Predation on two vole species by a shared predator: antipredatory response and prey preference
|
|
- Angela Merry Lawrence
- 6 years ago
- Views:
Transcription
1 Popul Ecol (2008) 50: DOI /s ORIGINAL ARTICLE Predation on two vole species by a shared predator: antipredatory response and prey preference Janne Sundell Æ Lenka Trebatická Æ Tarja Oksanen Æ Otso Ovaskainen Æ Marko Haapakoski Æ Hannu Ylönen Received: 24 July 2007 / Accepted: 31 March 2008 / Published online: 8 May 2008 Ó The Society of Population Ecology and Springer 2008 Abstract In prey communities with shared predators, variation in prey vulnerability is a key factor in shaping community dynamics. Conversely, the hunting efficiency of a predator depends on the prey community structure, preferences of the predator and antipredatory behavioural traits of the prey. We studied experimentally, under seminatural field conditions, the preferences of a predator and the antipredatory responses of prey in a system consisting of two Myodes species of voles, the grey-sided vole (M. rufocanus Sund.) and the bank vole (M. glareolus Schreb.), and their specialist predator, the least weasel (Mustela nivalis nivalis L.). To quantify the preference of the weasels, we developed a new modelling framework that can be used for unbalanced data. The two vole species were hypothesised to have different habitat-dependent vulnerabilities. We created two habitats, open and forest, to provide different escape possibilities for the voles. We found a weak general preference of the weasels for the grey-sided voles over the bank voles, and a somewhat stronger preference J. Sundell (&) O. Ovaskainen Metapopulation Research Group, Department of Biological and Environmental Sciences, University of Helsinki, P.O. Box 65, Helsinki, Finland janne.sundell@helsinki.fi L. Trebatická M. Haapakoski H. Ylönen Konnevesi Research Station, Department of Biological and Environmental Science, University of Jyväskylä, P.O. Box 35, Jyväskylä, Finland T. Oksanen Department of Ecology and Environmental Sciences, Umeå University, Umeå, Sweden specifically in open habitats. The weasels clearly preferred male grey-sided voles over females, whereas in bank voles, there was no difference. The activity of voles changed over time, so that voles increased their movements immediately after weasel introduction, but later adjusted their movements to times of lowered predation risk. Females that were more active had an elevated mortality risk, whereas in the case of males, the result was the opposite. We conclude that, in vulnerability to predation, the species- or habitatspecific characteristics of these prey species are playing a minor role compared to sex-specific characteristics. Keywords Antipredatory behaviour Apparent competition Clethrionomys Prey preference Introduction Predators have the potential to shape the dynamics of their prey. Differential predation on different prey types or species can also modify the prey community structure. One way in which this happens is predator-mediated competition, in which the increase of one prey type has a negative effect on the other prey type indirectly via a shared predator (i.e., apparent competition sensu Holt 1977). In a predator-mediated apparent competition, predators can react numerically or behaviourally to density changes of one of the coexisting prey species, which increases predation also on the other alternative prey species. This is observable as negative effects between prey species, even without actual competition (Holt 1977; Holt and Kotler 1987). The strength of a predator s power to modify the prey community depends on its specialisation (generalists vs. specialists; e.g., Hanski et al. 1991), prey preferences and the vulnerability of different prey types (e.g., Abrams
2 258 Popul Ecol (2008) 50: et al. 1998), together with competitive interactions (Sih et al. 1985; Chase et al. 2002) and environmental factors (e.g., Menge and Sutherland 1976, 1987). According to Holt s (1977) analysis, the vulnerability of a given prey species depends on the ratio of its reproductive capacity (r) to the predator s searching efficiency (a). In principle, a prey can increase its r/a ratio in two ways; first, by being elusive and, thus, avoiding predation, which reduces the value of a, or by having high reproductive capacity, i.e., a high value of r. If predation risk is unpredictable and short-termed, the latter approach to a high-risk situation is unlikely to be optimal (Oksanen and Lundberg 1995), and the vulnerability of a given species can largely be assessed on the basis of its a value. Given the importance of this parameter for survival under predation risk, it is reasonable to ask why evolution has not resulted in the minimisation of a for all prey species. However, elusiveness has its costs. For example, it is impossible for a herbivore to be both extremely elusive and have the capacity to handle low-quality forage, as there is a trade-off between these two characteristics. Because of this apparent trade-off, herbivore species living in unproductive environments tend to have larger digestive tracks than their congeners living in environments with high-quality food (Oksanen 1992). The grey-sided vole, Myodes rufocanus Sund., and the bank vole, M. glareolus Schreb. (genus name Myodes, was previously Clethrionomys; nomenclature follows Wilson and Reeder 2005), form a suitable species pair for testing the above-mentioned conjecture on differential vulnerability, as they share the same ancestry, but the former species is adapted to less productive habitats than the latter. The grey-sided vole lives in a wide variety of habitats, ranging from forests and mire edges to open tundra (Christensen and Hörnfeldt 2006). The bank vole is also not restricted to any particular habitat type, and can be found commonly in forests and bushy edge habitats. Especially during the peak years of abundance, the two species can be found in the same habitats, and they are often the two most common species in southern Lapland (Henttonen and Hansson 1984). The grey-sided vole is slightly larger, relatively slow, eats more green vegetation and breeds in higher densities than in the other Myodes voles (Viitala 1977; Henttonen 2000). Thus, ecologically, the grey-sided vole resembles more species of another common genus of voles, Microtus voles, than other Myodes species (Hanski and Henttonen 1996). The Microtus voles are mainly folivorous, live in dense matrilinear aggregations (Pusenius et al. 1998) and are heavier and probably also slower in movements than Myodes. The bank vole, as a typical representative of Myodes, is more agile, partly arboreal and does not attain such high densities as Microtus voles. Because of its lower densities and better escape behaviour, weasel predation should not be targeted towards bank vole and it is considered as the alternative prey (Henttonen 1987; Hanski and Henttonen 1996). Although both Myodes and Microtus are primarily preyed upon by small mustelids, and especially by the least weasel (Mustela nivalis nivalis L.), Microtus voles are assumed to be the pivotal prey for breeding weasels and the key prey species in small mustelid vole interactions in cyclic vole populations of Fennoscandia (e.g., Korpimäki et al. 1991). Our purpose was to investigate the underlying factors behind potentially different vulnerabilities of the two prey species to weasel predation. Based on the characteristics of these different types of voles described above, and the characteristics of the hunting behaviour of the least weasel (Jedrzejewski and Jedrzejewska 1990), we hypothesised that the Microtus-like grey-sided vole would be more prone to predation by the least weasel than the typical Myodes, the bank vole. We expected that the less agile grey-sided voles would suffer from heavier predation by the mostly ground-dwelling least weasel and that the partly arboreal bank voles would survive better, especially in forested habitats, because of the