Does size dimorphism reduce competition between sexes? The diet of male and female pine martens at local and wider geographical scales

Sex-specific niche segregation is often used to explain sexual size dimorphism (SSD). However, whether food niche partitioning between sexes occurs as a case of sexual size dimorphism or by other mechanisms, such as behavioural dimorphism or habitat segregation, remains poorly understood. To evaluate the nature and extent of food-niche differentiation between sexes in a solitary predator I examined variation in the diet of male and female pine martensMartes martes Linnaeus, 1758 in years of high and low rodent abundance. Small mammals were the most important prey for pine martens in years of both low and high rodent abundance (occurring in more than 49% of scats). Birds, invertebrates and plant material were relatively common food items in summer diet, whereas ungulate carcasses were often consumed in autumn—winter. In general, males consumed more ungulate carcasses, plant material, amphibians and reptiles than did females, whereas females preyed more on squirrels and birds than males. There was significant seasonally dependent, between-sex variation in the occurrence of shrews, small rodents, other mammals, birds and invertebrates in marten diet. Whereas the occurrence of bank vole, birds, carcasses and plant material changed between sexes, seasons and years with various rodent abundances, both sexes consumed larger prey and had increased food niche breadth in years of low compared with high rodent abundance. Neither prey size nor food niche breadth were significantly different between males and females. The food-niche overlap between sexes was consistently lower in spring and in years of low rodent abundance. A wider geographical comparison of different marten populations showed that the diet of males and females varied significantly between locations. Females consistently preyed on squirrels and birds, whereas males fed more often on ungulate carcasses and plant material. Local and geographical comparison of male and female diets suggest that food-niche partitioning between male and female pine martens changes across different habitat and food conditions, and is not related to sexual size dimorphism, but rather to behavioural differences between sexes.     

Sexual Niche Segregation and Gender-Specific Individual Specialisation in a Highly Dimorphic Marine Mammal

While sexual segregation is expected in highly dimorphic species, the local environment is a major factor driving the degree of resource partitioning within a population. Sexual and individual niche segregation was investigated in the Australian fur seal (Arctocephalus pusillus doriferus), which is a benthic foraging species restricted to the shallow continental shelf region of south-eastern Australia. Tracking data and the isotopic values of plasma, red blood cells and whiskers were combined to document spatial and dietary niche segregation throughout the year. Tracking data indicated that, in winter, males and females overlapped in their foraging habitat. All individuals stayed within central Bass Strait, relatively close (< 220 km) to the breeding colony. Accordingly, both genders exhibited similar plasma and red cell δ¹³C values. However, males exhibited greater δ¹³C intra-individual variation along the length of their whisker than females. This suggests that males exploited a greater diversity of foraging habitats throughout the year than their female counterparts, which are restricted in their foraging grounds by the need to regularly return to the breeding colony to suckle their pup. The degree of dietary sexual segregation was also surprisingly low, both sexes exhibiting a great overlap in their δ¹⁵N values. Yet, males displayed higher δ¹⁵N values than females, suggesting they fed upon a higher proportion of higher trophic level prey. Given that males and females exploit different resources (mainly foraging habitats), the degree of individual specialisation might differ between the sexes. Higher degrees of individual specialisation would be expected in males which exploit a greater range of resources. However, comparable levels of inter-individual variation in δ¹⁵N whisker values were found in the sampled males and females, and, surprisingly, all males exhibited similar seasonal and inter-annual variation in their δ¹³C whisker values, suggesting they all followed the same general dispersion pattern throughout the year.     

