Intraspecific and interspecific competition induces density‐dependent habitat niche shifts in an endangered steppe bird

Interspecific competition is a dominant force in animal communities that induces niche shifts in ecological and evolutionary time. If competition occurs, niche expansion can be expected when the competitor disappears because resources previously inaccessible due to competitive constraints can then be exploited (i.e., ecological release). Here, we aimed to determine the potential effects of interspecific competition between the little bustard (Tetrax tetrax) and the great bustard (Otis tarda) using a multidimensional niche approach with habitat distribution data. We explored whether the degree of niche overlap between the species was a density‐dependent function of interspecific competition. We then looked for evidences of ecological release by comparing measures of niche breadth and position of the little bustard between allopatric and sympatric situations. Furthermore, we evaluated whether niche shifts could depend not only on the presence of great bustard but also on the density of little and great bustards. The habitat niches of these bustard species partially overlapped when co‐occurring, but we found no relationship between degree of overlap and great bustard density. In the presence of the competitor, little bustard's niche was displaced toward increased use of the species' primary habitat. Little bustard's niche breadth decreased proportionally with great bustard density in sympatric sites, in consistence with theory. Overall, our results suggest that density‐dependent variation in little bustard's niche is the outcome of interspecific competition with the great bustard. The use of computational tools like kernel density estimators to obtain multidimensional niches should bring novel insights on how species' ecological niches behave under the effects of interspecific competition in ecological communities.     

Pulsed resource availability changes dietary niche breadth and partitioning between generalist rodent consumers

Identifying the mechanisms that structure niche breadth and overlap between species is important for determining how species interact and assessing their functional role in an ecosystem. Without manipulative experiments, assessing the role of foraging ecology and interspecific competition in structuring diet is challenging. Systems with regular pulses of resources act as a natural experiment to investigate the factors that influence the dietary niches of consumers. We used natural pulses of mast‐fruiting of American beech (Fagus grandifolia) to test whether optimal foraging or competition structure the dietary niche breadth and overlap between two congener rodent species (Peromyscus leucopus and P. maniculatus), both of which are generalist consumers. We reconstructed diets seasonally over a 2‐year period using stable isotope analysis (δ¹³C, δ¹⁵N) of hair and of potential dietary items and measured niche dynamics using standard ellipse area calculated within a Bayesian framework. Changes in niche breadth were generally consistent with predictions of optimal foraging theory, with both species consuming more beechnuts (a high‐quality food resource) and having a narrower niche breadth during masting seasons compared to nonmasting seasons when dietary niches expanded and more fungi (a low‐quality food source) were consumed. In contrast, changes in dietary niche overlap were consistent with competition theory, with higher diet overlap during masting seasons than during nonmasting seasons. Overall, dietary niche dynamics were closely tied to beech masting, underscoring that food availability influences competition. Diet plasticity and niche partitioning between the two Peromyscus species may reflect differences in foraging strategies, thereby reducing competition when food availability is low. Such dietary shifts may have important implications for changes in ecosystem function, including the dispersal of fungal spores.     

Seasonal dietary niche contraction in coexisting Neotropical frugivorous bats (Stenodermatinae)

Tropical dry forests are characterized by punctuated seasonal precipitation patterns that drive primary production and the availability of fruits, seeds, flowers, and insects throughout the year. In environments in which the quantity and quality of food resources varies seasonally, consumers should adjust their foraging behavior to maximize energy intake while minimizing overlap with competitors during periods of low food availability. Here, we investigated how the diets of frugivorous bats in tropical dry forests of NW Mexico varied in response to seasonal availability and how this affected dietary overlap of morphologically similar species. We performed stable isotope analyses to understand temporal and interspecific patterns of overall isotopic niche breadth, trophic position, and niche overlap in the diet of six frugivorous species of closely related New World leaf-nosed bats (family Phyllostomidae, subfamily Stenodermatinae). We estimated seasonal changes in resource abundance in two complementary ways: (a) vegetative phenology based on long-term remote sensing data and (b) observational data on food availability from previously published insect and plant fruiting surveys. In all species, there was a consistent pattern of reduced isotopic niche breadth during periods of low food availability. However, patterns of niche overlap varied between morphologically similar species. Overall, results from our study and others suggest that seasonal food availability likely determines overall dietary niche breadth in Phyllostomidae and that despite morphological specialization, it is likely that other mechanisms, such as opportunistic foraging and spatiotemporal niche segregation, may play a role in maintaining coexistence rather than simply dietary displacement.     

