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Vander Wal, E.

Publications and source records attributed to Vander Wal, E..

27 records · Page 2Linked to original sources

Death comes for us all: an interplay of habitat selection, movement, and social behavior relate to cause specific mortality among grey wolves

Avoiding death infects biological processes, including behavior. Habitat selection, movement, and sociality are highly flexible behaviors that influence the mortality risks and subsequent fitness of individuals. In the Anthropocene, animals are experiencing increased risks from direct human causes and increased spread of infectious diseases. Using integrated step selection analysis, we tested how the habitat selection, movement, and social behaviors of grey wolves vary as an individual dies due to humans or canine distemper virus (CDV) and how those behaviors may vary in the lead up to death. Behaviors that changed prior to death were strongly related to how an animal eventually died. Wolves killed by humans moved slower than wolves that survived and selected to be nearer roads closer in time to their death. Wolves that died due to CDV moved progressively slower as they neared death and reduced their avoidance of wet habitats. All animals, regardless of dying or not maintained strong selection to be near packmates across time, which seemingly contributed to disease dynamics in the packs that became infected with CDV. Habitat selection, movement, and sociality interact to put individuals and groups at greater risks, influencing their cause-specific mortality. Lay SummaryNot much is known about behaviors prior to death in wild animals. Grey wolves killed by humans selected to be in riskier areas increasingly prior to their deaths. Wolves that died due to disease moved slower and changed their habitat selection to be in areas with more water as they became sicker. Sick wolves also continued to select for packmates, increasing the chances that the whole pack would succumb to the disease.

animal behavior and cognition↗

Solving the Sample Size Problem for Resource Selection Analysis

O_LISample size sufficiency is a critical consideration for conducting Resource-Selection Analyses (RSAs) from GPS-based animal telemetry. Cited thresholds for sufficiency include a number of captured animals M [≥] 30 and as many relocations per animal N as possible. These thresholds render many RSA-based studies misleading if large sample sizes were truly insufficient, or unpublishable if small sample sizes were sufficient but failed to meet reviewer expectations. C_LIO_LIWe provide the first comprehensive solution for RSA sample size by deriving closed-form mathematical expressions for the number of animals M and the number of relocations per animal N required for model outputs to a given degree of precision. The sample sizes needed depend on just 2 biologically meaningful quantities: habitat selection strength and a novel measure of landscape complexity, which we define rigorously. The mathematical expressions are calculable for any environmental dataset at any spatial scale and are applicable to any study involving resource selection (including sessile organisms). We validate our analytical solutions using globally relevant empirical data including 5,678,623 GPS locations from 511 animals from 10 species (omnivores, carnivores, and herbivores living in boreal, temperate, and tropical forests, montane woodlands, swamps, and arctic tundra). C_LIO_LIOur analytic expressions show that the required M and N must decline with increasing selection strength and increasing landscape complexity, and this decline is insensitive to the definition of availability used in the analysis. Our results contradict conventional wisdom by demonstrating that the most biologically relevant effects on the utilization distribution (i.e. those landscape conditions with the greatest absolute magnitude of resource selection) can often be estimated with far fewer data than is commonly assumed. C_LIO_LIWe identify several critical steps in implementing these equations, including (i) a priori selection of expected model coefficients, and (ii) sampling intensity for background (absence/pseudo-absence) data within a given definition of availability. We show that random sampling of background data violates the underlying mathematics of RSA, leading to incorrect values for necessary M and N and potentially incorrect RSA model outputs. We argue that these equations should be a mandatory component for all future RSA studies. C_LI

ecology↗

Integrating plant stoichiometry and feeding experiments: state-dependent forage choice and its implications on body mass

Intraspecific feeding choices account for a large portion of herbivore foraging in many ecosystems. Plant resource quality is heterogeneously distributed, affected by nutrient availability and growing conditions. Herbivores navigate landscapes, making feeding decisions according to food qualities, but also energetic and nutritional demands. We test three non-exclusive foraging hypotheses using the snowshoe hare (Lepus americanus): 1) herbivores feeding choices and body conditions respond to intraspecific plant quality variation, 2) feeding responses are mitigated when energetic demands are high, and 3) feeding responses are inflated when nutritional demands are high. We measured black spruce (Picea mariana) nitrogen, phosphorus, and terpene compositions, as indicators of quality, within a snowshoe hare trapping grid and found plant growing conditions to explain spruce quality variation (R2 < 0.36). We then offered two qualities of spruce (H1) from the trapping grid to hares in cafeteria-style experiments and measured their feeding and body condition responses (n = 75). We proxied energetic demands (H2) with ambient temperature and coat insulation (% white coat) and nutritional demands (H3) with the spruce quality (nitrogen and phosphorus content) in home ranges. Hares that preferred higher-quality spruce lost less weight during experiments (p = 0.018). The results supported our energetic predictions: hares in colder temperatures and with less-insulative coats (lower % white) consumed more spruce and were less selective towards high-quality spruce. Collectively, we found variation in plant growing conditions within herbivore home ranges substantial enough to affect herbivore body conditions, but any plant-herbivore interactions are also mediated by animal energetic states.

