Can migrants escape from density dependence?

Migration is thought to maximize growth by enabling individuals to escape from density dependence, but this has rarely been tested at the individual level in natural populations. We employed linear mixed modeling of the spacing between consecutive scale growth rings to reconstruct individual growth profiles of a paradigmatic fish migrant, the sea trout (Salmo trutta) and related these to estimates of year class strength over a 13‐year period. Variation in scale growth was 1.3 times greater among individuals than within individuals in freshwater and 10 times greater at sea. Scale growth was inversely related to year class strength, both in freshwater (before migration) and at sea (after migration). Competition for patchily distributed resources is the most plausible explanation of the negative density‐dependent growth observed in freshwater and, to a lesser extent, in the marine environment. Our study provides some of the strongest evidence for a role of density dependence in determining partial migrations because although migrants can maximize growth by moving into the sea, they do not appear to become free from density dependence constraints completely. This has implications for conservation and suggests that sea trout and other anadromous fish displaying partial migrations may not be best managed on a river by river basis, but rather from a broader, coastal perspective.     

Partial migration in expanding red deer populations at northern latitudes – a role for density dependence?

Partial migration is common in ungulates living in highly seasonal environments. Typically, at higher latitudes, this involves movement between high elevation summer areas used during breeding and lowland areas with less snow used during winter. Snow depth is regarded the main cause of migration to low elevation, but it is less clear why deer migrate to high elevation in spring. The forage maturation hypothesis explains the upward migration due to plant phenology. We here present also an alternative and non‐exclusive hypothesis, that deer migrate uphill in summer to escape competition due to the high density in winter areas (the competition avoidance hypothesis). We also suggest that social fences may play a role at high population density. Based on a unique study of 141 GPS‐marked red deer from seven regions covering the main distribution in Norway, we found that the proportion of migrants in the populations varied from 38% to 100%. Migration was more common in areas with a diverse topography, i.e. for areas with access to high elevation. Further, we found evidence that migration was negatively density dependent, and that fall migration was delayed at high density. We suggest that a combination of avoidance of competition in high density winter ranges, social fencing during summer in addition to the forage maturation and predation risk avoidance hypotheses, is needed to explain migration patterns of northern ungulates.     

What's your move? Movement as a link between personality and spatial dynamics in animal populations

Recent studies have established the ecological and evolutionary importance of animal personalities. Individual differences in movement and space‐use, fundamental to many personality traits (e.g. activity, boldness and exploratory behaviour) have been documented across many species and contexts, for instance personality‐dependent dispersal syndromes. Yet, insights from the concurrently developing movement ecology paradigm are rarely considered and recent evidence for other personality‐dependent movements and space‐use lack a general unifying framework. We propose a conceptual framework for personality‐dependent spatial ecology. We link expectations derived from the movement ecology paradigm with behavioural reaction‐norms to offer specific predictions on the interactions between environmental factors, such as resource distribution or landscape structure, and intrinsic behavioural variation. We consider how environmental heterogeneity and individual consistency in movements that carry‐over across spatial scales can lead to personality‐dependent: (1) foraging search performance; (2) habitat preference; (3) home range utilization patterns; (4) social network structure and (5) emergence of assortative population structure with spatial clusters of personalities. We support our conceptual model with spatially explicit simulations of behavioural variation in space‐use, demonstrating the emergence of complex population‐level patterns from differences in simple individual‐level behaviours. Consideration of consistent individual variation in space‐use will facilitate mechanistic understanding of processes that drive social, spatial, ecological and evolutionary dynamics in heterogeneous environments.     

Does variation in patch use among butterfly species contribute to nestedness at fine spatial scales?

