Jana A. Eccard

Integrating movement ecology with biodiversity research - exploring new avenues to address spatiotemporal biodiversity dynamics

Movement of organisms is one of the key mechanisms shaping biodiversity, e.g. the distribution of genes, individuals and species in space and time. Recent technological and conceptual advances have improved our ability to assess the causes and consequences of individual movement, and led to the emergence of the new field of ‘movement ecology’. Here, we outline how movement ecology can contribute to the broad field of biodiversity research, i.e. the study of processes and patterns of life among and across different scales, from genes to ecosystems, and we propose a conceptual framework linking these hitherto largely separated fields of research. Our framework builds on the concept of movement ecology for individuals, and demonstrates its importance for linking individual organismal movement with biodiversity. First, organismal movements can provide ‘mobile links’ between habitats or ecosystems, thereby connecting resources, genes, and processes among otherwise separate locations. Understanding these mobile links and their impact on biodiversity will be facilitated by movement ecology, because mobile links can be created by different modes of movement (i.e., foraging, dispersal, migration) that relate to different spatiotemporal scales and have differential effects on biodiversity. Second, organismal movements can also mediate coexistence in communities, through ‘equalizing’ and ‘stabilizing’ mechanisms. This novel integrated framework provides a conceptual starting point for a better understanding of biodiversity dynamics in light of individual movement and space-use behavior across spatiotemporal scales. By illustrating this framework with examples, we argue that the integration of movement ecology and biodiversity research will also enhance our ability to conserve diversity at the genetic, species, and ecosystem levels.

Is the antipredatory response in behaviour reflected in stress measured in faecal corticosteroids in a small rodent

Predation risk has been shown to alter various behaviours in prey. Risk alters activity, habitat use and foraging, and weight decrease might be a consequence of that. In mammals, studies on physiological measures affected by risk of predation, other than weight, are rare. We studied in two separate laboratory experiments foraging, hoarding behaviour and expression of stress measured non-invasively from the faeces in the bank vole (Clethrionomys glareolus), a common boreal rodent. Voles were exposed to predation risk using odours of the least weasels (Mustela nivalis nivalis). Distilled water served as control. In the first experiment, we found that foraging effort, measured as sunflower seeds taken from seed trays filled with sand, was significantly lower in trays scented with weasel odour. Both immediate consumption of seeds and hoarding were affected negatively by the weasel odour. Females hoarded significantly more than males in autumn. In the second experiment, the negative effect of weasel odour on foraging was consistent over a 3-day experiment, but the strongest effect was observed in the first night. Foraging increased over the time of the experiment, which might reflect either energetic compensation during a longer period of risk, predicted in the predation risk allocation hypothesis, or habituation to the odour-simulated risk. Despite decreased foraging under predation risk, stress measured as corticosteroid metabolite concentration in vole faeces was not affected by the weasel odour treatment. In conclusion, we were able to verify predation-risk-mediated changes in the foraging effort of bank voles but no physiological stress response was measured non-invasively, probably due to great individual variation in secretion of stress hormones.

Interspecific competition in small rodents: from populations to individuals

The role of interspecific competition in shaping animal and plant communities has formed one of the major issues in ecology for decades. Small mammals, mainly rodents, have been among the model systems used for research on interspecific competition. Most studies within small mammal systems in the past have examined effects of competition on population attributes such as on population size, habitat use, or population dynamics. Population-level responses are the cumulative effects of individual responses, however, the influence of competition on individual life-history traits has rarely been studied. Research on life-histories may bridge gaps between population biology and effects of competition on individual behaviour. In this paper, we review recent research approaches to interspecific competition in rodents based on census data and species assemblages, that use regression analysis, time series analysis, removal and exclusion experiments, and showcase our own experimental research on the effects of interspecific competition on individual life-history traits in boreal voles.