ecology↗

Moving together, foraging apart: effects of the social environment on movement-integrated habitat selection

O_LIMovement links the distribution of habitats with the social environment of animals using those habitats; yet integrating movement, habitat selection, and socioecology remains an opportunity for further study. C_LIO_LIHere, our objective was to disentangle the roles of habitat selection and social association as drivers of collective movement in a gregarious ungulate. To accomplish this objective, we (1) assessed whether socially familiar individuals form discrete social communities and whether social communities have high spatial, but not necessarily temporal, overlap; and (2) we modelled the relationship between collective movement and selection of foraging habitats using socially informed integrated step selection analysis. C_LIO_LIWe used social network analysis to assign individuals to social communities and determine short and long-term social preference among individuals. Using integrated step selection functions (iSSF), we then modelled the effect of social processes, i.e., nearest neighbour distance and social preference, and movement behaviour on patterns of habitat selection. C_LIO_LIBased on assignment of individuals to social communities and home range overlap analyses, individuals assorted into discrete social communities, and these communities had high spatial overlap. By unifying social network analysis with iSSF, we identified movement-dependent social association, where individuals foraged with more familiar individuals, but moved collectively with any between foraging patches. C_LIO_LIOur study demonstrates that social behaviour and space use are inter-related based on spatial overlap of social communities and movement-dependent habitat selection. Movement, habitat selection, and social behaviour are linked in theory. Here, we put these concepts into practice to demonstrate that movement is the glue connecting individual habitat selection to the social environment. C_LI

animal behavior and cognition↗

Does where you live influence what you are made of? Spatial correlates of chemical traits across commonly occurring boreal plants

ContextSpatially explicit drivers of foliar chemical traits link plants to ecosystem processes to reveal landscape functionality. Specifically, foliar elemental, stoichiometric, and phytochemical (ESP) compositions represent key indicator traits. ObjectivesHere, we investigate the spatial drivers of foliar ESP at the species level and across species at the trait level for five commonly occurring boreal forest understory plants. MethodsOn the island of Newfoundland, Canada, we collected foliar material from four chronosequenced forest grids. Using response variables of foliar elemental (C, N, P, percent and quantity), stoichiometric (C:N, C:P, N:P), and phytochemical (terpenoids) composition, we tested multiple competing hypotheses using spatial predictors of land cover (e.g., coniferous, deciduous, mixedwood), productivity (e.g., enhanced vegetation index), biotic (e.g., stand age/height, canopy closure) and abiotic (e.g., elevation, aspect, slope) factors. ResultsWe found evidence to support spatial relationships of foliar ESP for most species (mean R2 = 0.22, max = 0.65). Spatial variation in elemental quantity traits of C, N, P were related to land cover along with biotic and abiotic factors for 2 of 5 focal species. Notably, foliar C, C:P, and sesquiterpene traits between different species were related to abiotic factors. Similarly, foliar terpenoid traits between different species were related to a combination of abiotic and biotic factors (mean R2 = 0.26). ConclusionsSpatial-trait relationships mainly occur at the species level, with some commonalities occurring at the trait level. By linking foliar ESP traits to spatial predictors, we can map plant chemical composition patterns that influence landscape-scale ecosystem processes.

ecology↗

Forage stoichiometry predicts the home range size of a small terrestrial herbivore

Consumers make space use decisions based on resource quality. Most studies that investigate the influence of resource quality on the spatial ecology of consumers use diverse proxies for quality including measures based on habitat classification, forage species diversity and abundance, and nutritional indicators, e.g., protein. Ecological stoichiometry measures resource quality in terms of elemental ratios, e.g., carbon (C):nitrogen (N) ratio, but rarely have these currencies been used to study consumer space use decisions. Yet, elemental ratios provide a uniquely quantitative way to assess resource quality. Consequently, ecological stoichiometry allows for investigation of how consumers respond to spatial heterogeneity in resource quality by changing their space use, e.g. their home range size, and how this may influence ecosystem dynamics and trophic interactions. Here, we test whether the home range size of a keystone boreal herbivore, the snowshoe hare (Lepus americanus), varies with differences in the C:N, C:phosphorus (P), and N:P ratios of two preferred forage species, lowland blueberry (Vaccinium angustifolium) and red maple (Acer rubrum). We consider forage resources with higher C content relative to N and P to be lower quality than resources with lower relative C content. We use a novel approach, combining elemental distribution models with herbivore home range size estimates to test our hypothesis that hare home range size will be smaller in areas with access to high, homogeneous resource quality compared to areas with access to low, heterogeneous resource quality during summer months. Our results support our prediction for lowland blueberry, but not for red maple. Herbivore home range size decreased with increasing blueberry foliage quality, but also with decreasing spatial heterogeneity in blueberry foliage quality, i.e. N or P content. Herbivores in the boreal forest face strong nutritional constraints due to the paucity of N and P. Access to areas of high, homogeneous resource quality is paramount to meeting their dietary requirements with low effort. In turn, this may influence community (e.g., trophic interactions) and ecosystem (e.g., nutrient cycling) processes. Paradoxically, our study shows that taking a reductionist approach of viewing resources through a biochemical lens can lead to holistic insights of consumer spatial ecology.

