Habitat selection,interspecific interactions and landscape composition

Summary I argue here that, from the perspective of any individual, most landscapes are composed of only three basic types of habitats. These are: (1) source habitat in which reproduction exceeds mortality and the expected per capita growth rate is greater than one; (2) sink habitat, in which limited, reproduction is possible but will not on average, compensate for mortality and the per capita rate of growth is between zero and one; and (3) unusable habitat, which comprises the matrix of all habitats that are never exploited by the species in question, and in which patches of source and sink habitats are embedded. Unlike earlier source-sink models, this model explicitly considers the effects that substituting one type of habitat for another has on the equilibrium size of a population and the interactions between species which can use both source and sink habitats. The model demonstrates that the equilibrium size of a species' population can sometimes be increased by substituting unusable habitat for sink habitat. Thus, even though the average patch quality in the landscape may be decreased, the overall quality of the landscape can increase. For two species with distinct habitat preferences, interactions between species can vary qualitatively as well as quantitatively as a function of the relative abundances of each of the habitat types. The model also shows that the interactions between species are particularly sensitive to the relative costs of moving between patches and sampling patches to determine their quality. Recent fragmentation of natural landscapes may increase the cost of searching for usable (source or sink) patches. Under some conditions, the interspecific interactions may be substantially more negative (competitive) than the interactions that evolved in the original natural landscape, further reducing population sizes and increasing the likelihood of competitive exclusion in fragmented modern landscapes.     

The cost of habitat selection in prairie voles: an empirical assessment using isodar analysis

The ideal free distribution assumes that habitat selection is without cost and predicts that fitness should be equal in different habitats. If habitat selection has a cost, then individuals should only move to another habitat when potential fitness in the new habitat exceeds that in the source habitat by an amount greater than the cost of habitat selection. We used isodar techniques to assess the cost of habitat selection. In an experimental landscape, we monitored density, movement, and reproductive success of adult female prairie voles, Microtus ochrogaster, in adjacent paired habitats with low and high cover. We tested the following hypotheses: (1) adult female prairie voles exhibited density-dependent habitat selection; (2) the cost of habitat selection was density-independent. Habitat quality based on population density and fitness of adult females was higher in high cover habitats. Net movement was from low cover to high cover habitats. The results indicated that adult female prairie voles exhibited density-dependent habitat selection. Furthermore, there was a significant cost of habitat selection, and the cost was density-independent.     

Do haddock select habitats to maximize condition?

Haddock Melanogrammus aeglefinus in the North sea increased their distributional range when more abundant, but this density dependent habitat selection (DDHS) explained only a small part of the year‐on‐year variation in distribution patterns. The condition of haddock was examined at 24 sites in the North Sea in August and September 2004 and related to their abundance, to examine if the ideal free distribution theory (IFD), which assumes that organisms select habitats that maximize their rate of food intake, can be used to explain this variation in large scale distribution patterns. At a given temperature, condition (hepato‐somatic index, I _H) was better at stations where haddock were most abundant. Therefore, haddock were not distributed perfectly according to the IFD in 2004. The positive correlation between abundance and I _H, however, indicated there was some habitat selection by haddock, as in the total absence of habitat selection no correlation between I _H and abundance, and no spatial variation in abundance was expected. DDHS may only explain a small part of the yearly variation in the distribution because haddock did not equalize and maximize their fitness at the scale of the North Sea. In addition, stable isotope analysis of muscle samples showed that haddock did not avoid competition for food when at high abundance by feeding at a lower or wider range of trophic levels.     

Habitat carrying capacity is reached for the European eel in a small coastal catchment: evidence and implications for managing eel stocks