Seabird species‐ and assemblage‐level isotopic niche shifts associated with changing prey availability during breeding in coastal Newfoundland

Shifting prey availability can lead to altered species interactions, indicated by variation in the dietary niche breadth and position of species within an assemblage. On the Newfoundland coast, annual inshore spawning migration of the dominant forage fish, Capelin Mallotus villosus, provides an excellent opportunity to investigate the influence of varying prey availability on dietary niche breadth and position among species. During June–August 2017, we investigated species‐ and assemblage‐level dietary responses to shifting Capelin availability of three Capelin‐eating, sympatrically breeding auk species, the Atlantic Puffin Fratercula arctica, Razorbill Alca torda and Common Murre Uria aalge. The diet of Leach's Storm Petrels Oceanodroma leucorhoa, which breed alongside the three auk species but are not known to rely on Capelin, was also examined to determine dietary shifts throughout breeding that were unrelated to Capelin availability. We quantified stable isotope ratios (δ¹⁵N, δ¹³C) in seabird blood components (plasma, cellular component) collected both before and after spawning Capelin arrived in the study area and compared isotopic niche breadth within a Bayesian framework. At the species level, auk trophic position increased and isotopic niche breadth narrowed after Capelin arrived, suggesting a more Capelin‐based diet. Simultaneously, trophic diversity of the auk assemblage, reflecting the extent of spacing among niches of species, decreased after spawning Capelin arrived inshore. Contrastingly, increased trophic position but broader isotopic niche breadth during higher relative to lower Capelin availability for Leach's Storm Petrel confirm that this species is probably not affected by the inshore arrival of Capelin, but instead that isotopic changes may be more related to a shift in breeding stage to chick‐rearing. Overall, our findings reiterate the importance of Capelin as a prey resource for breeding auks in coastal Newfoundland, but that the degree of reliance on Capelin varies among species, possibly allowing coexistence of these ecologically similar species. The findings highlight potential changing species interactions, such as increased competition, under declines in Capelin biomass.     

Effect of habitat degradation on competition,carrying capacity,and species assemblage stability

Changes in species’ trophic niches due to habitat degradation can affect intra‐ and interspecific competition, with implications for biodiversity persistence. Difficulties of measuring species’ interactions in the field limit our comprehension of competition outcomes along disturbance gradients. Thus, information on how habitat degradation can destabilize food webs is scarce, hindering predictions regarding responses of multispecies systems to environmental changes. Seagrass ecosystems are undergoing degradation. We address effects of Posidonia oceanica coverage reduction on the trophic organization of a macroinvertebrate community in the Tyrrhenian Sea (Italy), hypothesizing increased trophic generalism, niche overlap among species and thus competition and decreased community stability due to degraded conditions. Census data, isotopic analysis, and Bayesian mixing models were used to quantify the trophic niches of three abundant invertebrate species, and intra‐ and interspecific isotopic and resource‐use similarity across locations differing in seagrass coverage. This allowed the computation of (1) competition strength, with respect to each other and remaining less abundant species and (2) habitat carrying capacity. To explore effects of the spatial scale on the interactions, we considered both individual locations and the entire study area (“‘meadow scale”). We observed that community stability and habitat carrying capacity decreased as P. oceanica coverage declined, whereas niche width, similarity of resource use and interspecific competition strength between species increased. Competition was stronger, and stability lower, at the meadow scale than at the location scale. Indirect effects of competition and the spatial compartmentalization of species interactions increased stability. Results emphasized the importance of trophic niche modifications for understanding effects of habitat loss on biodiversity persistence. Calculation of competition coefficients based on isotopic distances is a promising tool for describing competitive interactions in real communities, potentially extendible to any subset of ecological niche axes for which specimens’ positions and pairwise distances can be obtained.     