A nested pattern occurs whenever the species observed in depauperate habitat patches are a subset of those found in more species‐rich patches. Ecologists have documented many instances of nestedness caused by population‐level processes such as colonization and extinction at biogeographic scales. However, few researchers have examined whether nestedness may exist at fine scales due to the ways in which individual organisms discriminate among potential habitat patches. In 1999, we experimentally fragmented an old‐field habitat into patches of varying size to test whether nestedness could exist on a fine spatial scale. Five treatments of differing patch size were replicated five times in a Latin square design by selectively mowing 15×15 m² plots within an old‐field (patch areas: 225, 180, 135, 90, and 45 m²). Specifically, we tested whether butterflies foraging within a network of patches differing in area conformed to a nested subset structure. We also classified species according to (1) their flight height while foraging (high or low), and (2) their adult habitat breadth (ubiquitous, general, or restricted) to determine whether nestedness could be explained by difference in species’ tendency to discriminate among patches differing in area.
We found significant evidence that a community of foraging Lepidoptera conformed to a nested subset structure based on the difference between the observed nestedness within the butterfly community and the nestedness obtained from randomly generated species presence/absence matrices. Poisson regression analyses demonstrated that high‐flying, habitat‐restricted species avoided the smallest patches (90 and 45 m²) in favor of larger remnants, whereas low‐flying, habitat generalists used all patch sizes. Thus, our study is one of the first to demonstrate that nestedness among species subsets can be observed at fine spatial scales (within a single 1.5 hectare field) and may be maintained by species behavioral differences: discriminating species (i.e. high‐flying, habitat restricted) avoided the smallest patches, and less discriminating species (i.e. low‐flying, ubiquitous) were distributed throughout the field without regard to patch size. Our results also suggest that nestedness should be viewed as yet another scalar pattern in ecology, generated by variation in patch use by individuals at fine‐scales as well as the more traditionally invoked processes of extinction and colonization of species at broad‐scales.     

Can altruism evolve in purely viscous populations?

Summary Limited dispersal is often thought to facilitate the evolution of altruism by increasing the degree of relatedness among interacting individuals. Limited dispersal can have additional effects, however, such as local population regulation, that inhibits the evolution of altruism. Many models of structured populations assume that a viscous stage of the life cycle alternates with a global mixing stage, which allows the advantages of interactions among close relatives without the disadvantages of local population regulation. Here we analyse a computer simulation model of pure population viscosity, in which offspring are always deposited close to parents and no global mixing stage exists. As expected, limited dispersal generates a high coefficient of relatedness among interacting individuals. Patches of altruists, however, are unable to export their productivity to other regions of the landscape and are easily invaded by selfish types from neighbouring patches. Unlike models of alternating viscosity, in which high relatedness and local population regulation can be decoupled, these two opposing effects are inextricably linked in purely viscous populations, which therefore are not conducive to the evolution of altruistic traits.     

Can balancing selection on MHC loci counteract genetic drift in small fragmented populations of black grouse?

The ability of natural populations to adapt to new environmental conditions is crucial for their survival and partly determined by the standing genetic variation in each population. Populations with higher genetic diversity are more likely to contain individuals that are better adapted to new circumstances than populations with lower genetic diversity. Here, we use both neutral and major histocompatibility complex (MHC) markers to test whether small and highly fragmented populations hold lower genetic diversity than large ones. We use black grouse as it is distributed across Europe and found in populations with varying degrees of isolation and size. We sampled 11 different populations; five continuous, three isolated, and three small and isolated. We tested patterns of genetic variation in these populations using three different types of genetic markers: nine microsatellites and 21 single nucleotide polymorphisms (SNPs) which both were found to be neutral, and two functional MHC genes that are presumably under selection. The small isolated populations displayed significantly lower neutral genetic diversity compared to continuous populations. A similar trend, but not as pronounced, was found for genotypes at MHC class II loci. Populations were less divergent at MHC genes compared to neutral markers. Measures of genetic diversity and population genetic structure were positively correlated among microsatellites and SNPs, but none of them were correlated to MHC when comparing all populations. Our results suggest that balancing selection at MHC loci does not counteract the power of genetic drift when populations get small and fragmented.     

Temporal β-diversity of zooplankton at various time scales in a small mountain lake

Limnology - To examine how compositional changes in a community vary depending on time scales, we estimated temporal β-diversity of zooplankton in Lake Hataya Ohnuma, a small lake in Yamagata,...     