Foraging behaviour of Magellanic Penguins during the early chick‐rearing period at Isla de los Estados,Argentina

Studies of the at‐sea distribution and trophic ecology of penguins are essential to understand their role in the broader marine food web. Magellanic Penguins Spheniscus magellanicus have a wide distribution and their foraging behaviour varies across breeding sites and between sexes, among others. In this study, we characterized the at‐sea areas, the diving strategies and the relative trophic level of Magellanic Penguins breeding at Isla de los Estados, Argentina, during the early chick‐rearing period. In addition, we quantified the interannual, sexual and individual variability in those parameters during three breeding seasons (2011–2013) using devices recording position and dive depth, and obtained blood samples for stable isotope analysis. During the early chick‐rearing period, Magellanic Penguins showed small differences between the sexes in their foraging behaviour and large overlap in the at‐sea areas used, suggesting no intraspecific variation between sexes. Although there was interannual variability in the foraging behaviour and the trophic level of the penguins, most of the studied nests managed successfully to raise both chicks during the first stage of the breeding cycle (guard stage). The foraging ecology of Magellanic Penguins from this colony was comparable with results of past studies at other breeding colonies. This study contributes to the identification of important at‐sea areas for Magellanic Penguins at the southern edge of their distribution and also to the identification of possible threats in the study area such as interaction with fisheries.     

The effect of social dominance on foraging by the Three-toed Woodpecker Picoides tridactylus

During the winter, female Three-toed Woodpeckers Picoides tridactylus , when unaccompanied by the male, foraged at a lower height above the ground and on tree trunks of greater diameter than when foraging together with a male. Among males, however, no such differences were found between those birds foraging in the absence of, or in company with, a female. The niche breadth of the males was less than that of the females, indicating a higher degree of specialization by the males. The niche overlap between the sexes was less when the birds foraged together in pairs, especially as regards foraging height and foraging site diameter. The intersexual segregation of foraging niche recorded for the Three-toed Woodpecker is therefore probably not determined genetically, but is due to social dominance by males which obliges the females to occupy the less preferred niche. Both sexes spent less time being vigilant when foraging together, than when alone.     

Magellanic woodpecker ( Campephilus magellanicus) abundance and foraging in Tierra del Fuego, Chile

The Magellanic woodpecker (Campephilus magellanicus) is a vulnerable and poorly studied bird in the sub-antarctic deciduous and evergreen beech (Nothofagus) forests of South America. On Tierra del Fuego island (Chile), we compared Magellanic woodpecker abundance and its foraging habitat in two forest types: pure N. pumilio and mixed forests composed by N. pumilio and N. betuloides, including managed and non managed stands. At a regional scale, abundance of woodpeckers was greater in landscapes including both forest types than in pure N. pumilio landscapes. When both forest types occurred together, woodpecker abundance did not differ between them. The number of trees with foraging signs was correlated with Magellanic woodpecker abundance and was also associated with N. betuloides and snag densities, but was not affected by forest management. Occurrence of pecking on foraging trees was greater in mixed Nothofagus than pure N. pumilio stands. Woodpeckers foraged disproportionately more on larger diameter and more decayed trees. Moreover, trees used for foraging were positively correlated with canopy cover and snag density and were negatively correlated with distance to nearby peatlands and beaver ponds. Direct observation revealed that the flying distance between trees was negatively correlated with proportion of trees with foraging signs. Woodpeckers chose trees that were visited before, suggesting a pattern of tree recognition within foraging territories.Communicated by F. Bairlein     

同域分布3种啄木鸟冬季取食的生态位差异

为了掌握黑啄木鸟、三趾啄木鸟和大斑啄木鸟的冬季取食行为特征,特别是三者之间取食生态位的差异,于2016年1月5-13日,在黑龙江省凉水国家级自然保护区以样线法结合样方法对3种啄木鸟的取食生境和取食行为进行了系统调查,收集了15个生境和行为特征变量数据。共布设45条样线,484个对照样方,收集312组啄木鸟取食数据,其中黑啄木鸟73组,三趾啄木鸟97组,大斑啄木鸟142组。多变量回归树和多分类逻辑斯谛分析结果显示,3种啄木鸟在所调查的15项特征上存在显著分异。采用基于利用-可利用方法的Bailey''s方法和双因子方差分析,分别对3种啄木鸟的生境选择和行为特征进行分析,结果显示：黑啄木鸟和三趾啄木鸟偏好在郁闭度较高的原始云、冷杉林中取食,而大斑啄木鸟则随机地在各种林型、生境中取食。黑啄木鸟、三趾啄木鸟多在树干取食,黑啄木鸟更常在倒木上取食,而大斑啄木鸟则多在树冠层取食。黑啄木鸟基本只在主干上凿洞,其他两种特别是大斑啄木鸟则可以在侧枝上取食。与黑啄木鸟和大斑啄木鸟凿洞取食昆虫不同,三趾啄木鸟多通过扒去树皮获得食物。黑啄木鸟的取食树基本为死树,单树取食时间最长,大斑啄木鸟多在活树上取食,单树取食时间最短,经常更换取食树,而三趾啄木鸟的取食树则死活参半,单树取食时间也较长。黑啄木鸟的冬季取食行为节律表现为双峰形,日出后和日落前各有一个活动高峰,其他两种则于白天持续取食。3种啄木鸟取食生境和行为生态位的差异,使它们能够更有效地利用有限的食物资源,共存于同一森林。     

