Summary 1.Describing distribution and abundance is requisite to exploring interactions between organisms and their environment. Recently, the resource selection function (RSF) has emerged to replace many of the statistical procedures used to quantify resource selection by animals. 2. A RSF is defined by characteristics measured on resource units such that its value for a unit is proportional to the probability of that unit being used by an organism. It is solved using a variety of techniques, particularly the binomial generalized linear model. 3. Observing dynamics in a RSF -obtaining substantially different functions at different times or places for the same species -alerts us to the varying ecological processes that underlie resource selection. 4. We believe that there is a need for us to reacquaint ourselves with ecological theory when interpreting RSF models. We outline a suite of factors likely to govern ecologically based variation in a RSF. In particular, we draw attention to competition and density-dependent habitat selection, the role of predation, longitudinal changes in resource availability and functional responses in resource use. 5. How best to incorporate governing factors in a RSF is currently in a state of development; however, we see promise in the inclusion of random as well as fixed effects in resource selection models, and matched case-control logistic regression. 6. Investigating the basis of ecological dynamics in a RSF will allow us to develop more robust models when applied to forecasting the spatial distribution of animals. It may also further our understanding of the relative importance of ecological interactions on the distribution and abundance of species.
Summary1. Density is a fundamental driver of many ecological processes including habitat selection. Theory on density-dependent habitat selection predicts that animals should be distributed relative to profitability of habitat, resulting in reduced specialization in selection (i.e. generalization) as density increases and competition intensifies. 2. Despite mounting empirical support for density-dependent habitat selection using isodars to describe coarse-grained (interhabitat) animal movements, we know little of how density affects fine-grained resource selection of animals within habitats [e.g. using resource selection functions (RSFs)]. 3. Using isodars and RSFs, we tested whether density simultaneously modified habitat selection and within-habitat resource selection in a rapidly growing population of feral horses (Equus ferus caballus Linnaeus; Sable Island, Nova Scotia, Canada; 42% increase in population size from 2008 to 2012). 4. Among three heterogeneous habitat zones on Sable Island describing population clusters distributed along a west-east resource gradient (west-central-east), isodars revealed that horses used available habitat in a density-dependent manner. Intercepts and slopes of isodars demonstrated a pattern of habitat selection that first favoured the west, which generalized to include central and east habitats with increasing population size consistent with our understanding of habitat quality on Sable Island. 5. Resource selection functions revealed that horses selected for vegetation associations similarly at two scales of extent (total island and within-habitat zone). When densities were locally low, horses were able to select for sites of the most productive forage (grasslands) relative to those of poorer quality. However, as local carrying capacity was approached, selection for the best of available forage types weakened while selection for lower-quality vegetation increased (and eventually exceeded that of grasslands). 6. Isodars can effectively describe coarse-grained habitat selection in large mammals. Our study also shows that the main predictions of density-dependent habitat selection are highly relevant to our interpretation of RSFs in space and time. At low but not necessarily high population size, density will be a leading indicator of habitat quality. Fitness maximization from specialist vs. generalist strategies of habitat and resource selection may well be apparent at multiple spatial extents and grains of resolution.
Fine-scale spatial variation in genetic relatedness and inbreeding occur across continuous distributions of several populations of vertebrates; however, the basis of observed variation is often left untested. Here we test the hypothesis that prior observations of spatial patterns in genetics for an island population of feral horses (Sable Island, Canada) were the result of spatial variation in population dynamics, itself based in spatial heterogeneity in underlying habitat quality. In order to assess how genetic and population structuring related to habitat, we used hierarchical cluster analysis of water sources and an indicator analysis of the availability of important forage species to identify a longitudinal gradient in habitat quality along the length of Sable Island. We quantify a west-east gradient in access to fresh water and availability of two important food species to horses: sandwort, Honckenya peploides, and beach pea, Lathyrus japonicas. Accordingly, the population clusters into three groups that occupy different island segments (west, central, and east) that vary markedly in their local dynamics. Density, body condition, and survival and reproduction of adult females were highest in the west, followed by central and east areas. These results mirror a previous analysis of genetics, which showed that inbreeding levels are highest in the west (with outbreeding in the east), and that there are significant differences in fixation indices among groups of horses along the length of Sable Island. Our results suggest that inbreeding depression is not an important limiting factor to the horse population. We conclude that where habitat gradients exist, we can anticipate fine-scale heterogeneity in population dynamics and hence genetics.
Movement away from an area or social group in response to increasing density (density-dependent dispersal) is known for most species; why it evolves is fundamental to our understanding of ecology and evolution. However, we have yet to fully appreciate how individuals of varying conditions (e.g., age and sex) might differently consider effects of density (quorum) when deciding to disperse or not, and scale dependence in their sense of quorum. We tracked movements of all individuals of a naturalized population of feral horses (Equus ferus caballus; Sable Island National Park Reserve, Nova Scotia, Canada) during a period of rapid population growth (N increased from 375 to 484 horses from 2008 to 2010). Permanent dispersal from breeding groups (bands) was positively density dependent for all age and sex categories with respect to local density (horses/km2, bounded by the 99th percentile of individual movements [8000 m]), but was negatively and positively density dependent for males and females, respectively, in relation to group (band) size. Dispersal was generally female biased, with the exception of foals which moved with their mothers (no sex effect), and for yearlings and subadults when band sizes were smaller than average, in which case males dispersed at higher rates than females. Dispersal distance was positively related to local density. We conclude that dispersal rate can be both positively and negatively density dependent for feral horses, contingent on the state of individuals and the scale at which quorum with respect to choosing to disperse or not is assessed. Scale effects and interactions of density-dependent and sex- and age-biased dispersal may have both ecological and evolutionary consequences through effects on resource and mate competition.