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.     

Niche partitioning and the effect of interspecific competition on microhabitat use by two sympatric galaxiid stream fishes

1. Numerous interacting abiotic and biotic factors influence niche use and assemblage structure of freshwater fishes, but the strength of each factor changes with spatial scale. Few studies have examined the role of interspecific competition in structuring stream fish assemblages across spatial scales. We used field and laboratory approaches to examine microhabitat partitioning and the effect of interspecific competition on microhabitat use in two sympatric stream fishes (Galaxias‘southern’ and Galaxias gollumoides) at large (among streams and among sites within streams) and small (within artificial stream channels) spatial scales. 2. Diurnal microhabitat partitioning and interspecific competition at large spatial scales were analysed among three sympatry streams (streams with allotopic and syntopic sites; three separate catchments) and four allopatry streams (streams with only allotopic sites; two separate catchments). Electro‐fishing was used to sample habitat use of fishes at 30 random points within each site by quantifying four variables for each individual: water velocity, depth, distance to nearest cover and substratum size. Habitat availability was then quantified for each site by measuring those variables at each of 50 random points. Diet and stable isotope partitioning was analysed from syntopic sites only. Diel cycles of microhabitat use and interspecific competition at small spatial scales were examined by monitoring water velocity use over 48 h in artificial stream channels for three treatments: (i) allopatric G. ‘southern’ (10 G. ‘southern’); (ii) allopatric G. gollumoides (10 G. gollumoides) and (iii) sympatry (five individuals of each species). 3. One hundred and ninety‐four G. ‘southern’ and 239 G. gollumoides were sampled across all seven streams, and habitat availability between the two species was similar among all sites. Galaxias‘southern’ utilised faster water velocities than G. gollumoides in both the field and in channel experiments. Both species utilised faster water velocities in channels at night than during the day. Diet differences were observed and were supported by isotopic differences (two of three sites). No interspecific differences were observed for the other three microhabitat variables in the field, and multivariate habitat selection did not differ between species. Interspecific competition had no effect on microhabitat use of either species against any variable either in the field (large scale) or in channels (small scale). 4. The results suggest that niche partitioning occurs along a subset of microhabitat variables (water velocity use and diet). Interspecific competition does not appear to be a major biotic factor controlling microhabitat use by these sympatric taxa at any spatial scale. The results further suggest that stream fish assemblages are not primarily structured by biotic factors, reinforcing other studies de‐emphasising interspecific competition.     

Niche segregation of coexisting Arctic charr (Salvelinus alpinus) and brown trout (Salmo trutta) constrains food web coupling in subarctic lakes

1. Generalist fish species are recognised as important couplers of benthic and pelagic food‐web compartments in lakes. However, interspecific niche segregation and individual specialisation may limit the potential for generalistic feeding behaviour. 2. We studied summer habitat use, stomach contents and stable isotopic compositions of the generalist feeder Arctic charr coexisting with its common resource competitor brown trout in five subarctic lakes in northern Norway to reveal population‐level and individual‐level niche plasticity. 3. Charr and trout showed partial niche segregation in all five lakes. Charr used all habitat types and a wide variety of invertebrate prey including zooplankton, whereas trout fed mainly on insects in the littoral zone. Hence, charr showed a higher potential to promote habitat and food‐web coupling compared to littoral‐dwelling trout. 4. The level of niche segregation between charr and trout and between pelagic‐caught and littoral‐caught charr depended on the prevailing patterns of interspecific and intraspecific resource competition. The two fish species had partially overlapping trophic niches in one lake where charr numerically dominated the fish community, whereas the most segregated niches occurred in lakes where trout were more abundant. 5. In general, pelagic‐caught charr had substantially narrower dietary and isotopic niches and relied less on littoral carbon sources compared to littoral‐caught conspecifics that included generalist as well as specialised benthivorous and planktivorous individuals. Despite the partially specialised planktivorous niche and thus reduced potential of pelagic‐dwelling charr to promote benthic–pelagic coupling, the isotopic compositions of both charr subpopulations suggested a significant reliance on both littoral and pelagic carbon sources in all five study lakes. 6. Our study demonstrates that both interspecific niche segregation between and individual trophic specialisation within generalist fish species can constrain food‐web coupling and alter energy mobilisation to top consumers in subarctic lakes. Nevertheless, pelagic and littoral habitats and food‐web compartments may still be highly integrated due to the potentially plastic foraging behaviour of top consumers.     

