Density-dependence by habitat heterogeneity: individual quality versus territory quality

The aim of this comment is to review the ecological issues concerning the role of individual and habitat heterogeneity as possible mechanisms explaining density-dependent fecundity in animal populations. Our intention is to discuss different approaches to determine whether or not studied populations are subjected to density-dependent processes and which mechanisms are involved. We show that because individual quality (e.g. measured in terms of age of breeding individuals) and territory quality can be correlated, untangling both effects on fecundity is frequently a difficult task. We discuss the misuse of statistical methods and other problems related to the specific characteristics of the studied populations that can have a strong influence on the conclusions reached by researchers working in this field.     

Trophic interactions and habitat segregation between competing Daphnia species

Summary Two commonly coexisting species of Daphnia segregate by habitat in many stratified lakes. Daphnia pulicaria is mostly found in the hypolimnion whereas D. galeata mendotae undergoes diel vertical migration between the hypolimnion and the epilimnion. I examined how habitat segregation between these two potentially competing species might be affected by trophic interactions with their resources and predators by performing a field experiment in deep enclosures in which I manipulated fish predation, nutrient levels, and the density of epilimnetic Daphnia. The results of the experiment indicate that habitat use by D. pulicaria can be jointly regulated by competition for food from epilimnetic Daphnia and predation by fishes. Patterns of habitat segregation between the two Daphnia species were determined by predation by fish but not by nutrient levels: The removal of epilimnetic fish predators resulted in higher zooplankton and lower epilimnetic phytoplankton densities and allowed D. pulicaria to expand its habitat distribution into the epilimnion. In contrast, increased resource productivity resulted in higher densities of both Daphnia species but did not affect phytoplankton levels or habitat use by Daphnia. The two species exhibit a trade-off in their ability to exploit resources and their susceptibility to predation by fish. D. g. mendotae (the less susceptible species) may thus restrict D. pulicaria (the better resource exploiter) from the epilimnion when fish are common due to lower minimum resource requirements than those needed by D. pulicaria to offset the higher mortality rate imposed by selective epilimnetic fish predators. D. g. mendotae does not appear to have this effect in the absence of fish.     

Spatial heterogeneity, source-sink dynamics, and the local coexistence of competing species

Patch occupancy theory predicts that a trade-off between competition and dispersal should lead to regional coexistence of competing species. Empirical investigations, however, find local coexistence of superior and inferior competitors, an outcome that cannot be explained within the patch occupancy framework because of the decoupling of local and spatial dynamics. We develop two-patch metapopulation models that explicitly consider the interaction between competition and dispersal. We show that a dispersal-competition trade-off can lead to local coexistence provided the inferior competitor is superior at colonizing empty patches as well as immigrating among occupied patches. Immigration from patches that the superior competitor cannot colonize rescues the inferior competitor from extinction in patches that both species colonize. Too much immigration, however, can be detrimental to coexistence. When competitive asymmetry between species is high, local coexistence is possible only if the dispersal rate of the inferior competitor occurs below a critical threshold. If competing species have comparable colonization abilities and the environment is otherwise spatially homogeneous, a superior ability to immigrate among occupied patches cannot prevent exclusion of the inferior competitor. If, however, biotic or abiotic factors create spatial heterogeneity in competitive rankings across the landscape, local coexistence can occur even in the absence of a dispersal-competition trade-off. In fact, coexistence requires that the dispersal rate of the overall inferior competitor not exceed a critical threshold. Explicit consideration of how dispersal modifies local competitive interactions shifts the focus from the patch occupancy approach with its emphasis on extinction-colonization dynamics to the realm of source-sink dynamics. The key to coexistence in this framework is spatial variance in fitness. Unlike in the patch occupancy framework, high rates of dispersal can undermine coexistence, and hence diversity, by reducing spatial variance in fitness.     