escape possibilities to trees. Further, we aimed to investigate how behavioural traits such as space use and activity affect the vulnerability of the prey. According to general theory, increasing mobility should lead to a higher encounter rate with the predator and, consequently, to higher mortality (Lima and Dill 1990), but on the other hand, also, reduced movements with the accumulation of odorous waste products may expose prey to olfactorily hunting predators (Banks et al. 2000). Materials and methods Study design The study was conducted in eight 0.25-ha ( m) enclosures in the vicinity of the Konnevesi Research Station of the University of Jyväskylä, Central Finland ( N, E) in September and October The walls of the enclosures were made of metal sheets reaching ca. 0.5 m below and 1.0 m above the ground. The enclosures were constructed on an old field in The main habitat is grassland, but bushes of Salix spp. cover about 1 10% of the enclosure areas. The terrain is generally flat with small dry ditches and some piles of stones. All of the enclosures had a few (ca. 10 per enclosure) saplings (2 6 m high) of birch (Betula spp.) and spruce (Picea abies). To create two different habitat types, we added young (3 5-m high) spruces by sticking them into the ground in four of the enclosures (to be called forest), while the remaining four were left as such (to be called
3 Popul Ecol (2008) 50: open). Spruces were erected within half of the area (0.125 ha) in each of the four enclosures, so that the total number of trees (added spruces and trees naturally growing in the enclosures) varied from 39 to 57 in the forest treatment. The trees in the forested parts of the enclosures were evenly distributed, with the distance between trees being never more than 7 m. The few trees naturally growing in the open enclosures and in the open half of the forested enclosures were treated by cutting off their lower branches. This procedure was evidently successful, as voles were never seen in these manipulated trees during the study. Small Salix bushes present in the open treatment were probably not suitable for climbing, as we did not observe in this study, nor have observed in our previous studies (e.g., Sundell et al. 2003), voles climbing on the thin and slippery branches mainly growing upwards. The aim of the habitat manipulation was to study the habitat-dependent vulnerability of the two vole species in the forest habitat providing arboreal escape possibilities. Both species have been considered to be able to climb on trees, but the observations of grey-sided voles in trees are largely anecdotal (e.g., Siivonen and Sulkava 1994). Bank voles are often seen collecting lichens for food from trees and are known to escape to trees when exposed to small mustelid predators (Jedrzejewski and Jedrzejewska 1990). The experiment was carried out in three subsequent runs, in which four enclosures were used simultaneously. Experimental animals For each enclosure in each replicate, eight adult mature voles ([19 g), four of each species, with sex ratio 1:1, were radio-collared (TW-4, Biotrack Ltd., Wareham, UK; ca. 1.5 g), marked subcutaneously with passive-induced transponders (Trovan, EID Aalten BV, Aalten, Holland) and released into the middle of a randomly assigned enclosure one day after marking and collaring. Experimental bank voles were captured near the Konnevesi Research Station or belonged to the first generation born in the laboratory or in the outdoor enclosures at the same place. All of the used bank voles had experienced natural conditions in enclosures as a part of an earlier experiment. Grey-sided voles were captured from Finnish, Swedish and Norwegian Lapland. They were brought to the Konnevesi Research Station before the experiment and were housed individually at the laboratory in standard mouse cages and fed with mouse pellets prior to the experiment. Even though the grey-sided vole is a slightly larger species than the bank vole (weight ranges from 20 to 50 g and from 7 to 40 g for grey-sided voles and bank voles, respectively, according to Jensen 1994), we wanted to control for the weight effect and minimise the proportional weight of the collar in this experiment by reducing the natural size difference between species. The mean weight was 24.9 g for bank voles (±2.4 g SD, range g; for females 23.3 ± 1.6 g and for males 26.5 ± 1.9 g), and 25.9 g for grey-sided voles (±3.9 SD, g; for females 25.0 ± 4.2 g and for males 26.9 ± 3.4 g). After 3 4 days habituation period in the enclosures, we started to monitor the space use of voles by locating them every second hour for a 24-h period. The possible use of trees was also recorded. Twenty trees were equipped with live traps (Ugglan Special, Grahnab AB, Hillerstorp, Sweden) to observe the use of trees between radio-trackings. After the habituation period of the voles, we released one least weasel per enclosure. The least weasels were wild-born or belonged to the first generation born in captivity. Altogether, 12 weasels, eight males and four females, were used in the experiment. The mean weights were 57.4 g (±4.4 SD, range g) for males and 43.8 g (±1.2 SD, g) for females. The radiocollared weasels (TW-4, Biotrack Ltd., Wareham, UK, ca. 2.2 g) were released in a randomly assigned enclosure s randomly assigned corner. Weasels were radio-located 3 4 times per 2 h. The weasels were let to hunt voles freely for 72 h or until they had captured four voles, after which, the experiment was terminated. Space use and activity To measure space use, we used the 100% minimum convex polygon method (MCP; Kenward 1987). We call in the sequel the area within the MCP as the home range, although in this kind of short-term enclosure experiment, the MCP reflects general space use rather than the size of an actual home range (Sundell et al. 2000). Home ranges were calculated with the program Ranges6 (Anatrack Ltd., Wareham, UK). We determined the activity of the experimental animals by measuring the mean distance moved between successive radiolocations (=interfix distance). We used data obtained during the entire experimental period or data collected within 24 h before and after the weasels were released, the latter for studying the effect of the predator on home ranges and activity patterns. The reason for restricting to a 24-h period after weasel release is a tendency for increasing home range size with increasing number of observations, especially when the number of observations is relatively small. The home range measures were log-transformed and proportions of habitat within a home range were arcsin-transformed to fulfil the assumptions of data for analysis of variance. Analyses were conducted using the MIXED procedure in the SAS Ò statistical software package, version 9.1. To control for variation in environmental conditions, the enclosures were nested within habitat types. Repeated measures analysis of variance was used when observations before and after