The coexistence of fish species in streams: relationships between assemblage attributes and trophic and environmental variables

Diet overlap and niche breadth are well-known species traits from trophic ecology that can assist in explaining how species interact and coexist as well as the ecological mechanisms that influence biodiversity. In the present study, we analyzed the relationships between these trophic variables and indicators of resource availability with some attributes of fish assemblages (species richness, Shannon diversity index, evenness, density and individual body size). The physical and chemical characteristics of the biotopes (topography, water quality and conservation of slopes) were examined to identify possible patterns. Monthly sampling using electrofishing was conducted in 2003 along five streams located in the Cuiabá River watershed. The relationships between environmental variables and attributes of fish assemblages were evaluated using Spearman correlation. Species richness and abundance varied among streams, with higher values (54 and 82 species) found in low-gradient streams that drained small swampy areas discharging in Cuiabá River. Diet overlap showed significant and negative correlations with species richness, the diversity index, abundance, variation in body size and food availability and positive correlations with evenness and niche breadth. Environments that had greater food availability had a reduced variety of food items (smaller amplitude) and distinct values in terms of dietary overlap. Nevertheless, to explain resource partitioning in a fish assemblage (overlap), it is necessary to consider food availability and niche breadth (degree of trophic specialization) beyond the interaction of these variables with species richness and density. In conclusion, high diversity and abundance values were strongly associated with a high degree of trophic specialization (low amplitude of trophic niche) and a small degree of overlap in the diet.     

Niche expansion and the niche variation hypothesis: does the degree of individual variation increase in depauperate assemblages?

The niche expansion and niche variation hypotheses predict that release from interspecific competition will promote niche expansion in depauperate assemblages. Niche expansion can occur by different mechanisms, including an increase in within-individual, among-individual, or bimodal variation (sexual dimorphism). Here we explore whether populations with larger niche breadth have a higher degree of diet variation. We also test whether populations from depauperate lizard assemblages differ in dietary resource use with respect to variation within and/or among individuals and sexual dimorphism. We found support for the niche expansion and niche variation hypotheses. Populations in assemblages with low phylogenetic diversity had a higher degree of individual variation, suggesting a tendency for niche expansion. We also found evidence suggesting that the mechanism causing niche expansion is an increase in variation among individuals rather than an increase in within-individual variation or an increase in bimodal variation due to sexual dimorphism.     

Stable isotope analysis reveals variation in trophic niche depending on altitude in an endemic alpine gecko

Interspecific competition is considered a major determinant of ecological niche. It is hypothesized that increased competition should reduce niche breadth. However, there are scarce field tests on this hypothesis. Here, we test this central hypothesis in ecology by using the Atlas day gecko Quedenfeldtia trachyblepharus. This alpine gecko faces fewer competitors as altitude increases, and thereby, we predict that this species should increase niche breadth and relevant fitness parameters with altitude. We tested this prediction by analysing the isotopic signature of carbon (δ¹³C) and nitrogen (δ¹⁵N). Our results reveal that specimens from higher altitudes showed higher values for both carbon (δ¹³C) and nitrogen (δ¹⁵N) isotopes, had better body condition and a greater isotopic breadth when compared to specimens from lower altitudes. Altitudinal variation in carbon values was not explained by variation in isotopic concentration in the baseline of the trophic chain. Therefore, our findings support the prediction that relaxed interspecific competition favours increased trophic niche breadth. These results also suggest that global warming may represent an important threat for this species, as it may provoke the ascent in altitude of competitors, with negative consequences for the conservation of this endemism.     