Spatial patterns of coexistence of competing species in patchy habitat

The single-species spatially realistic patch occupancy metapopulation model is, in this study, extended to a metacommunity of many competing species. Competition is assumed to reduce the local carrying capacity (effective patch area), which in turn increases local extinction rates and reduces colonization rates because of smaller population sizes. Each species is described by three parameters: pre-competitive abundance (equilibrium incidence of patch occupancy, which reflects the rate of colonization in relation to extinction rate), the spatial range of migration, and competitive ability. The model ignores spatio–temporal correlations caused by interspecific interactions, because in metacommunities of unequal competitors inhabiting heterogeneous landscapes, correlations in the occurrence of species are driven more by patch heterogeneity than by competition. The model allows the calculation of multispecies equilibria in patchy habitats without simulations. In general, the number of coexisting species in the metacommunity increases with decreasing strength of competition, increasing rate of colonization, and decreasing range of migration. Habitat heterogeneity in the form of spatial variation in patch areas tends to facilitate coexistence. Poor competitors may coexist with superior competitors in the patch network if the former have higher colonization rates (competition–colonization trade-off). When migration distances are short, competition leads to spatial pattern formation: Species tend to have restricted spatial distributions in the network, but contrary to intuitive expectations, often the distributions of many species are nested. Having more dispersive species enhances both local and global diversity, whereas more local migration decreases local but increases global diversity.     

Non-breeding season habitat quality mediates the strength of density-dependence for a migratory bird

Our understanding of when natural populations are regulated during their annual cycle is limited, particularly for migratory species. This information is needed for parametrizing models that can inform management and conservation. Here, we use 14 years of data on colour-marked birds to investigate how conspecific density and habitat quality during the tropical non-breeding period interact to affect body condition and apparent annual survival of a long-distance migratory songbird, the American redstart (Setophaga ruticilla). Body condition and survival of birds in high-quality mangrove habitat declined as density increased. By contrast, body condition improved and survival did not vary as density increased in adjacent, lower quality scrub habitat, although mean condition and survival were almost always lower than in mangrove. High rainfall enhanced body condition in scrub but not in mangrove, suggesting factors such as food availability outweighed consequences of crowding in lower quality habitat. Thus, survival of overwintering redstarts in mangrove habitat, disproportionately males, appears to be regulated by a crowding mechanism based on density-dependent resource competition. Survival of individuals in scrub, mostly females, appears to be limited by density-independent environmental factors but not regulated by crowding. The contrasting effects of density and food limitation on individuals overwintering in adjacent habitats illustrate the complexity of processes operating during the non-breeding period for migratory animals, and emphasize the need for long-term studies of animals in multiple habitats and throughout their annual cycles.     

Effects of habitat fragmentation and soil quality on reproduction in two heathland Genista species

Habitat fragmentation decreases plant population size and increases population isolation, as well as altering patterns of plant–animal interactions, all of which may reduce plant fitness. Here, we studied effects of habitat fragmentation (in terms of population size and isolation) and soil quality on the reproduction of two rare legume species, Genista anglica (13 populations) and Genista pilosa (14 populations), confined to remnants of acidic and nutrient‐poor Calluna heathlands. Single individuals of the Genista plants are impossible to distinguish; population size was therefore estimated according to the area occupied (referred to as population size hereafter). We collected soil samples in all heathland sites to determine content of soil water, C, N, P, Ca, K and Mg. In both species values of soil pH and C/N ratio, as well as content of soil P and base cations, reflected the highly acidic and nutrient‐poor environment of the heathlands. Population sizes were unrelated to soil quality. Although the two Genista species are similar in morphology and ecology, effects of explanatory variables on reproduction were largely inconsistent across species. In G. anglica, population size had a positive impact on all reproductive variables except germination rate, which, in contrast, was the only variable affected positively by population size in G. pilosa. In both species, mean total reproductive output, calculated as the product of total seed mass per shoot and total germination, increased with increasing water content and decreased with increasing P. In G. anglica, we found positive effects of the C/N ratio on all reproductive variables except mean single and total seed mass per shoot. In summary, in both species reproductive success per shoot decreased with increasing soil nutrient availability in the heathland sites. The infestation of two large populations of G. pilosa with the pre‐dispersal, seed‐predating weevil Apion compactum had no significant effect on reproduction of the populations.     

Environmental effects at two nested spatial scales on habitat choice and breeding performance of barn swallow

Parental effects comprise a wide range of mechanisms that individuals may adopt to enhance viability and adjust the phenotype of their offspring according to the conditions that the offspring will experience after birth. For example, individual choice of breeding habitat may mediate such parental effects via an effect of prenatal breeding conditions independently or in combination with offspring post-natal environment. However, ecological factors relevant to adaptive breeding habitat choice may vary at different spatial scales, which have been rarely investigated simultaneously. In the first part of the present study we use hierarchical linear models to disentangle micro- and macro-environmental variation in abundance and breeding performance of a small passerine bird, the barn swallow Hirundo rustica. We show that environmental conditions at the scale of nesting microhabitat are more influential than macro-environmental conditions at the scale of foraging range. We then experimentally investigate the effect of variation in micro-environmental conditions on growth and immunity of chicks by partially cross-fostering nestlings immediately after hatching between different nesting micro-habitats. Our results disclosed significant effects of environmental conditions where eggs were laid and incubated but not of those where nestlings grew-up on some components of nestling phenotype important for fitness. These results suggest that adults may enhance offspring quality by adjusting prenatal parental effects mediated by e.g., egg quality according to micro-habitat conditions where parents are breeding.     