Identifying the existence of population sinks is critical for conservation and management. However, because of density‐dependent dispersal, sinks can sometimes be masked by immigration events, especially during phases of population growth. We present a large‐scale, empirical demonstration of within‐population source‐sink dynamics using the feral horses (Equus ferus caballus) of Sable Island National Park Reserve, Nova Scotia, Canada, as a model. We tracked the fates and movements of 98.7% of the female population (n = 190–237) across 3 demographic clusters (subunits) during a period of rapid population growth (2008–2010; 24.7% increase in density). All subunits experienced increases in population size each year (λ > 1.0). Our individual‐based analysis showed that western Sable Island, where water availability was greatest, behaved as a source and would have grown with or without immigration in all years. However, the central (and fastest growing subunit) would have declined from 2008–2009 (λ = 0.951) without immigration. Further, the eastern subunit would have declined in 2 intervals (λ = 0.932, 0.999) without immigration. Our study demonstrates that the propensity of habitat to act as a sink can be masked during a period of population growth because of density‐dependent immigration from adjacent habitats. These findings present a caution to managers charged with conserving wide‐ranging species with long population cycles for which effects of immigration on local population growth rate can be difficult to isolate using standard methods of enumeration. © 2013 The Wildlife Society.
Young animals in a broad range of taxa solicit care from their parents with begging displays, which are used at least partly for competition among brood or litter mates. The effect of other begging offspring on an individual’s own begging display varies across studies, however, increasing its intensity in some, but not changing, or even decreasing it, in others. One possible reason for this discrepancy is that the potential pay‐off for more intense begging depends not only on how intensely an individual’s brood or littermates are begging, but also on how long that individual has been without food. Surprisingly, however, no studies have focused on how begging responses vary when both factors are varied simultaneously. We therefore examined how nestling tree swallows, Tachycineta bicolor, respond to nestmates in relation to both their own hunger levels and the begging intensity of nestmates. During a period of food deprivation, we played focal nestlings parental contact calls either alone (control) or with the begging calls of a nestling deprived of food for 30–50 (low intensity) or 100–110 min (high intensity). Nestlings called for longer in response to the low‐intensity playback, but, surprisingly, not in the high‐intensity playback, in which they instead delayed the onset of their calling. All these responses to nestmates were independent of how long the responding nestling had been deprived of food. Thus, even in the seemingly intensely competitive environment of a passerine brood, offspring do not necessarily respond to nestmates with escalation. This may be because de‐escalation is the best competitive option in some circumstances, or because begging has other functions besides advertisement of individual need and competition over food allocation. Certainly, the results illustrate the need for studies of how nestmate interactions vary across a broad range of contexts.
Abstract:Anthropogenic edge effects, whereby disturbance strength increases in proximity to ecotone boundaries, are known to strongly affect individual species but we lack a general understanding of how they vary by species, disturbance type and regional context. We deployed 46 camera-trap stations for a total of 3545 trap-days at two sites in Sulawesi, Indonesia, obtaining 937 detections of five vertebrate species. Anoa (Bubalus spp.) were more abundant near edges, booted macaque (Macaca ochreata) and red jungle fowl (Gallus gallus) were less abundant near edges, and edges did not impact Sulawesi warty pig (Sus celebensis) or Malay civet (Viverra tangalunga). But the relative importance of habitat disturbance from agriculture, roads and villages differed for each species, and edge-induced disturbances varied not only in magnitude but also in direction between the study areas. In the strongest instance, macaque local abundance was 3.5 times higher near villages than it was 3 km into the forest in one reserve, but 2.8 times higher 3 km into the forest than near villages in the other reserve. Our results suggest that responses to habitat edges among species and edge types are idiosyncratic, and that landscape-level context can strongly alter the influence of local disturbance on biodiversity.
There is a need to achieve sustainable agricultural production to secure food, fiber, and fuel for a growing global population. Climate-smart (CS) actions (no-till and cover crops) can reduce carbon emissions and promote soil organic carbon (SOC) storage. Contemporary voluntary carbon markets provide producers with a monetary incentive to adopt CS actions. However, SOC–yield dynamics under CS actions are not well known, making it difficult for producers to judge whether additional income from carbon credits will offset potential losses to yield income. We designed a SOC–yield framework that captures SOC–yield–income dynamics under traditional (reduced tillage, no cover crops) and CS actions. Using a modified structured decision-making approach, we applied the framework to a case study in which producers aim to increase income by selling carbon credits after adopting CS actions. Specifically, we demonstrated how to balance tradeoffs between yield and carbon credit income that arise from tillage and winter cover crop actions (cereal rye, Secale cereale L. and crimson clover, Trifolium incarnatum L.) in a soybean (Glycine max L.) production system in Mississippi. Results indicated that a producer could minimize losses to net yield income by adopting no-till if already using cover crops. There was also evidence that carbon credit income could offset losses to yield income when adopting CS in place of traditional actions. Identifying risks to yield income and SOC storage can help design carbon neutrality policies that have minimum impact on a producer’s income. History: This paper has been accepted for the Decision Analysis Special Issue on Further Environmental Sustainability. Funding: This work was supported by the USDA-ARS [Grants 58-0200-0-002 (Advancing Agricultural Research) and 58-6001-8-003] and the USDA National Institute of Food and Agriculture [McIntire Stennis Project 1020959]. Supplemental Material: The online appendix is available at https://doi.org/10.1287/deca.2023.0478 .
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