Testing the niche variation hypothesis with a measure of body condition

Individual variation and fitness are cornerstones of evolution by natural selection. The niche variation hypothesis (NVH) posits that when interspecific competition is relaxed, intraspecific competition should drive niche expansion by selection favoring use of novel resources and that among‐individual variation should confer a selective advantage. Population‐level niche expansion could be achieved by all individuals using all available resources, or by each individual using a unique combination of resources, thereby increasing among‐individual dietary niche variation. Although individual variation can lead to species‐level evolutionary and ecological change, observed variation does not ensure a beneficial outcome. We used carbon and nitrogen stable isotope analysis of claw keratin and a Bayesian stable isotope mixing model to estimate the summer (July–September) assimilated diet of individual female black Ursus americanus and brown U. arctos bears. We quantified variation in dietary niche in both populations, and assessed diet relative to percentage body fat. We hypothesized that if the NVH held, percentage body fat would be similar for individuals of the same species across much of the dietary range of observed proportional salmon contributions to individual bear diets. Although we found greater differences in dietary niches between than within species, we observed greater among‐individual dietary variation in the brown bear population. Moreover, we found that within each species individual female bears achieved similar ranges of percentage body fat at various levels of salmon in the diet. Our results provide support for the NVH. Linking individual dietary niches to measures of physiological condition related to fitness can offer new insights into eco‐evolutionary processes related to food resource use.     

择伐对思茅松天然林乔木种间与种内关系的影响

通过分析不同择伐强度下思茅松(Pinus kesiya)天然林乔木种群的生态位、种间联结、种内与种间竞争和群落稳定性, 探讨干扰强度对思茅松天然林乔木种群种间与种内关系的影响。结果表明: (1)择伐干扰改变了大多数乔木树种在群落中的优势地位, 思茅松优势地位未受影响, 随着择伐干扰强度的增加, 思茅松在群落中的生态位宽度逐渐变大, 在重度干扰的群落中与红木荷(Schima wallichii)一起成为生态幅度最大的树种。(2)通过种间联结与生态位重叠分析发现, 随着干扰强度的加剧, 乔木种群的种间关系发生变化, 中度干扰群落中物种之间的生态位重叠明显高于未干扰和重度干扰群落, 而在三类群落中, 思茅松与其他种群之间的重叠值较小。随着干扰强度增加, 种群间正联结与负联结的种对所占的比例逐渐增加。(3) Hegyi竞争指数表明, 三类群落中思茅松种群的竞争压力主要来自种间, 而干扰强度进一步增加了思茅松种群的种间竞争强度, 其种内平均竞争指数随着对象木胸径的增加而减少。(4)林窗是思茅松种群种间和种内关系变化的主要原因, 三类群落都处于较不稳定的阶段, 但中度干扰群落的物种丰富度与稳定性都高于未干扰和重度干扰的群落。择伐原则依据森林管理的目的而确定, 如以木材培育为主或森林生态系统服务功能为主。     

Grassland harvesting alters ant community trophic structure: An isotopic study in tallgrass prairies

Disturbances have long been recognized as important forces for structuring natural communities but their effects on trophic structure are not well understood, particularly in terrestrial systems. This is in part because quantifying trophic linkages is a challenge, especially for small organisms with cryptic feeding behaviors such as insects, and often relies on conducting labor‐intensive feeding trials or extensive observations in the field. In this study, we used stable isotopes of carbon and nitrogen to examine how disturbance (annual biomass harvesting) in tallgrass prairies affected the trophic position, trophic range, and niche space of ants, a widespread grassland consumer. We hypothesized that biomass harvest would remove important food and nesting resources of insects thus affecting ant feeding relationships and trophic structure. We found shifts in the feeding relationships inferred by isotopic signatures with harvest. In particular, these shifts suggest that ants within harvest sites utilized resources at lower trophic levels (possibly plant‐based resources or herbivores), expanded trophic breadth, and occupied different niche spaces. Shifts in resource use following harvest could be due to harvest‐mediated changes in both the plant and arthropod communities that might affect the strength of competition or alter plant nitrogen availability. Because shifts in resource use alter the flow of nutrients across the food web, disturbance effects on ants could have ecosystem‐level consequences through nutrient cycling.     