Compared distribution within a disturbed fishpond ecosystem of breeding ducks and bird species indicators of habitat quality

The distribution of breeding ducks versus three other bird species (Mute Swan Cygnus olor, Black-headed Gull Larus ridibundus and Purple Heron Ardea purpurea) among waterbodies was investigated in the Dombes area, Eastern France, where breeding duck populations have undergone a severe decline following a large-scale transformation of meadow habitat into cropland. Higher Pochard Aythya ferina, Tufted Duck Aythya fuligula, Red-crested Pochard Netta rufina and Gadwall Anas strepera pair densities were recorded in fishponds where Black-headed Gulls were nesting. In these ponds however, nesting success (assessed by the number of broods divided by the number of pairs) was not significantly higher. Similarly, Pochard pair density was higher in ponds with a Purple Heron colony, but brood densities were not. We hypothesise that, in the study region, clutch concentration in the most attractive areas could compensate for the anti-predation effect of gull colonies or overwater nesting in shore vegetation. We could not confirm the expected negative impact of Mute Swan aggressive behaviour on duck distribution. Even though Mallard Anas platyrhynchos pair density was lower in ponds with breeding swans, we did not observe difference for Mallard broods. Moreover, Red-crested Pochard pair density and nesting success were higher in ponds where swans were breeding, probably as a consequence of shared habitat preferences.     

Reproductive performance of little penguins Eudyptula minor in relation to year, age, pair-bond duration, breeding date and individual quality

We measured breeding performance of little penguins Eudyptula minor at Phillip Island, Victoria, Australia, during a 21-year period. All birds considered in this paper (n=307) were of known age (2–22 y) and sex, and most were of known, or closely-estimated, pair-bond status (1–8 mates per bird; pair-bond durations 1–13 y). Breeding dates and breeding performance varied markedly from year to year; measures of annual performance were not associated with early breeding. Measures of individual breeding performance (clutch-size, hatching success, chick masses and productivity) were related to early laying, parental age, and duration of pair-bond. Dependence of breeding performance on parental age was curvilinear, levelling off at about 8 y of age. Productivity declined significantly among birds older than 8 y; this decline was not due to events in the last year of breeding ("terminal illness"). Breeding performance increased with duration of pair-bond at least through y 5. Early breeding was significantly related to age and duration of pair-bond. Most of these relationships were stronger among males than among females, and many of them were not significant when females were considered alone. After controlling for other factors, breeding performance varied significantly among birds, but autocorrelations were low and limited to intervals of one year. Parental quality (defined for birds studied in six or more years as the individual bird term in a GLM for productivity controlling for other factors) was not correlated with lifespan or other demographic parameters, but high-quality birds were less prone to change mates and burrows than low-quality birds. We know of no previous study in which simultaneous effects of laying date, age and pair-bond duration on breeding performance were measured, while controlling for year, individual quality and terminal illness.     

Diversity and abundance of birds in relation to forest fragmentation,habitat quality and heterogeneity

This study investigated the importance of habitat quality and habitat heterogeneity for the abundance and diversity of breeding birds in continuous forest and in forest fragments surrounded by farmland in central Sweden. Positive correlations were found between species number and area, volume of Aspen Populus tremula and habitat heterogeneity. Spatial segregation of habitats at a relatively fine-grained scale is suggested to allow for the co-occurrence of more species. The abundance of at least 18 of the species in this study was influenced by fragmentation, and nine of these species preferred fragments to forest sites. The total density of birds was higher in fragments than in forest sites, probably because several fragment species forage in farmland surrounding the sites and a few also forage at edges. Nine species were more common in forest sites than in fragments, but only one species was restricted to continuous forest. However, several fragments were relatively close to forests (150 m) and forest was common in larger scale contexts. The abundance of most species (25 of 33 species) in this study was correlated with habitat quality variables (i.e. variables measuring the size, volume and diversity of ‘tree species’). Among these habitat variables the most important was the occurrence of deciduous trees which seemed to be important for 14 species. The second most important habitat factor seemed to be the diameter of trees, which was positively correlated with the abundance of eight species of which five are hole-nesters. Among coniferous trees, six species were positively correlated with the volume of Norway Spruce Picea abies, whereas no species seemed to be correlated with the volume of Pine Pinus sylvestris.     

