Small mammal personalities generate context dependence in the seed dispersal mutualism.

Mutualisms are foundational components of ecosystems with the capacity to generate biodiversity through adaptation and coevolution and give rise to essential services such as pollination and seed dispersal. To understand how mutualistic interactions shape communities and ecosystems, we must identify the mechanisms that underlie their functioning. One mechanism that may drive mutualisms to vary in space and time is the unique behavioral types, or personalities, of the individuals involved. Here, our goal was to examine interindividual variation in the seed dispersal mutualism and identify the role that different personalities play. In a field experiment, we observed individual deer mice (Peromyscus maniculatus) with known personality traits predating and dispersing seeds in a natural environment and classified all observed interactions made by individuals as either positive or negative. We then scored mice on a continuum from antagonistic to mutualistic and found that within a population of scatter hoarders, some individuals are more mutualistic than others and that one factor driving this distinction is animal personality. Through this empirical work, we provide a conceptual advancement to the study of mutualism by integrating it with the study of intraspecific behavioral variation. These findings indicate that animal personality is a previously overlooked mechanism generating context dependence in plant­animal interactions and suggest that behavioral diversity may have important consequences for the functioning of mutualisms.

Environmental heterogeneity modifies the link between personality and survival in fluctuating small mammal populations.

Despite numerous studies examining the fitness consequences of animal personalities, predictions concerning the relationship between personality and survival are not consistent with empirical observations. Theory predicts that individuals who are risky (i.e. bold, active and aggressive) should have higher rates of mortality; however, empirical evidence shows high levels of variation in behaviour-survival relationships in wild populations. We suggest that this mismatch between predictions under theory and empirical observations results from environmental contingencies that drive heterogeneity in selection. This uncertainty may constrain any universal directional relationships between personality traits and survival. Specifically, we hypothesize that spatiotemporal fluctuations in perceived risk that arise from variability in refuge abundance and competitor density alter the relationship between personality traits and survival. In a large-scale manipulative experiment, we trapped four small mammal species in five subsequent years across six forest stands treated with different management practices in Maine, United States. Stands all occur within the same experimental forest but contain varying amounts of refuge and small mammal densities fluctuate over time and space. We quantified the effects of habitat structure and competitor density on the relationship between personality traits and survival to assess whether directional relationships differed depending on environmental contingencies. In the two most abundant species, deer mice and southern red-backed voles, risky behaviours (i.e. higher aggression and boldness) predicted apparent monthly survival probability. Mice that were more aggressive (less docile) had higher survival. Voles that were bolder (less timid) had higher survival, but in the risky forest stands only. Additionally, traits associated with stress coping and de-arousal increased survival probability in both species at high small mammal density but decreased survival at low density. In the two less abundant study species, there was no evidence for an effect of personality traits on survival. Our field experiment provides partial support for our hypothesis: that spatiotemporal fluctuations in refuge abundance and competitor density alter the relationship between personality traits and survival. Our findings also suggest that behaviours associated with stress coping and de-arousal may be subject to density-dependent selection and should be further assessed and incorporated into theory.

Land-use change alters associations between personality and microhabitat selection.

Ecologists commonly assess ecological patterns at the population level, focusing on the average response of all individuals within a population, but to predict how populations will respond to land-use change we must understand how changes to habitat differentially affect individuals within a population. For example, forest management is a widespread type of land-use that impacts wildlife through the loss of key habitat features, but individuals within a population may vary in their responses to this loss due to differences in habitat selection among individuals. Specifically, intraspecific variation in habitat selection has been linked to animal personalities (i.e., consistent behavioral differences among conspecifics), but previous research has not examined whether the relationship between personality and habitat selection is influenced by land-use change. To address this knowledge gap, we tested the hypothesis that land-use change alters the association between personality and microhabitat selection in small mammals. Specifically, we investigated two main questions: (1) To what extent are personality type and microhabitat selection correlated among conspecifics? (2) Does land-use change alter individual patterns of microhabitat selection? To answer these questions, we conducted a large-scale field experiment over 4 years, contrasting unmanaged forest (control) with managed forest (two silvicultural treatments) in Maine, USA. We examined the relationships between habitat selection and personality traits in deer mice (Peromyscus maniculatus) and southern red-backed voles (Myodes gapperi). We found that personality traits were correlated with microhabitat selection at multiple spatial scales. Furthermore, land-use change altered these patterns of selection; resulting in either the loss of personality-associated selection or in novel patterns of selection in managed forests. These findings suggest that promoting structural complexity at multiple spatial scales, such as by interspersing stands of mature forest with managed stands, may maintain a variety of intraspecific habitat selection patterns and the associated ecological outcomes.

Environmental heterogeneity and population density affect the functional diversity of personality traits in small mammal populations.