Long-term fitness benefits of polyandry in a small mammal, the bank vole Clethrionomys glareolus

Polyandry, i.e. mating with multiple males within one reproductive event, is a common female mating strategy but its adaptive function is often unclear. We tested whether polyandrous females gain genetic benefits by comparing fitness traits of monandrous (mated twice with a single male) and polyandrous (mated twice with two different males) female bank voles Clethrionomys glareolus. We raised the offspring in the laboratory until adulthood and measured their body size, before releasing them to outdoor enclosures to overwinter. At the onset of the breeding season in the following spring, we found that offspring of polyandrous females performed significantly better at reproduction than those of monandrous females. This was mainly due to sons of polyandrous females producing significantly more offspring than those of monandrous females. No significant differences were found for offspring body mass or winter survival between the two treatments. Our results appear to provide evidence that bank vole females gain long-term benefits from polyandry.

Reproductive success of male bank voles (Clethrionomys glareolus): the effect of operational sex ratio and body size

The operational sex ratio (OSR) may influence the intensity of competition for mates and mate choice and is therefore thought to be a major factor predicting the intensity and direction of sexual selection. We studied the opportunity for sexual selection, i.e., the variance in male reproductive success and the direction and intensity of sexual selection on male body mass in bank vole (Clethrionomys glareolus) enclosure populations with experimentally manipulated sex ratios. The opportunity for sexual selection was high among male-biased OSRs and decreased towards female-biased OSRs. Paradoxically, selection for large male body mass was strongest in female-biased OSRs and also considerable at intermediate OSRs, whereas at male-biased OSRs, only a weak relationship between male size and reproductive success was found. Litters in male-biased OSRs were more likely to be sired by multiple males than litters in female-biased OSRs. Our results suggest that the intensity and direction of sexual selection in males differs among different OSRs. Although the direction of sexual selection on male body mass was opposite than predicted, large body mass can be favored by sexual selection. Naturally varying OSRs may therefore contribute to maintain variation in male sexually selected traits.

Do female bank voles (Clethrionomys glareolus) mate multiply to improve on previous mates

Contrary to classical sexual selection theories, females of many taxa mate with multiple males during one reproductive cycle. In this study, we conducted an experiment on the “trade-up hypothesis”, which proposes that females remate if a subsequently encountered male is potentially superior to previous mates to maximize the genetic quality of their offspring. We presented bank vole females (Clethrionomys glareolus) sequentially with two males of known dominance rank in different orders, i.e., either first subordinate and second dominant, first dominant and second subordinate, or two males that were equal in dominance (high ranking) and observed their mating behavior. We found that 92% of the females mated multiply and did not base their remating decision on male social status. Therefore, polyandry cannot be explained by the “trade-up hypothesis” based on dominance rank in this species. However, we found that dominant males sired significantly more offspring than subordinate males. This varied according to mating order: dominant males sired more offspring when they were second than when they were first. Moreover, litter sizes were significantly smaller when the dominant male was first (smallest relative success of dominant males) compared to litter sizes when mating order was reversed or both males equal in status. Our results suggest that even though multimale mating includes males that are of poorer quality and thus potentially decreases the fitness of offspring, most of a female’s offspring are sired by dominant males. Whether this is due to cryptic female choice, sperm competition, or a combination of both, remains to be tested.

WEASELS’ (MUSTELA NIVALIS NIVALIS) PREFERENCE FOR OLFACTORY CUES OF THE VOLE (CLETHRIONOMYS GLAREOLUS)

Many studies on life history strategies of small mammals under predation risk are based on assumptions that mammalian predators use scent marking from prey in searching and hunting. This is especially true for small mustelids hunting in the tunnels and cavities of their prey. It is assumed that weasels use the estrous signs of female voles as hunting cues, which exposes such females to a more pronounced risk of predation. We studied the preferences of 57 least weasels (Mustela nivalis nivalis) toward odor cues from four different reproductive categories of the bank vole (Clethrionomys glareolus). In the first experiment, weasels selected clearly for vole odors over clean bedding in a Y-maze arena. The second experiment demonstrated that there was no difference in weasel preference between estrous and pregnant or lactating females. Thirdly, there was no preference for odors from dominant vs. subordinate males. Finally, we offered weasels a dyadic choice between olfactory signs of six possible combinations ...