Seasonal dynamics of resource partitioning among foliage-gleaning passerines in northern Finland

Summary The five most abundant species were included in a year-round study with respect to six foraging niche dimensions. Approximately full multidimensional utilization functions were used for niche metrics. During summer foraging overlaps were invariably high, but in other seasons periodically lower. Foraging site breadth was lower in winter, when fewer sites are profitable for foraging than in summer. Feeding posture versatility, by contrast, was highest in winter. Seasonal foraging shifts were very prominent, as great in fact as between-species differences. Often seasonal trends were parallel in different species. Niche axes of macrohabitat-type (e.g. tree) were more open for foraging variation and axes of microhabitat-type (tree part) more rigid. Among the resident species seasonal variation in foraging was greatest in Regulus regulus and Parus cristatus, whereas the foraging behaviour was more stable for P. montanus (an abundant species with broad niche), perhaps owing to greater intraspecific competition. In these northern forests foliage-gleaners must be versatile generalists to cope with the unpredictable resources, and thus they overlap broadly in their general resource niches which are determined by their genetically fixed cost and benefit relations to each resource type. Anyhow, presumably during periodical food shortage, the actual resource uses adjusted by resource availability and competition may overlap narrowly.     

Reversed sexual dimorphism in tawny owls, Strix aluco, correlates with duty division in breeding effort

Even though most bird species with a raptorial feeding habit express varying extents of reversed sexual dimorphism (RSD: females bigger than males), the evolutionary basis for its maintenance, as well as its possible secondary consequences for the ecological adaptations of the different sexes, is debated. We studied pairs of tawny owls, Strix aluco (females 20% heavier than males), throughout the year by telemetry to test whether any inter-sexual differences in movement patterns, resource partitioning and breeding effort correlated with RSD. Females were larger than males in all body size measures and were 16% heavier than would be expected from the difference in wing length alone. In accordance with predictions from flight economics, males moved longer distances per time unit than females, in particular during the post-fledging season, when they also fed chicks more often than the females. Males had larger home ranges than females during the post-fledging period, whereas the sexes had home ranges of equal size during the non-breeding season. Until 10 days after fledging, females foraged much closer to the offspring than males, apparently balancing their distance to offspring between the needs of offspring guarding and foraging. In males, the parent–offspring distance only increased with decreasing brood condition. The sexes did not differ in habitat use or feeding habits, rendering no indications of food niche partitioning. The study provides further evidence that selection for males to be light and energetically efficient foragers is the main evolutionary force behind RSD in raptorial birds, even when the prey base is confined by territoriality.     

Sex‐specific winter distribution in a sexually dimorphic shorebird is explained by resource partitioning

Sexual size dimorphism (SSD) implies correlated differences in energetic requirements and feeding opportunities, such that sexes will face different trade‐offs in habitat selection. In seasonal migrants, this could result in a differential spatial distribution across the wintering range. To identify the ecological causes of sexual spatial segregation, we studied a sexually dimorphic shorebird, the bar‐tailed godwit Limosa lapponica, in which females have a larger body and a longer bill than males. With respect to the trade‐offs that these migratory shorebirds experience in their choice of wintering area, northern and colder wintering sites have the benefit of being closer to the Arctic breeding grounds. According to Bergmann's rule, the larger females should incur lower energetic costs per unit of body mass over males, helping them to winter in the cold. However, as the sexes have rather different bill lengths, differences in sex‐specific wintering sites could also be due to the vertical distribution of their buried prey, that is, resource partitioning. Here, in a comparison between six main intertidal wintering areas across the entire winter range of the lapponica subspecies in northwest Europe, we show that the percentage of females between sites was not correlated with the cost of wintering, but was positively correlated with the biomass in the bottom layer and negatively with the biomass in the top layer. We conclude that resource partitioning, rather than relative expenditure advantages, best explains the differential spatial distribution of male and female bar‐tailed godwits across northwest Europe.     