Herbivore overlap and competition in Kenya rangelands

Small herbivores such as lagomorphs, rodents and orthopterans are important consumers in grasslands. Techniques of data collection by microscope analysis of stomach contents and quantitative analysis of results are presented for assessing dietary overlap and potential competition for food resources among different classes of herbivores. Example analysis of data for small herbivores and ungulates from an area south of Nairobi National Park, Kenya, are presented. In the average growing season, diet overlap between angulates and small herbivores is low, and therefore, potential for competition slight. Considering sesonal variation in food resources availability and extreme shortages documented for occasional seasons, it is probable that periodic competition occurs. Hares are potentially the greatest competitors with ungulates in such conditions, because of their wide niche breadth and spatial overlap with ungulates. Assessment of the degree and implications of periodic competitive interactions requires careful analysis of niche breadth and overlap on the food resource gradient based upon a spatial and temporal evaluation of habitat utilization.     

Spatial,seasonal and individual variation in the diet of White‐tailed Eagles Haliaeetus albicilla assessed using stable isotope ratios

Many raptor species are considered to be generalists, taking a range of prey species. However, longitudinal dietary records are often scarce, although necessary for characterizing niche width of species at population and individual levels. Quantifying raptor diets at large spatio‐temporal scales is often necessary for refining conservation efforts, although it can be particularly difficult and may involve a great effort by conventional means. Therefore, we adopted the analysis of stable isotopes in tissues of predators and their potential food sources as a complementary methodology for assessing animals' diet. We examined the isotopic composition (δ¹³C and δ¹⁵N) of White‐tailed Eagles Haliaeetus albicilla from Germany, Finland and Greenland to detect patterns of dietary variation and quantify diet composition. The isotopic analysis included liver and muscle samples from Eagles of the three populations together with 16 potential food sources in the German population. Our results suggested dietary differences between German and Greenlandic Eagles, in accordance with the availability of freshwater and marine habitats in each population. Within the German population, we found seasonal shifts in isotopic ratios, suggesting the birds responded to temporal changes in food availabilities, and age‐related isotopic differences, indicating different diets in adults and juveniles. Isotopic values of liver and muscle tissues collected from the same animal showed intra‐individual short‐term changes in the German and Finnish but not Greenlandic population. This suggests that local feeding niches of this generalist predator may vary with local food supplies, which determines the niche width (from generalist to specialist) at the individual level. Our results also revealed that game mammal carcasses constitute an important food source (29.5% of diet) for the German Eagle population during the winter half‐year corresponding to the hunting season. This result is of relevance to management and conservation because the White‐tailed Eagle and other raptor species are affected by the ingestion of lead ammunition from shot mammalian carcasses.     

Individual dietary specialization in a generalist predator: A stable isotope analysis of urban and rural red foxes

Some carnivores are known to survive well in urban habitats, yet the underlying behavioral tactics are poorly understood. One likely explanation for the success in urban habitats might be that carnivores are generalist consumers. However, urban populations of carnivores could as well consist of specialist feeders. Here, we compared the isotopic specialization of red foxes in urban and rural environments, using both a population and an individual level perspective. We measured stable isotope ratios in increments of red fox whiskers and potential food sources. Our results reveal that red foxes have a broad isotopic dietary niche and a large variation in resource use. Despite this large variation, we found significant differences between the variance of the urban and rural population for δ¹³C as well as δ¹⁵N values, suggesting a habitat‐specific foraging behavior. Although urban regions are more heterogeneous regarding land cover (based on the Shannon index) than rural regions, the dietary range of urban foxes was smaller compared with that of rural conspecifics. Moreover, the higher δ¹³C values and lower δ¹⁵N values of urban foxes suggest a relatively high input of anthropogenic food sources. The diet of most individuals remained largely constant over a longer period. The low intraindividual variability of urban and rural red foxes suggests a relatively constant proportion of food items consumed by individuals. Urban and rural foxes utilized a small proportion of the potentially available isotopic dietary niche as indicated by the low within‐individual variation compared to the between‐individual variation. We conclude that generalist fox populations consist of individual food specialists in urban and rural populations at least over those periods covered by our study.     