Are raccoons limited by the availability of breeding resources? Evidence of variability in fecundity at fine spatial scales

Despite their behavioral plasticity, recent research suggests raccoons (Procyon lotor) exhibit variability in demography and genetic structure among individual habitat patches in landscapes heavily affected by anthropogenic land use. Consequently, elucidation of vital rates at fine-spatial scales is needed to implement appropriate management strategies for this species. To evaluate the degree of variability in productivity that exists among individual habitat patches, we collected reproductive (n = 170) and cementum annuli age data (n = 383) for raccoons occupying 30 forest patches varying in local and landscape-level habitat attributes within a highly fragmented agricultural ecosystem. Across all females sampled, pregnancy rates averaged 85% but were highly variable among ages (range: 47–100%). Average litter sizes ranged from 3.2 to 4.7, but did not differ as a function of age. At the landscape-level, we observed significant variability among habitat patches in the total number of offspring produced (range: 0–80), indicating that individual patches vary in their contribution to the overall size and stability of the global (landscape-level) population. Variability in productivity among habitat patches primarily was driven by local differences in the availability of denning resources, likely because of the influence of this variable on variability in the number, age structure, and reproductive rate of females. Our results suggest that in agricultural ecosystems, the reproductive potential and temporal stability of raccoons within individual habitat fragments is inextricably linked to the density of tree cavities. © 2012 The Wildlife Society.     

Does personality in small rodents vary depending on population density?

Personality means an individual's unique way of behaving and reacting to the environment. It is a stable and heritable trait, which is expressed consistently in different situations. The aim of our study was to develop novel tests to depict the personality structure of the bank vole Myodes glareolus, and to determine if the phase of the population cycle, i.e. population density, affects personality. We focused on some central aspects of bank vole behaviour: mobility, risk taking, exploratory behaviour, dominance, and aggressive behaviour towards pups. These behaviours were chosen because they directly affect bank vole survival or fitness or are classified as important factors of personality in other species. In total, 192 males from different populations went through four behavioural tests, in which 20 variables were measured. The tests were repeated after 3 weeks, which verified that all traits were stable, i.e. repeatable between trials. Three personality compounds emerged, named extroversion, novelty seeking and infanticide. Extroversion included dominance and mobility, while novelty seeking consisted of risk taking and exploration. Infanticide encompassed all indices measuring harmful behaviour towards pups. Mobility and dominance were connected, possibly because both seem to depend on condition. Time spent in captivity increased extroversion, which may be explained by good food, stable conditions and acclimation to strong social cues. Novelty seeking was connected to repeatability which could mean that novelty avoiding individuals adjust their behaviour to match new environments. Population density affected the infanticide trait but not novelty seeking or extroversion.     

Long-term trends in the population size of king penguins at Crozet archipelago: environmental variability and density dependence?

We examined the growth rate of the breeding population of king penguins of Crozet archipelago over 41 years. Most colonies showed positive growth rates from the 1960s. However, there was evidence for a decrease in the larger colonies since the early 1990s, and for lower growth rates in the smaller colonies during the 1990s. The overall population size was estimated using log linear models, and the average annual growth rate was +6.3% for the 41-year period. Four change points were detected in the annual growth rate: +21.1% during 1978–1985, +4.3% during 1985–1995, –19.2% during 1995–1999, and +10.9% during 1999–2003. Time-series analyses of the population-size estimates and the relationship between growth rate and population size both indicated density-dependence in population growth rate. Variations in population sizes are also discussed in relation to environmental fluctuations. Our results suggest that important changes occurred over the past 10 years.     

Can video images imitate real stimuli in animal behaviour experiments?

The use of video images in place of natural stimuli in animal behaviour experiments is reviewed. Unlike most other artificial means of stimulus presentation, video stimuli can depict complex moving objects such as other animals, preserving the temporal and spatial patterns of movement precisely as well as colour and sounds for repeated playback. Computer editing can give flexibility and control over all elements of the stimulus. A variety of limitations of video image presentation are also considered. Televisions and video monitors are designed with human vision in mind, and some non-human animals that differ in aspects of visual processing such as their colour vision, critical flicker-fusion threshold, perception of depth and visual acuity, may perceive video images differently to ourselves. The failure of video stimuli to interact with subjects can be a drawback for some studies. For video to be useful, it is important to confirm that the subject animal responds to the image in a comparable way to the real stimulus, and the criteria used to assess this are discussed. Finally, the contribution made by video studies to date in the understanding of animal visual responses is considered, and recommendations as to the future uses of video are made.     