Connectivity, habitat heterogeneity, and population persistence in Ranunculus nodiflorus, an endangered species in France

Here, we explore the role of habitat spatial structure in the maintenance of metapopulations of Ranunculus nodiflorus. This rare species grows in puddles that can be connected occasionally by flooded corridors. We monitored five locations in the Fontainebleau forest, France, since 2002 and recorded the presence of corridors among puddles and evaluated their impact on puddle demography and plant fitness. We showed that connections increased population size, by increasing both the number of puddles occupied by the species and the density of individuals within puddles, but seemed to have no direct influence on plant fitness. We found no evidence of a large persistent soil seed bank. Natural corridors are likely to decrease the extinction probability of the populations, most probably by allowing recolonization of empty puddles after extinctions. Therefore, the preservation of corridors appears crucial for the conservation of R. nodiflorus in its natural habitat.     

Oviposition behavior and offspring performance in herbivorous insects: consequences of climatic and habitat heterogeneity

The preference–performance hypothesis predicts that when female herbivorous insects determine where to position offspring of low mobility, they will select sites that maximize development and survival of those offspring. How this critical relationship responds to variation in climatic and habitat conditions remains untested, however, despite its important consequences for population and evolutionary dynamics. Here we report on 13 years of data totaling 1348 egg clusters of the montane Gillette's checkerspot butterfly Euphydryas gillettii (Lepidoptera: Nymphalidae). We used these data to test the hypothesis that, in environments with climatic and habitat heterogeneity, the oviposition behavior–offspring performance relationship should vary in both space and time. Orientation of egg clusters for maximum morning sun exposure is known to affect developmental rate. We therefore predicted female preference for morning sun orientation to be variable and a function of climatic and habitat conditions. We found that preference for egg cluster orientation on the leaf tracked the phenology of the start of the female flight season but that seasonal temperatures drove most of the variation in egg cluster development time. The relationship between behavior and performance was also dependent upon the climatic effects on survival; sun‐oriented egg clusters had higher survivorship in the coldest year of the four years for which measurements were made. We also examined how conifer cover affected larval survival and female oviposition behavior in one year. Females selected oviposition sites in more open habitat. However, when egg clusters were oriented to intercept morning sun, conifer cover increased survivorship to diapause. Finally, we found that predator activity was lower for morning sun‐oriented egg clusters suggesting that predation patterns may further influence habitat selection for oviposition. This study exemplifies how the relationship between oviposition behavior and offspring performance is context‐dependent: habitat and climate interact to determine preference–performance outcomes.     

Increased survival and breeding performance of double breeders in little penguins Eudyptula minor,New Zealand: evidence for individual bird quality?

The little penguin Eudyptula minor is unique among penguin species in being able to fledge chicks from two clutches in one breeding season. Pairs laying two clutches in a given season make a higher reproductive investment, and may be rewarded by a higher reproductive success as they may raise twice as many chicks as pairs laying one clutch. The higher effort made by pairs laying two clutches could correlate negatively with survival, future reproductive performance or offspring survival, indicating a cost of reproduction. Conversely, a positive relationship between the number of clutches produced in a given breeding season and survival, future reproductive performance or offspring survival would indicate that birds laying two clutches belonged to a category of birds with higher fitness, compared to birds laying only one clutch in the season. In this study we used a long‐term data set taken from an increasing population of little penguins in Otago, SE New Zealand. We modelled the relationship between the number of clutches laid in a breeding season and survival probability, reproductive performance in the next breeding season and first year survival of offspring using capture‐recapture modelling.
Birds laying two clutches produced 1.7 times more fledglings during a breeding season than pairs laying one clutch. We found that birds laying two clutches had a higher probability of breeding in the following breeding season, a higher probability of laying two clutches in the following breeding season and a higher survival probability. There was no overall difference in post‐fledging survival between the young of birds producing one clutch and the young of birds producing two clutches. However, the survival of young of single clutch breeders declined with laying date, whereas the young of double clutch breeders had the same survival rate irrespective of laying date. For a subset of data with birds of known age, we found evidence that the probability of laying two clutches increased with age. However, there were also indications for differences among birds in the tendency to lay two clutches that could not be attributed to age. We tentatively interpret our results as evidence of quality difference among little penguin breeders.     