1. Despite carrying capacity being one of the most important parameters in population management and modelling, we lack substantial evidence for habitat limitations on freshwater species. Here we tested the ideal free distribution (IFD) hypothesis using an indirect behaviour‐based method for small closed populations assuming that animals can effectively estimate habitat suitability and distribute themselves accordingly in time and space. 2. We analysed spatiotemporal variations in the density of the European eel Anguilla, a catadromous species with good colonisation abilities in a small coastal catchment in France. The general linear model used enabled us to test simultaneously the effect of temporal, macro‐ and meso‐scale habitat factors on the presence and abundance of eels at 30 sites over an 8‐year period. 3. Almost every site sampled had eels, whatever its location on the catchment and its habitat characteristics. Density estimates (overall mean ± SD of 0.40 ± 0.48 m^?2) were at the upper range of other values for European catchments. Moreover, eel densities were mainly influenced by the availability of suitable habitats (rocky substratum and instream cover), which suggests that their distribution reflects an IFD. 4. Despite marked variability in recruitment, the density of the oldest size‐class remained stable over the study, suggesting that density‐dependent mortality occurred, probably due to intraspecific competition for space and food and to predation. 5. These findings suggest that eel habitats are saturated in the Frémur. Therefore, we suggest that the mean abundance of eels observed could serve as a threshold value for other male‐dominated river stocks (provided they have a similar overall percentage of suitable habitats) that are common in small, low gradient streams on the north‐Atlantic coast of Europe.     

A generalized habitat matching rule

Summary When fitness of a resource-limited animal depends only on that individual's share of the total resource in a habitat patch and individuals are free to move to the patch where their gains are highest, population density matches resource availability under the simple assumption that individual fitness increases with resource use. Previous theory on habitat matching required the stronger assumption that individual fitness was directly proportional to (rather than monotonically increasing with) resource use. The basic theory suggests conditions under which population density empirically indicates habitat quality. Extensions of this basic theory apply when individuals that are free to move among resource patches interact by interfering with each other's resource extraction or by competing unequally. Analysis of existing models of such ideal free competition yields conditions for a single general matching rule in which the logarithm of crowding is a linear function of the logarithm of resource abundance. Double logarithmic plots of empirical data on habitat use and habitat quality based on this rule furnish possible graphical indicators of the occurrence and intensity of competition in nature.     

Movement between patches, unequal competitors and the ideal free distribution

Researchers have often commented on the ability of the original ideal free distribution (IFD) model to approximate observed animal distributions even though the critical assumption that competitors are of equal ability is usually violated. We provide an explanation by recognizing that animals will occasionally move between patches for reasons other than to simply maximize their resource payoffs, given perfect (i.e. ideal) information about the current payoff in each patch, and that these movements will continue to occur even after an equilibrium is reached. When such movements are incorporated into an unequal competitors IFD model, a single, stable distribution of each competitor type is predicted. This equilibrium will usually be characterized by under-matching of total competitive units relative to the distribution of resources (i.e. too few competitive units in the good patch). More importantly, it will often resemble the original, equal competitors IFD, in that total competitor numbers will come close to matching the distribution of resources. We argue that researchers claiming to have observed an IFD of equal competitors have actually observed this equilibrium distribution of unequal competitors. Our model predicts that the deviation from input-matching will usually be an under-matching of total competitor numbers relative to resources (i.e. too few competitors in the good patch). Examination of published data reveals that post-equilibrium movement between patches occurs frequently and, although the reported distributions are similar to those predicted by input-matching, under-matching is usually observed.     

Distribution of a naturally fluctuating ungulate population among heterogeneous plant communities: ideal and free?

1. Herbivore distribution is often assumed to follow the ideal free distribution (IFD) model. This assumes that organisms are omniscient about forage quality and availability within the area available to them and are free to move, with negligible cost, throughout this environment. If this were the case we would expect that, at lowest densities, all animals would be found in the best habitat patches, with less desirable habitats being occupied stepwise as population density increases. We test this using data from a naturally fluctuating population of feral Soay sheep. 2. We show that, although the distribution of individuals is correlated positively with food quality, in line with patterns reported for hill sheep in Scotland, their distribution does not conform to the predictions of the IFD model. We argue that it is the dynamic nature of their food resource that causes this departure from the predictions of the IFD model and make the case that the IFD model, in its unmodified form, is inappropriate for use in modelling distribution among patches containing dynamic resources.     

Linking Resource Matching and Dispersal

We study theoretically the effect of inter-habitat migration on the distribution of population sizes between two habitats, and compare this distribution with the expected ideal free distribution (IFD). Whenever emigration from the two habitats is asymmetric, or when there is a survival cost during migration, the resulting equilibrium distribution of population sizes deviates from the IFD. This result holds irrespective of emigration rule, even though a density-dependent fraction of emigrants generally produces a distribution closer to the IFD than a constant fraction of emigrants. Environmental stochasticity causes a linear relation between population sizes in the two habitats, with slope and intercept only identical to the IFD when net inter-habitat exchange is zero. The type and asymmetry of inter-habitat migration will influence how we should interpret data on population distribution in different habitats. The resulting resource matching is also critically contingent on the relative time-scales of population renewal and dispersal, and when population size is measured in relation to reproduction and dispersal. Therefore, data on population sizes cannot be used uncritically to assess habitat quality.     