4 260 Popul Ecol (2008) 50: weasel release were compared. Linear regression analysis was used to analyse the dependence of vole activity on weasel activity (SPSS Ò statistical software package, version ). Prior to the analysis of activity changes (r t = x t - x t-1 ), the data was log(x + 1) transformed for linearising the relationship. Cause and timing of death About half of the voles carried radio-collars with temperature sensors. Therefore, it was possible to determine the time of death with an accuracy of a few minutes, as the signal pulse gets faster with decreasing temperature. In the case of voles carrying ordinary radio-collars, we estimated the time of death using the following procedure. If the vole was inactive during three consecutive checks or if a weasel was observed near to its location, we tried to locate the body of the vole. The cause of death was deduced by searching for weasel bite marks from the vole and collar. Also, the location of the dead vole could be used to determine its cause of death, as voles killed by a weasel were usually hidden under the ground or amongst vegetation. All voles killed by weasels were left in the place where they were found. The timings of kills were used to investigate the killing order and preference of the weasels. For the sake of simplicity, here, we use the words preference and prefer, although the probability of an individual vole to fall as a prey for the weasels is likely to be the result of both the vulnerability of the vole and the weasel s active preference. The effect of different factors on the fate of voles (killed by weasel or survived) was examined with a generalised linear model (GLM) using the GENMOD procedure of the SAS Ò statistical software package, version 9.1. To examine the effect of the vole s activity on its vulnerability, we included as an explanatory variable the activity of the vole during the 24 h before weasel release. Since many of the voles died soon after the weasel release, the vole activity during a period of 24 h before the weasel release, in which all of the voles have approximately the same amount of fixes, was used as an explanatory variable. In the analysis, the enclosures were nested within habitat types and a logit link function with binary error distribution was assumed. Weasels hunting preference model The order in which the weasels killed voles was used to examine whether the weasels showed a preference to either prey species or to either sex within a species. As we did not replace the killed prey with new ones, the relative numbers of different prey types available for the weasels did not stay in balance during the experiment. For this reason, the use of preference indices developed earlier (Krebs 1999) would be problematic, and we develop here a new modelling framework that can be used for unbalanced data. We describe the modelling approach for the case of two prey types (e.g., species or sex, denoted by A and B), but it applies also for cases in which there are more types of prey. We assume that the predator moves around searching for prey. If the predator encounters an individual of type A, we assume that it has the probability q A of detecting and catching it, whereas the corresponding probability for type B is denoted by q B. Let us assume that there are currently n A individuals of type A and n B individuals of type B, and denote by x A = n A /(n A + n B ) the relative abundance of type A. Then, the probability p A that the predator will catch next an individual of type A follows the equation: p A ¼ x A ðq A þ ð1 q A Þp A Þþð1 x A Þð1 q B Þp A ð1þ The first term on the right-hand side represents the probability x A that the predator first encounters an individual of type A and either catches it right away (with probability q A ) or does not catch it on that occasion (with probability 1 - q A ), but still catches an individual of type A next (p A ). The second term represents the possibility that the predator encounters an individual of type B next (1 - x A ), but does not catch it on that occasion (1 - q B ), and catches an individual of type A next (p A ). Solving p A from Eq. 1 gives: x A z A p A ¼ ð2þ 1 x A þ x A z A where z A = q A /q B measures the relative preference for prey A. We transform z A into a logarithmic scale by defining the preference that the predator shows for type A over type B as w A = log 10 z A, so that w A = 0 corresponds to no preference, w A [ 0 to preference for type A and w A \ 0 to preference for type B (note that w A + w B = 0). Figure 1 illustrates how the probability p A depends both on the preference w A and on the relative numbers of types A and B. We used a Bayesian approach to relate the model given by Eq. 1 to the data assuming an uninformative (flat) prior for w A [ [-2, 2] (which covers already biologically unrealistically strong preferences) and zero prior probability for w [ 2. Let us denote by y ij the type of prey captured as the jth prey in the replicate i, so that y ij = 1 if the prey is of type A and y ij = 0 if the prey is of type B. The likelihood of the data is then given by: Pr ðyjp A Þ ¼ Yn Y k i i¼1 j¼1 p ij A yij A þ 1 pij A 1 y ij A ð3þ where n is the number of replicates, k i is the total number ij of prey caught in replicate i and p A is the probability of catching next a prey of type A (Eq. 1) in the situation where j - 1 prey were already caught in replicate i. To
5 Popul Ecol (2008) 50: Statistical power Pr[Pr(w > 0) > 0.95] Catching probability (p A ) w = 0.1 w = 0.3 w = Number of replicates w = 1.0 w = 0.5 w = 0 w = -0.5 w = Relative abundance of prey type A (x A ) Fig. 1 Upper panel The statistical power of the estimation method, measured as the probability that a data set contains high statistical support (posterior probability greater than 0.95) for the existence of a positive preference (w [ 0) at different levels of true values of the preference parameter w. Lower panel The model-based probability (p A ) that the predator will next catch an individual of type A (Eq. 1) with different preference levels (w) construct the posterior distribution for w A, we used a discrete grid approximation with grid size 1/50. We first tested the performance of the estimation scheme by performing a power analysis, in which we generated 100 replicate data sets for each of the values of n = 5, 10 and 15, with each experiment initially containing four individuals of both types A and B. We assumed that the preference of the predator was either mild (w A = 0.1), intermediate (w A = 0.3) or strong (w A = 0.5), and we let it catch four prey with the probabilities given by Eq. 1. We estimated the posterior distribution for the parameter w for each of the created data sets and calculated the posterior probability Pr(w A [ 0), by which the data suggests preference for prey type A. We then calculated the fraction of the 100 replicates for which Pr(w A [ 0) [ 0.95, in order to measure how often the results from a given experiment would give a strong signal for the preference for species A. We estimated the posterior distribution of the preference parameter w A for the real data set in two ways. First, we let the two species represent the types A and B, and estimated w A separately for the forest enclosures and for the open enclosures to see if the predator prefers either of the species and if the preference depends on the habitat type. Second, we examined whether there was a preference for either sex by treating the data for each species separately, but pooling the data for the two habitat types. In the real data set, there was an additional complication that some of the prey were recorded dead also for reasons other than predation by the weasels. We excluded such prey from the likelihood (Eq. 3), but accounted for the change that they caused to the relative numbers of the two prey types. One of the replicates (open habitat) was omitted from the analysis of preference, as, in this case, we were not able to determine reliably the killing order of the voles and as one of the male grey-sided voles was missing. Results Preference of weasels In both open and forest enclosures, the median estimates for w suggest that the weasels prefer the grey-sided voles over bank voles (Table 1 for the raw data and Table 2 for the results of analysis), the posterior probability for the pooled data being Pr(w [ 0) = The estimated preference was stronger in open areas (w = 0.30) than in forests (w = 0.14). Comparing the two sexes, the weasels clearly prefer male grey-sided voles over female grey-sided voles (w =-0.39; Pr(w \ 0) = 0.97), whereas in the bank voles, there is no difference between the sexes (Table 2). As expected, the results of the power analysis (Fig. 1) showed that the possibility of obtaining a strong support for the existence of a preference increases with increasing number of replicates and with increasing strength of the preference. Assuming an intermediate strength for the preference (w A = 0.30), which was suggested by the model comparing the two species in the open enclosures, the probability of obtaining a result with Pr(w [ 0) [ 0.95 Table 1 Number of voles killed by weasels in the two enclosure types (forest or open) Grey-sided vole Bank vole Total Female Male Female Male Forest Open 3 (5) 6 3 (4) 3 (4) 15 (19) Total 6 (8) 15 8 (9) 8 (9) 37 (41) The numbers in parentheses include also individuals for which the killing order was not possible to determine and which were, therefore, left out from the analysis of preference (see Materials and methods section). The total number of voles in the experiment was 96 (=12 9 8)
