to increase fitness and that broadening the trophic niche may be an adaptive strategy in unpredictable environments.