Understanding factors affecting the functional diversity of ecological communities is an important goal for ecologists and conservationists. Previous work has largely been conducted at the community level; however, recent studies have highlighted the critical importance of considering intraspecific functional diversity (i.e. the functional diversity of phenotypic traits among conspecifics). Further, a major limitation of existing literature on this topic is the lack of empirical studies examining functional diversity of behavioural phenotypes-including animal personalities. This is a major shortcoming because personality traits can affect the fitness of individuals, and the composition of personalities in a population can have important ecological consequences. Our study aims to contribute to filling this knowledge gap by investigating factors affecting the functional diversity of personality traits in wild animal populations. Specifically, we predicted that the richness, divergence and evenness associated with personality traits would be impacted by key components of forest structure and would vary between contrasting forest types. To achieve our objective we conducted a fully replicated large-scale field experiment over a 4 year period using small mammal populations as a model system. We found that greater heterogeneity in the cover of shrubs, coarse woody debris and canopy cover was associated with a greater richness, lower divergence and lower evenness in personality traits. Greater population density was associated with greater functional richness and lower functional divergence and evenness of personality traits. To maintain a behaviourally diverse population and its associated functions, managers may promote heterogeneity in vegetation and increased population density, which we found to be the most important determinants driving functional diversity of personality traits.

Land-use change and the ecological consequences of personality in small mammals.

Many plants rely on animals for seed dispersal, but are all individuals equally effective at dispersing seeds? If not, then the loss of certain individual dispersers from populations could have cascade effects on ecosystems. Despite the importance of seed dispersal for forest ecosystems, variation among individual dispersers and whether land-use change interferes with this process remains untested. Through a large-scale field experiment conducted on small mammal seed dispersers, we show that an individual's personality affects its choice of seeds, as well as how distant and where seeds are cached. We also show that anthropogenic habitat modifications shift the distribution of personalities within a population, by increasing the proportion of bold, active, and anxious individuals and in-turn affecting the potential survival and dispersal of seeds. We demonstrate that preserving diverse personality types within a population is critical for maintaining the key ecosystem function of seed dispersal.

Extensive behavioral data contained within existing ecological datasets.

Long-term ecological datasets contain vast behavioral data, enabling the quantification of among-individual behavioral variation at unprecedented spatiotemporal scales. We detail how behaviors can be extracted and describe how such data can be used to test new hypotheses, inform population and community ecology, and address pressing conservation needs.

Umbrella effect of monitoring protocols for mammals in the Northeast US.

Developing cost-effective monitoring protocols is a priority for wildlife conservation agencies worldwide. In particular, developing protocols that cover a wide range of species is highly desirable. Here we applied the 'umbrella species' concept to the context of ecological monitoring; specifically testing the hypothesis that protocols developed for the American marten would contextually allow detecting occupancy trends for 13 other mammalian species (i.e., an umbrella effect). We conducted a large-scale four-year camera trapping survey across a gradient of forest disturbance in Maine, USA. We sampled 197 sites using a total of 591 cameras and collected over 800,000 photographs to generate detection histories for the most common terrestrial species. By combining multi-season occupancy modelling and power analyses, we estimated the required sampling effort to detect 10%, 25% and 50% declines in the fourteen species. By conducting a spatially explicit comparison of sampling effort, we found evidence that monitoring protocols for American marten would provide an umbrella effect for up to 11 other mammal species. The capacity of the umbrella effect varied among species, with fisher, snowshoe hare, red squirrel, and black bear consistently covered under several scenarios. Our results support the application of the umbrella species concept to monitoring (here defined as 'umbrella monitoring species'), providing empirical evidence for its use by management agencies.

Correction: Effects of trap confinement on personality measurements in two terrestrial rodents.

[This corrects the article DOI: 10.1371/journal.pone.0221136.].

Effects of trap confinement on personality measurements in two terrestrial rodents.

In recent years, consistent individual differences in behavior, or personalities, have been a topic of increasing interest as researchers strive to understand and predict the responses of individuals and populations to anthropogenic changes. Behavioral studies in wild populations often require that animals are live trapped before behavioral observation can occur, and this is especially true in studies investigating animal personalities. However, it is unknown whether the amount of time confined to a live trap may regulate the behavior of trapped individuals. Specifically, if the duration of trap confinement directly influences behavior, then by obtaining wild animals through live trapping we may be confounding the very measurements of greatest interest. To investigate whether the duration of trap confinement influences the behavior of trapped individuals, we performed a study on two small mammal species, focusing specifically on personality traits. We positioned high-definition trail cameras facing Longworth small mammal traps in the field to observe capture events and record the time of capture. We then measured personality in captured deer mice (Peromyscus maniculatus) and southern red-backed voles (Myodes gapperi) using three standardized tests, and through linear and generalized linear models we found that the time an animal had spent confined to a trap before testing did not affect 86% of behaviors exhibited. Our results showed two weak behavioral effects of confinement duration on boldness and docility resulting from an interaction between the duration of confinement and whether or not an individual was naïve to trapping. Our results suggest that behavioral measurements of wild, trapped small mammals are not determined by the time spent confined to a trap. However, researchers should use caution and consider whether an animal is naïve to trapping during analysis since habituation to the live trap may play a role in the effects of confinement duration on behavior.