6 262 Popul Ecol (2008) 50: Table 2 Preference (w) of weasels for the grey-sided vole against the bank vole in different enclosure types (forest or open) and for females against males in different species 400 Forest w (95% HPDI) Pr(w [ 0) Preference (w) of weasels for the grey-sided vole against the bank vole Habitat type Forest 0.14 ( ) 0.78 Open 0.30 ( ) 0.90 Pooled data 0.20 ( ) 0.92 Preference (w) of weasels for females against males Prey species Grey-sided vole ( ) 0.03 Bank vole 0.02 ( ) 0.53 The estimates shown for w are the median of the posterior distribution and the 95% highest posterior density interval (HPDI) is around half for the size of our data set (5 replicates in open areas). Performing 15 replicates would increase the probability to only three quarters, which illustrates that it is laborious to demonstrate the existence of a relatively mild preference. If the preference level would be stronger (w A = 0.5), then a set of five replicates would already produce a result with Pr(w [ 0) [ 0.95 with an 80% chance. As the preference of the weasels might depend on the size of the voles, we compared the weight of the voles that were killed by the weasels and those that survived, but no statistically significant difference was found (t-test: t = 1.60, df = 85, P = 0.114). Effect of weasels on space use and activity In the analysis of variance including all of the observations, the sizes of home ranges did not differ between species (F 1,80 = 0.01, P = 0.904), sexes (F 1,80 = 0.62, P = 0.434) or habitats (F 1,86 = 0.08, P = 0.777). All interactions were statistically non-significant and were excluded from the final model. Furthermore, when comparing home range sizes one day before and one day after the release of the weasels, no statistically significant differences were found (Fig. 2; Table 3). However, there was a tendency for larger home ranges in grey-sided voles (time 9 species, F 1,81 = 2.57, P = 0.113) after weasel release, especially in the forested enclosures. No statistically significant effects were found in the proportion of forest within the home ranges in forest enclosures (Table 3). In voles activity, measured as distances moved between consecutive fixes, no differences were found between species (ANOVA, F 1,74 = 0.88, P = 0.351), sexes (F 1,74 = 2.55, P = 0.114) or habitats (F 1,74 = 2.57, P = 0.114) when data from the entire experimental period were Home range MCP (m 2 ) Open Grey-sided vole Bank vole Species Before After Fig. 2 Home range sizes of voles measured as minimum convex polygons (MCP) one day before weasel release and one day after the weasel release. Voles were located every second hour during a 24-h period. The upper panel shows the mean (+SE) home ranges in forest enclosures and the lower panel in open enclosures included. All interactions were statistically non-significant and were excluded from the final model. When the mean interfix distance moved during one day before and after the weasel release was examined, statistically significant differences were found. Generally, voles moved more after the weasel release (time, F 1,84 = 4.52, P = 0.036), but there was a difference between species (time 9 species, F 1,84 = 6.66, P = 0.012; Table 3) so that grey-sided voles moved longer distances when weasels were present (before 6.8 ± 0.7 m and during weasel presence 9.3 ± 0.8 m), while bank voles tended to move less (before 7.4 ± 0.7 m and during 7.1 ± 0.6 m). The fate of the voles (survived or killed by the weasels) was explained by their sex (GLM, v 2 = 7.07, df = 1, P = 0.008) and by their activity (sex 9 activity, v 2 = 5.48, df = 1, P = 0.019; Table 4); generally, more males were killed, but in females, more active voles were killed by weasels, while in males, more active ones survived (Fig. 3). As females moving long distances suffered higher mortality than females moving short distances, and as the pattern was the opposite in males, pooling the data
7 Popul Ecol (2008) 50: Table 3 Repeated measures ANOVA table on log-transformed home range sizes, arcsin-transformed proportions of forest in forest enclosures and interfix distances one day before and after the weasel release (time) Dependent variable Source of variation ndf ddf F P Home range Time (T) Sex Species (S) Habitat (H) Enclosure (habitat) T 9 S Proportion of forest in forest enclosures Time (T) Sex Species (S) Enclosure Interfix distance Time (T) Sex Species (S) Habitat (H) Enclosure (habitat) T 9 S Other factors are species (grey-sided vole or bank vole), sex, habitat (forest or open enclosure) and enclosure. In two of the models, enclosures were nested within habitat types to control the variation among enclosures (not in proportions of forest in forest enclosure). Non-significant interactions were not included in the final model, except in the model of home range, in which the interaction term closest to the 0.05 level of significance was included Table 4 Results of the generalised linear model (GLM) for the effects of activity (interfix distance before weasel release), sex, species (grey-sided vole or bank vole) and habitat (forest or open habitat enclosure) on the fates of the voles (survived or killed by the weasels) Source of variation df v 2 P Activity (A) Sex Species (S) Habitat (H) Enclosure (habitat) A 9 sex In the model, the enclosures were nested within habitat types to control variation among enclosures. Only the significant interactions are included in the model led to a non-linear relationship between the mortality and activity. The highest proportions of voles killed were found in the highest and the lowest distance categories, although voles that moved intermediate distances suffered substantial mortality by the weasels (Fig. 4). Voles activity increased considerably soon after the release of the weasels, but, later on, the voles tended to concentrate their movements to times of low weasel activity (linear regression R 2 = 0.166, F 1,35 = 6.76, P = 0.014; Fig. 5). Observations of voles in trees were rare. Before the weasel release, only one bank vole female was found in a tree twice. The same individual was observed in a tree five times after the release of the weasels. After the weasel release, altogether, three (3/24) grey-sided voles and five (5/24) bank voles were found in trees, with most of these on several occasions. Discussion Preference of weasels Largely based on different strategies and dissimilar r/a ratios (Holt 1977), it has been suggested that the grey-sided vole, as a representative of Microtus-type voles, would be more vulnerable to weasel predation than a typical Myodes vole the bank vole (Hanski and Henttonen 1996; Henttonen 2000). In line with this expectation, we found a slight preference of weasels for grey-sided voles over bank voles. We assumed that, in forested habitat, the bank vole, a good climber, would have gained from the availability of trees more than the grey-sided vole. In these species, the arboreal escape tactic has been observed previously only in bank voles (Jedrzejewska and Jedrzejewski 1990; Jedrzejewski et al. 1993; Sundell and Ylönen 2004). However, contrary to expectations, the preference for the grey-sided vole was stronger in open habitats than in forests. Our observations on climbing suggest that grey-sided voles also use arboreal escape, which may explain the unexpected result of only weak preference for grey-sided voles and a smaller difference in forested enclosures.