Shared dietary niche between sexes in Magellanic Penguins

Prey use by male and female Magellanic Penguins (Spheniscus magellanicus) was quantified during chick rearing in northern Golfo San Jorge, Argentina and the dietary niches and diet composition of each sex were assessed and compared. A total of 67 males and 45 females were flushed during the chick stages of 2011 and 2012. Argentine Anchovy (Engraulis anchoita) was the main prey in terms of importance by mass in males and females in all stages and years (52.4–86.6% and 83.8–88.7%, respectively), except during the old chick stage in 2011 when the main prey in males and females was the Shortfin Squid (Illex argentinus) (63.1% and 44.4%, respectively). Analyses of similarity showed that diet composition in terms of contribution by mass of the main prey species was similar between sexes. Based on the isotopic values of δ¹³C and δ¹⁵N corresponding to 38 males and 21 females, the Bayesian mixing model outputs showed that Argentine Anchovy was also the main prey. In both years and chick stages, the isotopic niche of males and females was similar. The similarity in the dietary niche between male and female Magellanic Penguins raising chicks may be due to the high availability of Argentine Anchovy during the study period and the consequent dominance of this forage species in the diet of both sexes.     

Specialisation in prey capture drives coexistence among sympatric spider‐hunting wasps

1. Sister taxa that coexist in the same space and time often face competition due to the use of similar resources. However, some closely related species can adopt fine‐grained specialisation in resource use to coexist. This study investigated niche overlap between three sympatric spider‐hunting wasp species of the genus Trypoxylon (Hymenoptera: Crabronidae) known to nest in three of the habitats found in the study area. 2. First, the co‐occurrence of these wasp species in the three habitats was estimated, as a proxy for potential competition. Then, the following hypotheses were tested: (i) niche partitioning is seen more often between species that co‐occur in a habitat, whereas there is niche overlap between species nesting in distinct habitats (prey specialisation hypothesis); and (ii) wasp species capture prey according to their size (physical constraint hypothesis). 3. Two pairs of wasp species were found consistently nesting in the same habitat. Niche partitioning based on prey taxa occurred regardless of the habitat preference. It was also found that differences in the size of wasps reflected distinctions in the size of their prey. 4. These findings were consistent over the years, showing that the significance of specialisation in foraging activities and physical constraints during prey capture can play key roles in the coexistence of sympatric species. The distinctions in the foraging strategies of these wasps are discussed, as well as potential mechanisms driving the evolution in prey specialisation, with insights for future studies.     

Intragroup competition predicts individual foraging specialisation in a group‐living mammal

Individual foraging specialisation has important ecological implications, but its causes in group‐living species are unclear. One of the major consequences of group living is increased intragroup competition for resources. Foraging theory predicts that with increased competition, individuals should add new prey items to their diet, widening their foraging niche (‘optimal foraging hypothesis’). However, classic competition theory suggests the opposite: that increased competition leads to niche partitioning and greater individual foraging specialisation (‘niche partitioning hypothesis’). We tested these opposing predictions in wild, group‐living banded mongooses (Mungos mungo), using stable isotope analysis of banded mongoose whiskers to quantify individual and group foraging niche. Individual foraging niche size declined with increasing group size, despite all groups having a similar overall niche size. Our findings support the prediction that competition promotes niche partitioning within social groups and suggest that individual foraging specialisation may play an important role in the formation of stable social groupings.     