Forecasting ecological and evolutionary strategies to global change: an example from habitat selection by lemmings

Ecologists and evolutionary biologists must develop theories that can predict the consequences of global warming and other impacts on Earth's biota. Theories of adaptive habitat selection are particularly promising because they link distribution and density with fitness. The evolutionarily stable strategy that emerges from adaptive habitat choice is given by the system's habitat isodar, the graph of densities in pairs of habitats such that the expectation of fitness is the same in each. We illustrate how isodars can be converted into adaptive landscapes of habitat selection that display the density‐ and frequency‐dependent fitness of competing strategies of habitat use. The adaptive landscape varies with the abundance of habitats and can thus be used to predict future adaptive distributions of individuals under competing scenarios of habitat change. Application of the theory to three species of Arctic rodents living on Herschel Island in the Beaufort Sea predicts changes in selection gradients as xeric upland increases in frequency with global warming. Selection gradients will become more shallow for brown lemming (Lemmus trimucronatus) and tundra vole (Microtus oeconomus) strategies that preferentially exploit mesic habitat. Climate change will cause selection gradients for the alternative strategy of using mostly xeric habitat to become much steeper. Meanwhile, the adaptive landscape for collared lemmings (Dicrostonyx groenlandicus), which specialize on xeric Dryas‐covered upland, will become increasingly convex. Changes in the adaptive landscapes thus predict expanding niches for Lemmus and Microtus, and a narrower niche for Dicrostonyx. The ability to draw adaptive landscapes from current patterns of distribution represents one of the few methods available to forecast the consequences of climate change on the future distribution and evolution of affected species.     

Ecological principles of species distribution models: the habitat matching rule

Most species distribution models (SDMs) assume that habitats are closed, stable and without competition. In that environmental context, it is ecologically correct to assume that members of a species will be distributed in direct relation to the suitability of the habitat, that is, according to the so‐called habitat matching rule. This paper examines whether it is possible to maintain the assumption of the habitat matching rule in the following circumstances: (1) when habitats are connected and organisms can move between them, (2) when there are disturbances and seasonal cycles that generate instability, and (3) when there is inter‐specific and intra‐specific competition. Here I argue that it is possible as long as the following aspects are taken into account. In open habitats at equilibrium, in which habitat selection and competition operate, the habitat matching rule can be applied in some conditions, while competition tends to homogenize the species distribution in other environmental contexts. In the latter case, two methods can be used to incorporate these effects into SDMs: new parameters can be incorporated into the response functions, or the occurrence of proportions of categories of individuals (adult/young, male/female, or dominant/subordinate species in guilds) can be used instead of the occurrence of organisms. The habitat matching rule is not fulfilled in non‐equilibrium environments. The solution to this problem lies in the design of SDMs with two strategies that depend on scale. Locally, the disequilibrium can be encapsulated using average environmental conditions, with sufficiently large cells (in the case of metapopulations) and/or long enough sampling periods (in the case of seasonal cycles). At coarse scales, the use of presence‐only models can in some cases avoid the destabilizing effect of catastrophic historical processes. The matching law is a strong assumption of SDMs because it is based on population ecology theory and the principle of evolution by natural selection.     

Density-dependent habitat selection in migratory passerines during stopover: what causes the deviation from IFD?

We studied the distribution of migratory warblers (genus: Sylvia) in poor and high quality habitat patches at a stopover site in the northern Negev, Israel. The purpose of our study was to test predictions based on the ideal free distribution (IFD) model by using a natural ecosystem which has a high turnover of individuals moving between unfamiliar foraging patches. We trapped birds in two groves of Pistacia atlantica embedded within a coniferous forest. The fruit-density ratio between these groves was 45:1. We compared bird density, body condition and habitat matching (the ratio between bird density and resource density) at the two sites. To analyse the data we integrated two approaches to density-dependent habitat selection: the isodar method and the habitat matching rule. As predicted by the IFD model, we found that habitat suitability decreased with bird density with a high correlation between warbler densities in the two habitat patches. Contrary to IFD predictions, warbler density in the poor patch was higher than expected by the habitat-matching rule. This habitat under-matching, had a cost: in the rich habitat the average energy gain per individual bird was higher than in the poor habitat. Further analysis suggests that the apparent habitat under-matching is not due to interference or differences in warbler competitive abilities. Therefore, we suggest that this migratory bird community is not at equilibrium because the birds possess imperfect knowledge of resource distribution. We propose that this lack of knowledge leads to free, but not ideal distributions of migrant birds in unfamiliar stop over sites.     