8 264 Popul Ecol (2008) 50: Interfix distance (m) Female Grey-sided vole 7 11 Weasel Alive Male Male Female A factor not included in the present experiment is variation in body size. As we decided to control for size, partly to minimise the effect of the radio-collar s proportional weight (Kenward 1987), the natural weight difference between the species was reduced. In this experiment, within the size range variation present in the experimental animals, we did not find a statistically significant difference in weights between the killed voles and those that survived. Similarly, in a previous experiment (Sundell et al. 2003), the weight difference between experimental bank voles and field voles (Microtus agrestis L.) did not turn out to play any role, although it should be noted that, in that experiment, there was no possibility for arboreal escape. To obtain more conclusive evidence on the potential role of Bank vole Fig. 3 Mean activity of voles (±SE) measured as interfix distance (distance moved between consecutive locations). Voles were located every second hour during the day before the release of weasels. The white bars correspond to voles which survived the entire experiment, while the black bars indicate the voles which were killed by the weasels. The numbers represent the sample sizes Proportion killed 1.0 Grey-sided vole Bank vole < Interfix distance (m) Fig. 4 Proportions of grey-sided voles (black bars) and bank voles (white bars) killed by weasels as a function of the voles movement distances. Interfix distances (distance moved between consecutive locations) were averaged over the entire experiment 12 body size and individual behaviour, prey individuals of a wider range of sizes should be used. However, size is usually age-dependent, which further complicates a study focussing on the size effect only. We found that weasels preferred male grey-sided voles over females, while such a preference was not found in bank voles. This is interesting, as in previous studies, female voles have been found to be more vulnerable than male voles for mammalian predation in general, and especially for small mustelid predation (Klemola et al. 1997; Norrdahl and Korpimäki 1998; Sundell 2003), or no preference has been found (Ylönen et al. 2003). However, it should be noted that none of these studies have used grey-sided voles as a prey species. Instead, in previous studies, observations have been obtained in nature and on Microtus voles. In the field, Microtus voles are forming dense matrilinear kin-clusters during the breeding season (Pusenius et al. 1997), which are likely to attract predators and may, therefore, explain the preference for Microtus females in nature (Norrdahl and Korpimäki 1993). If the common view that female voles in breeding condition should be more prone to weasel predation is true (e.g., Cushing 1985), then our experiment may better describe the situation outside the breeding season. It might well be that the marking behaviour characteristic for Myodes males (e.g., Horne and Ylönen 1998), together with a greater mobility (Norrdahl and Korpimäki 1998), may expose them to weasel predation more than females, but it does not explain why we did not find such a preference for male bank voles. Effect of weasels on space use and activity Grey-sided voles moved longer distances between consecutive radiolocations after the weasels were released into the enclosure, but the same trend was not found in bank voles. Those female grey-sided voles that moved more were killed by the weasels, while in males, more mobile or active individuals survived better than less active individuals. The same tendency was evident also in bank voles. Another result of our study was that the activity rhythm of voles depended on the activity rhythm of weasels; voles were active when weasels were inactive and vice versa. Soon after the weasel release, voles became active and started to move longer distances. This observation might reflect an attempt to move away from the weasels. Our visual observations support this, as we saw, on several occasions, voles running along the fences after the weasel release. Unfortunately, we did not have controls for these observations, i.e., voles that would have stayed for the same period in the enclosures as in our weasel treatment but without weasels. However, the activity peak right after weasel release was the highest one we observed and the
9 Popul Ecol (2008) 50: Fig. 5 The mean activity of all voles and weasels measured as interfix distances (distance moved between consecutive locations). The animals were located every second hour during the entire experiment. The vertical line shows the time of the weasel release. The mean activity of the voles (±SE) is shown as the black line with dots, while the grey bars represent the mean activity of the weasels. The error bars for the weasel observations are omitted for the sake of clarity Interfix distance (m) 18 Weasels Before weasel After weasel Voles Time (h) later peaks that coincided with weasels inactivity were not during the same time of the day in different days, which implies that they were likely to be direct reactions to weasel activity. The observation that activity patterns may change over time due to temporal variation in predation risk has also been made in earlier studies. For example, prey activity may first decrease, but if predation risk is present for a long period, the prey has to become active at some point to be able to, e.g., seek food (Lima and Bednekoff 1999). Previous studies on the activity of prey under predator presence have produced conflicting observations. Commonly, prey reduces activity, but, sometimes, an increase in activity has also been observed (e.g., Wooster and Sih 1995). Some of the differences can be caused by different measures used for activity, potentially reflecting very different types of activities (e.g., mobility, foraging, emigration). In general, reduced mobility should lead to lower encounter rates with predators and, consequently, to higher survival rates (Lima and Dill 1990). Norrdahl and Korpimäki (1998) found that voles moving long distances were killed more often than voles that moved less. This seemed to also be the case in females, but not in males, in the present experiment. Our observations of voles moving along the walls of the enclosures may indicate that they attempted to disperse longer distances than was possible in the closed enclosures. The U-shaped survival curve as a function of vole activity observed by Banks et al. (2000) was also apparent in our experiment if the two sexes were combined. When the sexes were investigated separately, the patterns of survival were mirror images. Banks et al. (2000) explained the phenomenon of nonlinearity in the predation risk so that reduced prey mobility leads to the accumulation of odorous waste products, which are used as cues by the predators. In the other extreme, very mobile individuals have an increased risk of random predator encounter. In our study, the same strategies to avoid weasel predation lead to a different success rates in female and male voles. In general, male voles have been observed to move more than females (Norrdahl and Korpimäki 1998). This is often explained by the mating system of voles, as male voles try to include as many female home ranges as possible within their own home ranges to be able to increase the fitness in terms of matings (Klemme et al. 2006). This implies that males are better adapted to longdistance and females to short-distance movements. In the light of this assumption, our results make sense, showing that females, adapted to move less, were vulnerable when they move more, while the opposite is evident for males. Conclusions We observed only weak evidence for the higher vulnerability of Microtus-type voles, here, the grey-sided vole, to small mustelid predation, and the evidence was unexpectedly stronger in open habitats than in forested habitats. Combined with observations of grey-sided voles in trees, this indicates that arboreal escape tactics are also used in this clumsy field-layer species. Thus, our results suggest that grey-sided voles may not be a true representative of Microtus-type voles, but, rather, an intermediate type between the two types. However, under natural conditions, grey-sided voles may have a higher vulnerability compared to bank voles due to their tendency to form relatively high-