Environmental effects on vertebrate species richness: testing the energy, environmental stability and habitat heterogeneity hypotheses

Background

Explaining species richness patterns is a central issue in biogeography and macroecology. Several hypotheses have been proposed to explain the mechanisms driving biodiversity patterns, but the causes of species richness gradients remain unclear. In this study, we aimed to explain the impacts of energy, environmental stability, and habitat heterogeneity factors on variation of vertebrate species richness (VSR), based on the VSR pattern in China, so as to test the energy hypothesis, the environmental stability hypothesis, and the habitat heterogeneity hypothesis.

Methodology/Principal Findings

A dataset was compiled containing the distributions of 2,665 vertebrate species and eleven ecogeographic predictive variables in China. We grouped these variables into categories of energy, environmental stability, and habitat heterogeneity and transformed the data into 100×100 km quadrat systems. To test the three hypotheses, AIC-based model selection was carried out between VSR and the variables in each group and correlation analyses were conducted. There was a decreasing VSR gradient from the southeast to the northwest of China. Our results showed that energy explained 67.6% of the VSR variation, with the annual mean temperature as the main factor, which was followed by annual precipitation and NDVI. Environmental stability factors explained 69.1% of the VSR variation and both temperature annual range and precipitation seasonality had important contributions. By contrast, habitat heterogeneity variables explained only 26.3% of the VSR variation. Significantly positive correlations were detected among VSR, annual mean temperature, annual precipitation, and NDVI, whereas the relationship of VSR and temperature annual range was strongly negative. In addition, other variables showed moderate or ambiguous relations to VSR.

Conclusions/Significance

The energy hypothesis and the environmental stability hypothesis were supported, whereas little support was found for the habitat heterogeneity hypothesis.     

Dynamics of biomass partitioning in two competing meadow plant species

The optimal allocation theory predicts that growth is allocated between the shoot and the roots so that the uptake of the most limiting resource is increased. Allocation is dynamic due to resource depletion, interaction with competitors, and the allometry of growth. We assessed the effects of intra- and inter-specific competition on growth and resource allocation of the meadow species Ranunculus acris and Agrostis capillaris, grown in environments with high (+) or low (−) availability of light (L) and nutrients (N). We took samples twice a week over the 7 weeks experiment, to follow the changes in root-to-shoot ratios in plants of different sizes, and carried out a larger scale harvest at the end of the experiment. Of all the tested factors, availability of nutrients had the largest effect on the growth rate and shoot-to-root allocation in both species, although both competition and light had significant effects as well. The highest root-to-shoot ratios were measured from the L+N− treatment, and the lowest from the L−N+ treatment, as predicted by the optimal allocation theory. Competition changed resource allocation, but not always toward acquiring the resource that is most limiting to growth. We thus conclude that the greatest variation in shoot-to-root allocation was due to the resource availability and the effects of competition were small, probably due to low density of plants in the experiment.     

Effects of habitat fragmentation on provisioning rates, diet and breeding success in two species of tit (great tit and blue tit)

The aim of this study was to examine the effects of forest fragmentation on the ability of parent birds to provide their young with an adequate food supply. To examine whether prey population densities of the great tit (Parus major L.) and the blue tit (P. caeruleus L.) vary between study areas in different forest size classes we compared provisioning rates and chick diet and related these parameters to breeding success. We filmed 217 nests over two breeding seasons and collected data on frass fall as a general estimate of caterpillar availability. Nests which were attended by none or one parent only during filming (n = 46) were excluded from the analyses. In both years and for both species feeding rates were highest in the smallest fragments and lowest in the large forest. There was also a suggestion that differences in feeding rates between areas vary between years. We found no consistent tendency for prey size to change with forest size, although both species brought slightly smaller prey items to the nest in the smallest forest fragments and feeding rates correlated negatively with prey size. Caterpillars were the main item fed to nestlings, in both species. We found no evidence to suggest that either frass fall or the proportion of caterpillars in the diet varied with forest size. There was also no correlation between mean frass fall and the total number of caterpillars brought to the nests, in either species. Breeding success, as measured by clutch size, brood size, fledging weight and fledging success, did not differ between the small fragments and the large forest, in either species. There was also no relationship between provisioning rate (as concerns volume of prey fed to nestlings and the quality of chick diet) and breeding success parameters. In conclusion, this study does not suggest suboptimal foraging or breeding conditions in small fragments compared to a nearby large forest, for either species. Received: 23 June 1997 / Accepted: 29 December 1997     