Consistency in boldness, activity and exploration at different stages of life

BackgroundAnimals show consistent individual behavioural patterns over time and over situations. This phenomenon has been referred to as animal personality or behavioural syndromes. Little is known about consistency of animal personalities over entire life times. We investigated the repeatability of behaviour in common voles (Microtus arvalis) at different life stages, with different time intervals, and in different situations. Animals were tested using four behavioural tests in three experimental groups: 1. before and after maturation over three months, 2. twice as adults during one week, and 3. twice as adult animals over three months, which resembles a substantial part of their entire adult life span of several months.ResultsDifferent behaviours were correlated within and between tests and a cluster analysis showed three possible behavioural syndrome-axes, which we name boldness, exploration and activity. Activity and exploration behaviour in all tests was highly repeatable in adult animals tested over one week. In animals tested over maturation, exploration behaviour was consistent whereas activity was not. Voles that were tested as adults with a three-month interval showed the opposite pattern with stable activity but unstable exploration behaviour.ConclusionsThe consistency in behaviour over time suggests that common voles do express stable personality over short time. Over longer periods however, behaviour is more flexible and depending on life stage (i.e. tested before/after maturation or as adults) of the tested individual. Level of boldness or activity does not differ between tested groups and maintenance of variation in behavioural traits can therefore not be explained by expected future assets as reported in other studies.

Why do female bank voles, Clethrionomys glareolus, mate multiply?

Females of many species actively engage in multiple mating, with either a single male or several males, but the adaptive function of this behaviour is often unclear. We conducted a laboratory experiment on a small mammal species, the bank vole, testing the possible benefits of multiple mating on a females short-term reproductive success (pregnancy rate, litter size and early postnatal survival). Such benefits may affect a females fitness either directly or indirectly (genetic benefit). We assigned females to three treatments: a single mating treatment in which females mated once with a single male and two multiple mating treatments in which females mated either twice with a single male or twice with two different males. We found a significant reduction in pregnancy rate of females that mated only once compared to females that mated twice. This direct benefit is most likely explained by an increased stimulus gained from multiple mating. However, we found no difference in reproductive success of females mated twice with the same male or once with each of two males. Our study supports the importance of direct benefits of multiple mating. Although our study cannot rule out effects of genetic benefits on future survival and reproductive success of offspring, we found no fitness benefits of polyandry for the traits studied here.

SPACE USE, CIRCADIAN ACTIVITY PATTERN, AND MATING SYSTEM OF THE NOCTURNAL TREE RAT THALLOMYS NIGRICAUDA

Abstract We present results of a radiotracking study of the black-tailed tree rat Thallomys nigricauda, based on 3 males and 4 females in the breeding season and 2 males and 5 females in the nonbreeding season. The study was conducted in the southern Kalahari thornveld, South Africa, a savanna landscape of acacia trees and patchy acacia bush. Nocturnal activity patterns, vegetation use, space use, and interactions were followed for 8–17 days. During the breeding season, males were active with 2 synchronous activity peaks shortly after sunset and late at night. Home ranges included the nests of 8–10 females, 6 different daytime resting places, and 3–5 areas of high activity. Males were mobile within home ranges of 5–10 ha (90% minimal convex polygons) that overlapped with other males and covered home ranges of several females. Activity of females was high shortly after sunset and before sunrise with 6 h of low activity between. Home ranges of nursing females were 0.001–0.03 ha and included 2 small areas of high activity (nest and foraging site) where no other adult females were observed. Animals preferred small acacia shrubs (Acacia mellifera, A. luederitzi) for foraging. During the nonbreeding season, activity of both sexes was low, with only 1 peak shortly after sunset. Although small acacia shrubs did not have foliage, they were preferred by both sexes for foraging, and rats were observed feeding on buds. Home-range size of males was 1–2 ha and of females was 0.1–0.3 ha. Contrary to earlier reports, we found tree rats living solitarily or with young offspring. We suggest that the species has a promiscuous mating system.