10 266 Popul Ecol (2008) 50: density aggregations attracting predators. What part the generally larger size of the grey-sided vole compared to the bank vole plays in species-specific vulnerability remains to be tested with a wider size range of different sized individuals. Acknowledgments J. Gustafsson, A. Hämäläinen, J. Häyrinen, K. Juutilainen, E. Knuutila, S. Niskanen, A. Reckordt and H. Vihervaara are acknowledged for their assistance during the field work. We thank J.O. Wolff for the valuable comments on the manuscript and A.-L. Laine for helping with the statistics analysis. The study was funded by the Academy of Finland (project nos to JS, to HY and MH, and to OO), Swedish Research Council to TO and CIMO to LT. The experiment complies with the current laws of Finland. References Abrams PA, Holt RD, Roth JD (1998) Apparent competition or apparent mutualism? Shared predation when populations cycle. Ecology 79: Banks PB, Norrdahl K, Korpimäki E (2000) Nonlinearity in the predation risk of prey mobility. Proc R Soc Lond B 267: Chase JM, Abrams PA, Grover JP, Diehl S, Chesson P, Holt RD, Richards SA, Nisbet RM, Case TJ (2002) The interaction between predation and competition: a review and synthesis. Ecol Lett 5: Christensen P, Hörnfeldt B (2006) Habitat preferences of Clethrionomys rufocanus in boreal Sweden. Landscape Ecol 21: Cushing BS (1985) Estrous mice and vulnerability to weasel predation. Ecology 66: Hanski I, Henttonen H (1996) Predation on competing rodent species: a simple explanation of complex patterns. J Anim Ecol 65: Hanski I, Hansson L, Henttonen H (1991) Specialist predators, generalist predators, and the microtine rodent cycle. J Anim Ecol 60: Henttonen H (1987) The impact of spacing behavior in microtine rodents on the dynamics of least weasels Mustela nivalis a hypothesis. Oikos 50: Henttonen H (2000) Long-term dynamics of the bank vole Clethrionomys glareolus at Pallasjärvi, northern Finnish taiga. Polish J Ecol 48(Suppl):87 96 Henttonen H, Hansson L (1984) Interspecific relations between small rodents in European boreal and subarctic environments. Acta Zool Fenn 172:61 65 Holt RD (1977) Predation, apparent competition, and the structure of prey communities. Theor Popul Biol 12: Holt RD, Kotler BP (1987) Short-term apparent competition. Am Nat 130: Horne TJ, Ylönen H (1998) Heritabilities of dominance-related traits in male bank voles (Clethrionomys glareolus). Evolution 52: Jedrzejewska B, Jedrzejewski W (1990) Antipredatory behaviour of bank voles and prey choice of weasels enclosure experiments. Ann Zool Fenn 27: Jedrzejewski W, Jedrzejewska B (1990) Effect of a predator s visit on the spatial distribution of bank voles: experiments with weasels. Can J Zool 68: Jedrzejewski W, Rychlik L, Jedrzejewska B (1993) Responses of bank voles to odours of seven species of predators: experimental data and their relevance to natural predator vole relationships. Oikos 68: Jensen B (1994) Suomen ja Pohjolan nisäkkäät. WSOY, Porvoo (in Finnish) Kenward R (1987) Wildlife radio tagging: equipment, field techniques and data analysis. Academic Press, London Klemme I, Eccard JA, Gerlach G, Horne TJ, Ylönen H (2006) Does it pay to be a dominant male in a promiscuous species? Ann Zool Fenn 43: Klemola T, Koivula M, Korpimäki E, Norrdahl K (1997) Small mustelid predation slows population growth of Microtus voles: a predator reduction experiment. J Anim Ecol 66: Korpimäki E, Norrdahl K, Rinta-Jaskari T (1991) Responses of stoats and least weasels to fluctuating food abundances: is the low phase of the vole cycle due to mustelid predation? Oecologia 88: Krebs CJ (1999) Ecological methodology, 2nd edn. Benjamin Cummings, Menlo Park, California Lima SL, Bednekoff PA (1999) Temporal variation in danger drives antipredator behavior: the predation risk allocation hypothesis. Am Nat 153: Lima SL, Dill LM (1990) Behavioral decisions made under the risk of predation: a review and prospectus. Can J Zool 68: Menge BA, Sutherland JP (1976) Species diversity gradients: synthesis of the roles of predation, competition, and temporal heterogeneity. Am Nat 110: Menge BA, Sutherland JP (1987) Community regulation: variation in disturbance, competition, and predation in relation to environmental stress and recruitment. Am Nat 130: Norrdahl K, Korpimäki E (1993) Predation and interspecific competition in two Microtus voles. Oikos 67: Norrdahl K, Korpimäki E (1998) Does mobility or sex of voles affect risk of predation by mammalian predators? Ecology 79: Oksanen L (1992) Evolution of exploitation ecosystems I. Predation, foraging ecology and population dynamics in herbivores. Evol Ecol 6:15 33 Oksanen L, Lundberg P (1995) Optimization of reproductive effort and foraging time in mammals: the influence of resource level and predation risk. Evol Ecol 9:45 56 Pusenius J, Viitala J, Marienberg T, Ritvanen S (1998) Matrilineal kin clusters and their effect on reproductive success in the field vole Microtus agrestis. Behav Ecol 9:85 92 Sih A, Crowley P, McPeek M, Petranka J, Strohmeier K (1985) Predation, competition, and prey communities: a review of field experiments. Annu Rev Ecol Syst 16: Siivonen L, Sulkava S (1994) Mammals of northern Europe, 5th edn. Otava, Helsinki (in Finnish) Sundell J (2003) Population dynamics of microtine rodents: an experimental test of the predation hypothesis. Oikos 101: Sundell J, Ylönen H (2004) Behaviour and choice of refuge by voles under predation risk. Behav Ecol Sociobiol 56: Sundell J, Norrdahl K, Korpimäki E, Hanski I (2000) Functional response of the least weasel, Mustela nivalis nivalis. Oikos 90: Sundell J, Eccard JA, Tiilikainen R, Ylönen H (2003) Predation rate, prey preference and predator switching: experiments on voles and weasels. Oikos 101: Viitala J (1977) Social organization in cyclic subarctic populations of the vole Clethrionomys rufocanus (Sund.) and Microtus agrestis (L.). Ann Zool Fenn 14:53 93 Wilson DE, Reeder DM (2005) Mammal species of the world, 3rd edn. Johns Hopkins University Press, Baltimore, Maryland Wooster D, Sih A (1995) A review of the drift and activity responses of stream prey to predator presence. Oikos 73:3 8 Ylönen H, Sundell J, Tiilikainen R, Eccard JA, Horne TJ (2003) Weasels (Mustela nivalis nivalis) preference for olfactory cues of the vole (Clethrionomys glareolus). Ecology 84:
Behaviour and resource use of two competing vole species under shared predation risk
Oecologia (2008) 157:707 715 DOI 10.1007/s00442-008-1099-6 BEHAVIORAL ECOLOGY - ORIGINAL PAPER Behaviour and resource use of two competing vole species under shared predation risk Lenka Trebatická Janne
More informationIn multipredator environments, animals encounter the risk
Behavioral Ecology Vol. 7 No. : 0-4 Microhabitat use and behavior of voles under weasel and raptor predation risk: predator facilitation? Erkki Korpimaki, Vesa Koivunen, and Hani Hakkarainen Laboratory
More informationPopulation dynamics of small game. Pekka Helle Natural Resources Institute Finland Luke Oulu
Population dynamics of small game Pekka Helle Natural Resources Institute Finland Luke Oulu Populations tend to vary in size temporally, some species show more variation than others Depends on degree of
More informationHabitat selection of grey-sided voles and bank voles in two subalpine populations in southern Norway
Ann. Zool. Fennici 36: 215 222 ISSN 0003-455X Helsinki 17 December 1999 Finnish Zoological and Botanical Publishing Board 1999 Habitat selection of grey-sided voles and bank voles in two subalpine populations
More informationUral owl predation on field voles and bank voles by size, sex and reproductive state
Ann. Zool. Fennici 47: 90 98 ISSN 0003-455X (print), ISSN 1797-2450 (online) Helsinki 30 April 2010 Finnish Zoological and Botanical Publishing Board 2010 Ural owl predation on field voles and bank voles
More informationBehavioural response of field voles under mustelid predation risk in the laboratory: more than neophobia
Ann. Zool. Fennici 37: 169 178 ISSN 0003-455X Helsinki 27 October 2000 Finnish Zoological and Botanical Publishing Board 2000 Behavioural response of field voles under mustelid predation risk in the laboratory:
More informationrodent species in Australia to the fecal odor of various predators. Rattus fuscipes (bush
Sample paper critique #2 The article by Hayes, Nahrung and Wilson 1 investigates the response of three rodent species in Australia to the fecal odor of various predators. Rattus fuscipes (bush rat), Uromys
More informationFungal-Mediated Multitrophic Interactions - Do Grass Endophytes in Diet Protect Voles from Predators?
Fungal-Mediated Multitrophic Interactions - Do Grass Endophytes in Diet Protect Voles from Predators? Susanna Saari 1,2 *, Janne Sundell 3,6, Otso Huitu 4, Marjo Helander 2, Elise Ketoja 5, Hannu Ylönen
More informationSupplementary Fig. 1: Comparison of chase parameters for focal pack (a-f, n=1119) and for 4 dogs from 3 other packs (g-m, n=107).