Can small wildlife conservancies maintain genetically stable populations of large mammals? Evidence for increased genetic drift in geographically restricted populations of Cape buffalo in East Africa

The Cape buffalo (Syncerus caffer caffer) is one of the dominant and most widespread herbivores in sub‐Saharan Africa. High levels of genetic diversity and exceptionally low levels of population differentiation have been found in the Cape buffalo compared to other African savannah ungulates. Patterns of genetic variation reveal large effective population sizes and indicate that Cape buffalos have historically been interbreeding across considerable distances. Throughout much of its range, the Cape buffalo is now largely confined to protected areas due to habitat fragmentation and increasing human population densities, possibly resulting in genetic erosion. Ten buffalo populations in Kenya and Uganda were examined using seventeen microsatellite markers to assess the regional genetic structure and the effect of protected area size on measures of genetic diversity. Two nested levels of genetic structure were identified: a higher level partitioning populations into two clusters separated by the Victoria Nile and a lower level distinguishing seven genetic clusters, each defined by one or two study populations. Although relatively small geographic distances separate most of the study populations, the level of genetic differentiation found here is comparable to that among pan‐African populations. Overall, correlations between conservancy area and indices of genetic diversity suggest buffalo populations inhabiting small parks are showing signs of genetic erosion, stressing the need for more active management of such populations. Our findings raise concerns about the future of other African savannah ungulates with lower population sizes and inferior dispersal capabilities compared with the buffalo.     

Seed viability in declining populations of Caladenia rigida (Orchidaceae): are small populations doomed?

Despite comparatively good rates of pollination and seed production, some populations of the endangered terrestrial orchid Caladenia rigida continue to decline. To determine whether seed quality may be limiting reproductive potential, we assessed seed viability among declining populations of C.?rigida (in the southern part of its distribution) and among populations that are regarded as stable (in the northern part of its distribution). We also compared differences in seed viability to plant traits, population size and habitat characteristics (soil properties, canopy cover, presence of proximate vegetation). Seed capsules from southern populations were significantly smaller, with only 9% of seeds being viable, compared to 36% in capsules from northern populations. Soil phosphorus concentrations differed between regions, but other habitat characteristics did not correlate with seed viability. Using calculations based on seedling recruitment data from other Caladenia species, we predict that seed output is insufficient to ensure the long-term persistence of the smallest C.?rigida populations.     

Does predation risk influence habitat use by northern redbelly dace Phoxinus eos at different spatial scales?

This study investigated the relationship between spatial variations in predation risk and abundance of northern redbelly dace Phoxinus eos at both macroscale (littoral v. pelagic zones) and microscale (structured v. open water habitats in the littoral zone) of Canadian Shield lakes. Minnow traps were placed in both structured and open water habitats in the littoral zone of 13 Canadian Shield lakes, and estimates of the relative predation risk of P. eos in both the pelagic and the littoral zones were obtained from tethering experiments. Results showed that (1) the mean abundance of P. eos in the littoral zone was positively correlated with the relative predation risk in the pelagic zone, (2) P. eos preferentially used structured over open water habitats in the littoral zone and (3) this preference was not related to the relative predation risk in the littoral zone but decreased as the relative predation risk increased in the pelagic zone. At the lake level, these results support the hypothesis that P. eos enter the littoral zone to avoid pelagic piscivores. At the littoral zone level, the results do not necessarily contradict the widely accepted view that P. eos preferentially use structured over open habitats to reduce their predation risk, but suggest that flexibility in antipredator tactics ( e.g. shelter use v. shoaling) could explain the spatial distribution of P. eos between structured and open water habitats.     

Geographical separation of two Ulex species at three spatial scales: does competition limit species' ranges?