Realised niche changes in a native herbivore assemblage associated with the presence of livestock

Habitat partitioning is a common ecological mechanism to avoid competition among coexisting species, and the introduction of new species into existing assemblages can increase competitive pressures. However, situations of species in allopatry and sympatry only differing in species presence but not in environmental conditions are scarce. Thus, discerning whether niche segregation arises from competition or from different habitat preferences is usually unfeasible. Here, we analyse species’ habitat niches in an assemblage of native and introduced herbivores in southern Patagonia. We test if niche overlap is higher between native and domestic herbivores than among natives as expected from the relatively short time of coexistence, and we evaluate the effect of intra‐ and interspecific competition on niche breadth. We use a probabilistic multidimensional approach and null models to evaluate overlap and changes in niche dimensions. Overlap among native species is low as expected for species coexisting in evolutionary time. In native‐domestic species pairs, niche overlap was higher than among natives, although showing some niche segregation indicating niche differentiation in ecological time. Moreover, the presence of domestic species was associated with niche narrowing of both native and introduced species, revealing interspecific density‐dependent effects on their habitat niche during resource shortage periods.     

Regional occupancy in unicellular eukaryotes: a reflection of niche breadth,habitat availability or size‐related dispersal capacity?

1. The distribution patterns of unicellular and multicellular organisms have recently been shown to differ profoundly, with the former probably being mostly cosmopolitan, whereas the latter are mostly restricted to certain regions. However, the within‐region distribution patterns of these two organism groups may be rather similar. 2. We predicted that the degree of regional occupancy in unicellular eukaryotes would be related to niche characteristics, dispersal ability and size, as has been found previously for multicellular organisms. The niche characteristics we considered were niche position, that measures marginality in species habitat distribution, and niche breadth, that measures amplitude in species habitat distribution. Niche characteristics were determined using Outlying Mean Index (OMI) analysis. 3. We found that the regional occupancy in our model group of unicellular eukaryotes, stream diatoms, was primarily a reflection of the niche position of a species or, more generally, habitat availability. Thus, non‐marginal species (i.e. species that occupied common habitat conditions across the region) tended to be more widely distributed than marginal species (i.e. species that were restricted to a limited range of rare habitat conditions). This finding was further supported by the general linear model, with niche position, niche breadth, maximum size and attachment mode as explanatory variables: niche position was by far the most important variable accounting for variability in regional occupancy, with significant amounts of additional variation related to niche breadth and maximum size of diatoms. 4. Thus, the degree of regional occupancy among unicellular eukaryotes may be primarily governed by habitat availability, supporting former findings for multicellular organisms.     

Co-existence of nine gill ectoparasites (Dactylogyrus: monogenea) parasitising the roach (Rutilus rutilus l.): history and present ecology

Co-existence among potentially competing species can be favoured by niche specialisation and/or by reducing the overall intensity of competition via aggregated utilisation of fragmented resources. We investigated the respective roles of niche specialisation and aggregation in the case of nine congeneric monogenean parasites on the gills of Roach (Rutilus rutilus L.) belonging to the genus Dactylogyrus. The position of each individual parasite of the nine Dactylogyrus species was recorded. Niche breadth and niche overlap of parasite species were estimated. Comparative methods, which take into account phylogenetic information of the analysed species, were used. We reconstructed a phylogeny of the nine Dactylogyrus species based on morphological characters. We used the 'aggregation model of co-existence' in the model to test if species co-existence is facilitated when intraspecific aggregation exceeds interspecific aggregation. We observed a lack of negative correlation in abundance between pairs of parasites, and a negative correlation between niche size and parasite aggregation, for both intraspecific and interspecific aggregation. Our comparative analysis showed that parasite abundance is positively correlated with niche breadth. Then parasite abundance, and not interactions between Dactylogyrus species, seems to be the most important factor determining niche size This result gives some support to niche segregation by specialisation. Niche size was negatively correlated with both intraspecific and interspecific aggregation. No relationship was found between an increase of interspecific aggregation with an increase of niche overlapping, which suggests that competition may play little role. A lack of competition could be also confirmed by the lack of negative correlation in abundance between species pairs. A parsimony analysis of the evolution of gill distribution indicates a change in one parameter of the niche (arch, segment and/or area) at each branching event.