Supplementary Fig. 1: Comparison of chase parameters for focal pack (a-f, n=1119) and for 4 dogs from 3 other packs (g-m, n=107). (a,g) Maximum stride speed, (b,h) maximum tangential acceleration, (c,i)
More informationLynx Update May 25, 2009 INTRODUCTION
Lynx Update May 25, 2009 INTRODUCTION In an effort to establish a viable population of Canada lynx (Lynx canadensis) in Colorado, the Colorado Division of Wildlife (CDOW) initiated a reintroduction effort
More informationTHE WOLF WATCHERS. Endangered gray wolves return to the American West
CHAPTER 7 POPULATION ECOLOGY THE WOLF WATCHERS Endangered gray wolves return to the American West THE WOLF WATCHERS Endangered gray wolves return to the American West Main concept Population size and makeup
More informationSCIENTIFIC REPORT. Analysis of the baseline survey on the prevalence of Salmonella in turkey flocks, in the EU,
The EFSA Journal / EFSA Scientific Report (28) 198, 1-224 SCIENTIFIC REPORT Analysis of the baseline survey on the prevalence of Salmonella in turkey flocks, in the EU, 26-27 Part B: factors related to
More information2 Introduction. in this web service Cambridge University Press. Fig. 1.1.
1 Introduction Vertebrate predators, particularly large charismatic apex predators, have traditionally elicited much esteem and deference, because they have been depicted as being beautiful, majestic and
More informationMice alone and their biodiversity impacts: a 5-year experiment at Maungatautari
Mice alone and their biodiversity impacts: a 5-year experiment at Maungatautari Deb Wilson, Corinne Watts, John Innes, Neil Fitzgerald, Scott Bartlam, Danny Thornburrow, Cat Kelly, Gary Barker, Mark Smale,
More informationAmes, IA Ames, IA (515)
BENEFITS OF A CONSERVATION BUFFER-BASED CONSERVATION MANAGEMENT SYSTEM FOR NORTHERN BOBWHITE AND GRASSLAND SONGBIRDS IN AN INTENSIVE PRODUCTION AGRICULTURAL LANDSCAPE IN THE LOWER MISSISSIPPI ALLUVIAL
More informationYounger bank voles are more vulnerable to avian predation
1074 NOTE / NOTE Younger bank voles are more vulnerable to avian predation Taru Meri, Matti Halonen, Tapio Mappes, and Jukka Suhonen Abstract: The importance of predation on prey populations is mainly
More informationSelection for Egg Mass in the Domestic Fowl. 1. Response to Selection
Selection for Egg Mass in the Domestic Fowl. 1. Response to Selection H. L. MARKS US Department of Agriculture, Science & Education Administration, Agricultural Research, uthern Regional Poultry Breeding
More informationY Use of adaptive management to mitigate risk of predation for woodland caribou in north-central British Columbia
Y093065 - Use of adaptive management to mitigate risk of predation for woodland caribou in north-central British Columbia Purpose and Management Implications Our goal was to implement a 3-year, adaptive
More informationEffects of Cage Stocking Density on Feeding Behaviors of Group-Housed Laying Hens
AS 651 ASL R2018 2005 Effects of Cage Stocking Density on Feeding Behaviors of Group-Housed Laying Hens R. N. Cook Iowa State University Hongwei Xin Iowa State University, hxin@iastate.edu Recommended
More informationLimitation of reproductive success by food availability and litter size in the bank vole, Clethrionomys glareolus
Limitation of reproductive success by food availability and litter size in the bank vole, Clethrionomys glareolus Esa Koskela 1*, Pernilla Jonsson 2, Tommi Hartikainen 1 and Tapio Mappes 1 1 Department
More informationBROOD REDUCTION IN THE CURVE-BILLED THRASHER By ROBERTE.RICKLEFS
Nov., 1965 505 BROOD REDUCTION IN THE CURVE-BILLED THRASHER By ROBERTE.RICKLEFS Lack ( 1954; 40-41) has pointed out that in species of birds which have asynchronous hatching, brood size may be adjusted
More informationJournal of Animal Ecology (1988), 57, THE IMPACT OF PREDATION ON BOREAL TETRAONIDS DURING VOLE CYCLES: AN EXPERIMENTAL STUDY
Journal of Animal Ecology (1988), 57, 859-872 THE IMPACT OF PREDATION ON BOREAL TETRAONIDS DURING VOLE CYCLES: AN EXPERIMENTAL STUDY BY V. MARCSTROM*, R. E. KENWARD$ AND E. ENGREN* * Institute of Zoophysiology,
More informationSheikh Muhammad Abdur Rashid Population ecology and management of Water Monitors, Varanus salvator (Laurenti 1768) at Sungei Buloh Wetland Reserve,
Author Title Institute Sheikh Muhammad Abdur Rashid Population ecology and management of Water Monitors, Varanus salvator (Laurenti 1768) at Sungei Buloh Wetland Reserve, Singapore Thesis (Ph.D.) National
More informationWater Vole Translocation Project: Abberton ReservoirAbout Water Voles Population Dynamics
Water Vole Translocation Project: Abberton ReservoirAbout Water Voles Measuring up to 24cm, water voles (Arvicola amphibius) are the largest of the British voles and at a quick glace, are often mistaken
More informationAN APPLIED CASE STUDY of the complexity of ecological systems and process: Why has Lyme disease become an epidemic in the northeastern U.S.
AN APPLIED CASE STUDY of the complexity of ecological systems and process: Why has Lyme disease become an epidemic in the northeastern U.S. over the last few decades? What causes Lyme disease? 1 Frequency
More informationOverview of some of the latest development and new achievement of rabbit science research in the E.U.