Density-dependent habitat selection: Testing the theory with fitness data

Summary According to density-dependent habitat selection theory, reproductive success should decline with increased density. Fitness should be similar between habitats if habitat selection follows an ideal free distribution; fitness should be dissimilar between habitats if habitat selection is modified by territorial behavior. I tested these assumptions by examining a variety of fitness estimates obtained from white-footed mice living in nest boxes in forest, forest edge and fencerow habitats in southwestern Ontario. As expected, mean litter size declined with increased population density. Litter sizes, adult longevity and the proportion of adult animals in breeding condition were not significantly different among the three habitats. The success at recruiting at least one offspring to the adult population and the number of recruits per litter were much greater in the forest than in either of the other two habitats. Fitness was thus unequal among habitats and the results confirm both assumptions of density-dependent habitat selection theory for territorial white-footed mice.     

Habitat assessment by parasitoids: consequences for population distribution

The ideal free distribution (IFD) is a stable distribution ofcompetitors among resource patches. For equally efficient competitors,equilibrium is reached when the per capita rate of intake equalizesacross patches. The seminal version of the IFD assumes omniscience,but populations may still converge toward the equilibrium providedthat competitors 1) accurately assess their environment by learningand 2) remain for an optimal (rate-maximizing) time on eachencountered patch. In the companion article (Tentelier C, DesouhantE, Fauvergue X. 2006. Habitat assessment by parasitoids: mechanismsfor patch time allocation. Behav Ecol. Forthcoming), it is shownthat the parasitoid wasp Lysiphlebus testaceipes adapts itsexploitation of aphid host colonies based on previous experience,in a manner consistent with these two conditions. We thereforepredicted that a randomly distributed population of initiallynaive wasps should converge toward the IFD. We tested this predictionby introducing 1300 L. testaceipes females into a 110-m² greenhousecontaining 40 host patches. Just after introduction, the parasitoidrate of gain was positively affected by host number and negativelyaffected by parasitoid number but, as predicted, these effectsvanished in the course of the experiment. Six hours after introduction,the expected rate of gain reached a constant. Surprisingly,this passage through equilibrium was not accompanied by a decreasein the coefficient of variation among gain rates or by a shiftfrom a random to an aggregated distribution of parasitoids.These results challenge our understanding of the link betweenindividual behavior and population distribution.     

Incorporating density dependence into the oviposition preference-offspring performance hypothesis

1. Although theory predicts a positive relationship between oviposition preferences and the developmental performance of offspring, the strength of this relationship may depend not only on breeding site quality, but also on the complex interactions between environmental heterogeneity and density-dependent processes. Environmental heterogeneity may not only alter the strength of density dependence, but may also fundamentally alter density-dependent relationships and the preference-performance relationship. 2. Here I present results from a series of field experiments testing the effects of environmental heterogeneity and density-dependent feedback on offspring performance in tree-hole mosquitoes. Specifically, I asked: (i) how do oviposition activity, patterns of colonization and larval density differ among habitats and among oviposition sites with different resources; and (ii) how is performance influenced by the density of conspecifics, the type of resource in the oviposition site, and the type of habitat in which the oviposition site is located? 3. Performance did not differ among habitats at low offspring densities, but was higher in deciduous forest habitats than in evergreen forest habitats at high densities. Oviposition activity and larval densities were also higher in deciduous forests, suggesting a weak preference for these habitats. 4. The observed divergence of fitness among habitats with increasing density may select for consistent but weak preferences for deciduous habitats if regional abundances vary temporally. This would generate a negative preference-performance relationship when population densities are low, but a positive relationship when population densities are high. 5. This study demonstrates that failure to recognize that fitness differences among habitats may themselves be density-dependent may bias our assumptions about the ecological and evolutionary processes determining oviposition preferences in natural systems.