Identifying key habitats for the conservation of Bonelli's Eagle Aquila fasciata using radiotracking

Identifying the priority habitats of endangered species is crucial to implementing effective conservation actions. We characterize the key habitats used by Bonelli's Eagle Aquila fasciata, an umbrella and flagship species that is endangered in Mediterranean countries. We radiotracked 17 breeding individuals (10 males and seven females) in Catalonia (NE Iberian Peninsula) and used compositional analysis to determine the key habitats in home‐ranges of both sexes. The main habitats identified within the home‐range area were scrubland, coniferous forests, cropland, sclerophyllous forests, rock outcrops and urban areas, with little difference in habitat use between the sexes. Bonelli's Eagles preferred rocky habitats as breeding areas and scrubland as foraging areas, as these hold the highest abundance of their main prey, Rabbits Oryctolagus cuniculus and Red‐legged Partridges Alectoris rufa. Habitat selection varied seasonally in foraging areas: scrubland was the most preferred habitat type during the breeding season, whereas rocky areas were preferred during the non‐breeding season (although scrubland was also highly selected). Urban areas were avoided both as breeding and as foraging areas. Home‐range size was inversely correlated with the proportion of scrubland, suggesting that this is a key habitat for Bonelli's Eagle. To conserve this species effectively, policies that ensure the preservation of the cliffs used as breeding sites, as well as suitable management of the scrubland used for foraging, should be implemented in the areas inhabited by this species. The promotion of traditional land uses and management techniques that will enhance open areas in Mediterranean landscapes should in the future play an essential role in the conservation of Bonelli's Eagle in Mediterranean habitats.     

Foraging ecologies of giraffe (Giraffa camelopardalis reticulata) and camels (Camelus dromedarius) in northern Kenya: effects of habitat structure and possibilities for competition?

The foraging ecologies of reticulated giraffe (Giraffa camelopardalis reticulata) and domestic camels (Camelus dromedarius) were examined in the Laikipia District of Kenya, where these species have recently become sympatric. Camels increased popularity in the region has lead to concerns about their environmental impacts and possible competition with wild giraffe for resources. We gathered foraging data on both species using 2‐min group scans that recorded feeding heights and plant food preferences. Transects sampled the vegetation in areas where foraging observations were recorded. Giraffe females feed at lower elevations than males, while female camels feed below both sexes of giraffe. There was very little observed overlap in food preferences between the species. However, habitat type has an effect on foraging ecologies of both giraffe sexes, but habitat did not influence camel foraging. Camel herder husbandry techniques also influence camel foraging dynamics. These findings have important implications in achieving the twin objectives of wildlife conservation and pastoralist livestock production in northern Kenya.     

Intersexual trophic niche partitioning in an ant-eating spider (Araneae: Zodariidae)

Background

Divergence in trophic niche between the sexes may function to reduce competition between the sexes (“intersexual niche partitioning hypothesis”), or may be result from differential selection among the sexes on maximizing reproductive output (“sexual selection hypothesis”). The latter may lead to higher energy demands in females driven by fecundity selection, while males invest in mate searching. We tested predictions of the two hypotheses underlying intersexual trophic niche partitioning in a natural population of spiders. Zodarion jozefienae spiders specialize on Messor barbarus ants that are polymorphic in body size and hence comprise potential trophic niches for the spider, making this system well-suited to study intersexual trophic niche partitioning.

Methodology/Principal Findings

Comparative analysis of trophic morphology (the chelicerae) and body size of males, females and juveniles demonstrated highly female biased SSD (Sexual Size Dimorphism) in body size, body weight, and in the size of chelicerae, the latter arising from sex-specific growth patterns in trophic morphology. In the field, female spiders actively selected ant sub-castes that were larger than the average prey size, and larger than ants captured by juveniles and males. Female fecundity was highly positively correlated with female body mass, which reflects foraging success during the adult stage. Females in laboratory experiments preferred the large ant sub-castes and displayed higher capture efficiency. In contrast, males occupied a different trophic niche and showed reduced foraging effort and reduced prey capture and feeding efficiency compared with females and juveniles.

Conclusions/Significance

Our data indicate that female-biased dimorphism in trophic morphology and body size correlate with sex-specific reproductive strategies. We propose that intersexual trophic niche partitioning is shaped primarily by fecundity selection in females, and results from sex-differences in the route to successful reproduction where females are selected to maximize energy intake and fecundity, while males switch from foraging to invest in mating effort.     