Distinguishing area and habitat heterogeneity effects on species richness: Birds in Victorian buloke remnants

Abstract Resolving whether area per se or habitat heterogeneity has the greater influence in controlling species richness remains a controversial yet important question. Here we show that avian species richness of same-sized transects (1 ha) is independent of the remnant area (of buloke woodland) within which a transect is positioned. We also show that avi-faunal similarity of pairs of transects randomly placed within the largest remnants (≥ 48 ha) is not consistently related to either proximity (i. e. being within the same remnant) nor to physiognomic characteristics of the transects. We believe that much of the controversy over area/habitat heterogeneity effects is probably related to scalar issues and propose a protocol by which some resolution of the question might be reached. The protocol involves ‘zoom’ sampling in which successively larger transect sizes are used, and measures of faunal richness and habitat heterogeneity are made at these different grains of resolution. One of our intentions is to stimulate discussion on how heterogeneity might be measured when grains increase from typical transect sizes (ca 1 ha) up to much larger grains (ca 128 ha).     

The impact of two non-native plant species on native flora performance: potential implications for habitat restoration

Both Impatiens glandulifera and Fallopia japonica are highly invasive plant species that have detrimental impacts on native biodiversity in areas where they invade and form dense monocultures. Both species are weakly dependent on arbuscular mycorrhizal fungi (AMF) for their growth and, therefore, under monotypic stands, the AMF network can become depauperate. We evaluated the impact of I. glandulifera and F. japonica on the performance (expressed as shoot biomass) of three UK native species (Plantago lanceolata, Lotus corniculatus and Trifolium pratense) grown in soil collected from under stands of both invasive plants and compared to plants grown in soil from under stands of the corresponding native vegetation. All native species had a higher percentage colonisation of AMF when grown in uninvaded soil compared to the corresponding invaded soil. P. lanceolata and L. corniculatus had a higher biomass when grown in uninvaded soil compared to corresponding invaded soil indicating an indirect impact from the non-native species. However, for T. pratense there was no difference in biomass between soil types related to I. glandulifera, suggesting that the species is more reliant on rhizobial bacteria. We conclude that simply managing invasive populations of non-native species that are weakly, or non-dependent, on AMF is inadequate for habitat restoration as native plant colonisation and establishment may be hindered by the depleted levels of AMF in the soil below invaded monocultures. We suggest that the reintroduction of native plants to promote AMF proliferation should be incorporated into future management plans for habitats degraded by non-native plant species.     

Cannibalistic interactions in two co-occurring decapod species: Effects of density, food, alternative prey and habitat

Cannibalism is a potentially important factor in the regulation of populations in a range of habitats. The intensity of this biotic factor may be determined by both intra and interspecific interactions. Cancer pagurus and Porcellana platycheles are two co-occurring decapods on Atlantic rocky shores. In laboratory mesocosms, we investigated intra and intersize class cannibalistic and interspecific predatory behaviours in those species. We addressed the effects of prey and predator densities, food, starvation, alternative prey and habitat type. No agonistic behaviour was noted in P. platycheles, suggesting a non-aggressive co-existence between gregarious individuals. Predation of C. pagurus on P. platycheles was intense, possibly accounting for the spatial segregation observed in the natural environment. Cannibalism among C. pagurus juveniles was low and only on vulnerable prey (i.e. at moulting), suggesting a non-aggressive co-existence among juveniles. However, intersize class cannibalism in C. pagurus was intense (ontogenetic shift), possibly reflecting the juvenile-adult segregation in the natural environment. Prey and predator densities, food and habitat type strongly influenced this behaviour. Possible interference among cannibals was noted, with lower prey consumption at high predator density. Food supply alone had more effect on cannibalistic rate than did alternative prey (P. platycheles) and predator starvation. Structurally complex habitats (small pebble and Fucus serratus habitats) yielded higher prey survival than the sandy habitat, and the behaviours of both prey and cannibals reflected the small-scale spatial distribution of individuals in the wild. Intersize class cannibalism and interspecific agonistic relationships may account for the intertidal distribution of crab species at low tide.