ecology↗

Patterns and potential drivers of intraspecific variability in the body elemental composition of a terrestrial consumer, the snowshoe hare (Lepus americanus)

O_LIIntraspecific variability in ecological traits is widespread in nature. Recent evidence, mostly from aquatic ecosystems, shows individuals differing at the most fundamental level, that of their chemical composition. Age, sex, or body size may be key drivers of intraspecific variability in the body concentrations of carbon (C), nitrogen (N), and phosphorus (P). However, we still have a rudimentary understanding of the patterns and drivers of intraspecific variability in chemical composition of terrestrial consumers, particularly vertebrates.\nC_LIO_LIHere, we investigate the whole-body chemical composition of snowshoe hare Lepus americanus, providing one of the few studies of patterns of stoichiometric variability and its potential drivers for a terrestrial vertebrate. Based on snowshoe hare ecology, we expected higher P and N concentrations in females, as well as in larger and older individuals.\nC_LIO_LIWe obtained whole-body C, N, and P concentrations and C:N, C:P, N:P ratios from a sample of 50 snowshoe hares. We then used general linear models to test for evidence of a relationship between age, sex, or body size and stoichiometric variability in hares.\nC_LIO_LIWe found considerable variation in the C, N, and P concentrations and elemental ratios within our sample. Contrary to our predictions, we found evidence of N content decreasing with age. As expected, we found evidence of P content increasing with body size. As well, we found no support for a relationship between sex and N or P content, nor for variability in C content and any of our predictor variables.\nC_LIO_LIDespite finding considerable stoichiometric variability in our sample, we found no substantial support for age, sex, or body size to relate to this variation. The weak relationship between body N concentration and age may suggest varying nutritional requirements of individuals at different ages. Conversely, Ps weak relationship to body size appears in line with recent evidence of the potential importance of P in terrestrial systems. Snowshoe hares are a keystone herbivore in the boreal forest of North America. The substantial stoichiometric variability we find in our sample could have important implications for nutrient dynamics in both boreal and adjacent ecosystems.\nC_LI

ecology↗

Conducting social network analysis with animal telemetry data: applications and methods using spatsoc

O_LIWe present spatsoc: an R package for conducting social network analysis with animal telemetry data.\nC_LIO_LIAnimal social network analysis is a method for measuring relationships between individuals to describe social structure. Using animal telemetry data for social network analysis requires functions to generate proximity-based social networks that have flexible temporal and spatial grouping. Data can be complex and relocation frequency can vary so the ability to provide specific temporal and spatial thresholds based on the characteristics of the species and system is required.\nC_LIO_LIspatsoc fills a gap in R packages by providing flexible functions, explicitly for animal telemetry data, to generate gambit-of-the-group data, perform data-stream randomization and generate group by individual matrices.\nC_LIO_LIThe implications of spatsoc are that current users of large animal telemetry or otherwise georeferenced data for movement or spatial analyses will have access to efficient and intuitive functions to generate social networks.\nC_LI

ecology↗

Trends and perspectives on the use of social network analysis in behavioural ecology: a bibliometric approach

The increased popularity and improved accessibility of social network analysis has improved our ability to test hypotheses about the complexity of animal social structure. To gain a deeper understanding of the use and application of social network analysis, we systematically surveyed the literature and extracted information on publication trends from articles using social network analysis. We synthesize trends in social network research over time and highlight variation in the use of different aspects of social network analysis. Our primary finding highlights the increase in use of social network analysis over time and from this finding, we observed an increase in the number of review and methods of social network analysis. We also found that most studies included a relatively small number (median = 15, range = 4-1406) of individuals to generate social networks, while the number and type of social network metrics calculated in a given study varied zero to nine (median = 2, range 0-9). The type of data collection or the software programs used to analyze social network data have changed; SOCPROG and UCINET have been replaced by various R packages over time. Finally, we found strong taxonomic and conservation bias in the species studied using social network analysis. Most species studied using social networks are mammals (111/201, 55%) or birds (47/201, 23%) and the majority tend to be species of least concern (119/201, 59%). We highlight emerging trends in social network research that may be valuable for distinct groups of social network researchers: students new to social network analysis, experienced behavioural ecologists interested in using social network analysis, and advanced social network users interested in trends of social network research. In summary we address the temporal trends in social network publication practices, highlight potential bias in some of the ways we employ social network analysis, and provide recommendations for future research based on our findings.

animal behavior and cognition↗