It is a common assumption that species' ranges are limited by their physiological tolerances to climatic factors, Biotic factors, such as competition, are rarely considered. We investigated the distributions of Ulex minor and U. gallii at three spatial scales from geographic ranges to individual heaths - to examine whether the species are negatively associated, as predicted by the hypothesis that the ranges of the species are limited by competition with each other. Distribution maps for the British Isles and France (100 400 km² survey units) show the two species have largely separated, but slightly overlapping ranges. A region of range overlap on the heaths of Dorset, southern England was mapped using 4 ha survey squares. There was strong negative association between the species, and the heaths could be divided into zones where one species was dominant. There was some indication of edaphic differences between the U. minor -dominated zones and the U. gallii zones. The few heaths where the species co-occurred were surveyed using 4 m² quadrats placed along transects. Usually one species was widespread over the heath, while the other occurred in patches. The species were strongly negatively associated in all transects. Therefore, the two species showed strong negative associations at three mapping scales. Apparent co-occurrences detected at one spatial scale largely disappeared when species were mapped at finer scales, emphasising the fractal nature of distributions. This provides evidence that the distributions of the two species are not independent and that they cannot coexist, and therefore that their ranges are limited by competition. Over their ranges, competitive superiority is probably determined by the climate. At the range boundaries in the region of overlap, climate is not important, but other physical factors such as edaphic conditions may determine the outcome of competition.     

Does tiger shark predation risk influence foraging habitat use by bottlenose dolphins at multiple spatial scales?

Prey availability and predation risk are important determinants of habitat use, but their importance may vary across spatial scales. In many marine systems, predator and prey distributions covary at large spatial scales, but do no coincide at small spatial scales. We investigated the influences of prey abundance and tiger shark ( Galeocerdo cuvier ) predation risk on Indian Ocean bottlenose dolphin ( Tursiops aduncus ) habitat use across multiple spatial scales, in Shark Bay, Western Australia. Dolphins were distributed between deep and shallow habitats and across microhabitats within patches approximately proportional to prey density when shark abundance was low. When shark abundance was high, foraging dolphins greatly reduced their use of dangerous, but productive, shallow patches relative to safer deep ones. Also, dolphins reduced their use of interior portions of shallow patches relative to their edges, which have higher predator density but lower intrinsic risk (i.e. a higher probability of escape in an encounter situation). These results suggest that predation risk and prey availability influence dolphin habitat use at multiple spatial scales, but intrinsic habitat risk, and not just predator encounter rate, is important in shaping dolphin space use decisions. Therefore, studies of habitat use at multiple spatial scales can benefit from integrating data on prey availability and the subcomponents of predation risk.     

Can zebra mussels change stratification patterns in a small reservoir?

Colonization and proliferation of zebra mussel (Dreissena polymorpha) population in Hargus lake, a small thermally stratified reservoir in Ohio, U.S.A., caused a significant increase in water clarity and a remarkable decrease in phytoplankton biomass during the period from 1993 to 1995. Increased light penetration and reduced organic matter loading to the meta-and hypolimnion were reflected in the lake stratification patterns, particularly in the temperature and oxygen profiles in the metalimnion. The meta- and hypolimnetic water temperature increased significantly over three years, irrespective of variation in surface water temperature. The epilimnion depth (mixing depth) increased by about the same magnitude as did the average Secchi depth. However, the total heat content of the lake did not show a consistent trend to increasing zebra mussel abundance, as it was largely influenced by the temperature of the large water volumes near the surface, which were in turn affected by weather conditions. Concurrent with the thermal structure change, the dissolved oxygen structure also changed over three years, though to a lesser extent. The changes in oxygen stratification pattern were reflected by increased oxygen concentrations in the metalimnion and a lowered depth of 3 mg l^–1 DO isopleth. These observed changes were likely attributed to increased water mixing depth, metalimnion photosynthesis and reduced oxygen consumption by organic matter. With increased epilimnion thickness and improved oxygen conditions in the metalimnion, the habitable space for aquatic macro-organisms (including fish) expanded substantially. Our results suggest that the indirect impacts of zebra mussels on small lake stratification patterns may have much broader implications than do the direct trophic interactions to the whole ecosystem.