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.     

Persistent bill and corolla matching despite shifting temporal resources in tropical hummingbird‐plant interactions

By specialising on specific resources, species evolve advantageous morphologies to increase the efficiency of nutrient acquisition. However, many specialists face variation in resource availability and composition. Whether specialists respond to these changes depends on the composition of the resource pulses, the cost of foraging on poorly matched resources, and the strength of interspecific competition. We studied hummingbird bill and plant corolla matching during seasonal variation in flower availability and morphology. Using a hierarchical Bayesian model, we accounted for the detectability and spatial overlap of hummingbird‐plant interactions. We found that despite seasonal pulses of flowers with short‐corollas, hummingbirds consistently foraged on well‐matched flowers, leading to low niche overlap. This behaviour suggests that the costs of searching for rare and more specialised resources are lower than the benefit of switching to super‐abundant resources. Our results highlight the trade‐off between foraging efficiency and interspecific competition, and underline niche partitioning in maintaining tropical diversity.     

Intra‐ and inter‐individual variation show distinct trends as drivers of seasonal changes in the resource use of a neotropical marsupial

Individual specialization can influence important ecological and evolutionary traits and both inter‐ and intra‐individual variation in resource use can drive niche shifts in natural populations. We evaluated the predominance of these two factors for determining seasonal differences in the trophic niche of the didelphid marsupial Gracilinanus agilis (Burmeister, 1854) in the highly seasonal Brazilian savanna. In the three sampled sites, the population of G. agilis increased its dietary niche width in the warm–wet season, when food resources are more abundant, and there were no differences between sexes and no interaction between season and sex. However, the evaluation of intra‐individual variation indicated that females reduce the number of items consumed during the warm–wet season, whereas males show no seasonal differences. Inter‐individual variation nonetheless followed the overall population pattern because both sexes increased their spread with respect to food‐item consumption in the warm–wet season. Additionally, we found positive relationships between body length and diet only in the warm–wet season, when larger animals fed more on invertebrates and less on fruits than the small ones. Our results show a previously unknown pattern for mammals, in which the trophic niche is wider during the high‐resource season as a result of inter‐individual variation along the body‐size axis. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 737–747.     

Feeding ecology and seasonal diet overlap between Stellifer brasiliensis and Stellifer stellifer in a tropical estuarine ecocline

Seasonal freshwater discharge was important for defining habitat utilization by different ontogenetic phases of Stellifer brasiliensis and Stellifer stellifer along the estuarine ecocline. The middle estuary was important as a nursery and feeding ground for young‐of‐the‐year, and a feeding ground for sub‐adults and adults of both species. These species are zoobenthivorous, but during their life cycle and between different habitats and seasons, their trophic guild can change to opportunist and zooplanktivore. During the late rainy season in the lower estuary, all phases of both species, except juveniles of S. brasiliensis and adults of S. stellifer, showed a niche overlap indicating similarity in prey utilization. The diet composition was qualitatively similar, showing an evident niche overlap of intra and interspecific competition among the Stellifer spp. Although the niches of these species appeared to significantly overlap, some resource partitioning patterns were apparent. The niche overlap was significantly reduced due to the seasonal difference in habitat use and prey consumption along the ecocline of the estuary by different ontogenetic phases. The ingestion of blue nylon fragments by both species was observed and quantified.     

Effects of habitat fragmentation on foraging behaviour of tits and related species: does niche space vary in relation to size and degree of isolation of forest fragments?

We studied the winter foraging niches of tits and related species in deciduous forest fragments varying in size between 1 and 30 ha (plus one forest of 200 ha) in order to investigate the influence of forest fragmentation on foraging niches Very few correlations between niche structure (foraging niche, width and overlap) and forest size or isolation turned out to be significant This implies that either the species that disappear in small fragments are those that suffer most from competition (making the effect unmeasurable), or that competition is relatively unimportant for niche structure In any case we find no evidence that foraging niches are strongly affected by the changes (in habitat and/or community structure) associated with fragmentation