First Jilin Rabbit Fair and Conference on Asian Rabbit Production Development, Changchun (China), 8-10 Septembre 2009. Overview of some of the latest development and new achievement of rabbit science research
More informationDoug Manzer, Kyle Prince, Blair Seward, Layne Seward and Mike Uchikura
Alberta Conservation Association (ACA) Date: 2014-2015 Project Name: Upland Gamebird Studies Wildlife Program Manager: Doug Manzer Project Leader: Layne Seward Primary ACA staff on project: Doug Manzer,
More informationAdjustments In Parental Care By The European Starling (Sturnus Vulgaris): The Effect Of Female Condition
Proceedings of The National Conference on Undergraduate Research (NCUR) 2003 University of Utah, Salt Lake City, Utah March 13-15, 2003 Adjustments In Parental Care By The European Starling (Sturnus Vulgaris):
More informationOriginal Draft: 11/4/97 Revised Draft: 6/21/12
Original Draft: 11/4/97 Revised Draft: 6/21/12 Dear Interested Person or Party: The following is a scientific opinion letter requested by Brooks Fahy, Executive Director of Predator Defense. This letter
More information4B: The Pheasant Case: Handout. Case Three Ring-Necked Pheasants. Case materials: Case assignment
4B: The Pheasant Case: Handout Case Three Ring-Necked Pheasants As you can see, the male ring-necked pheasant is brightly colored. The white ring at the base of the red and green head stand out against
More informationHabitat Use and Survival of Gray Partridge Pairs in Bavaria, Germany
National Quail Symposium Proceedings Volume 6 Article 19 2009 Habitat Use and Survival of Gray Partridge Pairs in Bavaria, Germany Wolfgang Kaiser Ilse Storch University of Freiburg John P. Carroll University
More informationGREATER SAGE-GROUSE BROOD-REARING HABITAT MANIPULATION IN MOUNTAIN BIG SAGEBRUSH, USE OF TREATMENTS, AND REPRODUCTIVE ECOLOGY ON PARKER MOUNTAIN, UTAH
GREATER SAGE-GROUSE BROOD-REARING HABITAT MANIPULATION IN MOUNTAIN BIG SAGEBRUSH, USE OF TREATMENTS, AND REPRODUCTIVE ECOLOGY ON PARKER MOUNTAIN, UTAH Abstract We used an experimental design to treat greater
More informationGUIDELINES ON CHOOSING THE CORRECT ERADICATION TECHNIQUE
GUIDELINES ON CHOOSING THE CORRECT ERADICATION TECHNIQUE PURPOSE... 2 1. RODENTS... 2 1.1 METHOD PROS AND CONS... 3 1.1. COMPARISON BETWEEN BROUDIFACOUM AND DIPHACINONE... 4 1.2. DISCUSSION ON OTHER POSSIBLE
More informationPopulation Dynamics: Predator/Prey Teacher Version
Population Dynamics: Predator/Prey Teacher Version In this lab students will simulate the population dynamics in the lives of bunnies and wolves. They will discover how both predator and prey interact
More informationIntegrated Management of Invasive Geese Populations in an International Context: a Case Study
Integrated Management of Invasive Geese Populations in an International Context: a Case Study Tim Adriaens, Frank Huysentruyt, Sander Devisscher, Koen Devos & Jim Casaer Neobiota 2014 4/11/2014, Antalya
More informationEvaluation of Horn Flies and Internal Parasites with Growing Beef Cattle Grazing Bermudagrass Pastures Findings Materials and Methods Introduction
Evaluation of Horn Flies and Internal Parasites with Growing Beef Cattle Grazing Bermudagrass Pastures S. M. DeRouen, Hill Farm Research Station; J.E. Miller, School of Veterinary Medicine; and L. Foil,
More informationEgg laying site preferences in Pterostichus melanarius Illiger (Coleoptera: Carabidae)
Egg laying site preferences in Pterostichus melanarius Illiger (Coleoptera: Carabidae) H. Tréfás & J.C. van Lenteren Laboratory of Entomology, Wageningen University and Research Centre, Binnenhaven 7,
More informationTesting the Ideal Free Distribution on Turtles in the Field
Testing the Ideal Free Distribution on Turtles in the Field Justin Carasa Nicole Cinquino Christopher Contreras Santiago Londoño Michelle Ortiz Andrea Remiro Alexander Rodriguez Research in Ecology University
More informationThis article is downloaded from.
This article is downloaded from http://researchoutput.csu.edu.au It is the paper published as: Author: A. Wichman, L. Rogers and R. Freire Title: Visual lateralisation and development of spatial and social
More informationP VASANTA KUMARI and JAMIL AHMAD KHAN Department of Zoology, Aligarh Muslim University, Aligarh
Proc. Indian Acad. Sci., Vol. 87 B, No.9, (Animal Sciences-J), September 1978, pp. 285-291, printed in 1ndia. Retrieval of young by lactating Indian gerbil, indica (Hardwicke) Tatera indica P VASANTA KUMARI
More informationRELATIONSHIPS AMONG WEIGHTS AND CALVING PERFORMANCE OF HEIFERS IN A HERD OF UNSELECTED CATTLE
RELATIONSHIPS AMONG WEIGHTS AND CALVING PERFORMANCE OF HEIFERS IN A HERD OF UNSELECTED CATTLE T. C. NELSEN, R. E. SHORT, J. J. URICK and W. L. REYNOLDS1, USA SUMMARY Two important traits of a productive
More informationDO BROWN-HEADED COWBIRDS LAY THEIR EGGS AT RANDOM IN THE NESTS OF RED-WINGED BLACKBIRDS?
Wilson Bull., 0(4), 989, pp. 599605 DO BROWNHEADED COWBIRDS LAY THEIR EGGS AT RANDOM IN THE NESTS OF REDWINGED BLACKBIRDS? GORDON H. ORTANS, EIVIN RDSKAPT, AND LES D. BELETSKY AssrnAcr.We tested the hypothesis
More informationLAB. NATURAL SELECTION
Period Date LAB. NATURAL SELECTION This game was invented by G. Ledyard Stebbins, a pioneer in the evolution of plants. The purpose of the game is to illustrate the basic principles and some of the general
More informationNathan A. Thompson, Ph.D. Adjunct Faculty, University of Cincinnati Vice President, Assessment Systems Corporation
An Introduction to Computerized Adaptive Testing Nathan A. Thompson, Ph.D. Adjunct Faculty, University of Cincinnati Vice President, Assessment Systems Corporation Welcome! CAT: tests that adapt to each
More informationComparative Evaluation of Online and Paper & Pencil Forms for the Iowa Assessments ITP Research Series
Comparative Evaluation of Online and Paper & Pencil Forms for the Iowa Assessments ITP Research Series Catherine J. Welch Stephen B. Dunbar Heather Rickels Keyu Chen ITP Research Series 2014.2 A Comparative
More informationEnrichments for captive Andean Condor (Vultur gryphus) in Zuleta, North Ecuador. Yann Potaufeu (2014)
Enrichments for captive Andean Condor (Vultur gryphus) in Zuleta, North Ecuador Yann Potaufeu (2014) 1 Introduction Over recent decades, enrichment has been shown to be an important component for the well-being
More informationESTIMATING NEST SUCCESS: WHEN MAYFIELD WINS DOUGLAS H. JOHNSON AND TERRY L. SHAFFER
ESTIMATING NEST SUCCESS: WHEN MAYFIELD WINS DOUGLAS H. JOHNSON AND TERRY L. SHAFFER U.S. Fish and Wildlife Service, Northern Prairie Wildlife Research Center, Jamestown, North Dakota 58402 USA ABSTRACT.--The
More informationCIWF Response to the Coalition for Sustainable Egg Supply Study April 2015
CIWF Response to the Coalition for Sustainable Egg Supply Study April 2015 The Coalition for Sustainable Egg Supply study seeks to understand the sustainability impacts of three laying hen housing systems
More informationPilot study to identify risk factors for coprophagic behaviour in dogs
Pilot study to identify risk factors for coprophagic behaviour in dogs Joanne A.M. van der Borg and Lisette Graat Wageningen University Introduction According to several training centres of guide dogs
More informationMate protection in pre-nesting Canada Geese Branta canadensis
Mate protection in pre-nesting Canada Geese Branta canadensis I. P. JOHNSON and R. M. SIBLY Fourteen individually marked pairs o f Canada Geese were observedfrom January to April on their feeding grounds
More informationSurvivorship. Demography and Populations. Avian life history patterns. Extremes of avian life history patterns
Demography and Populations Survivorship Demography is the study of fecundity and survival Four critical variables Age of first breeding Number of young fledged each year Juvenile survival Adult survival
More informationCiccaba virgata (Mottled Owl)
Ciccaba virgata (Mottled Owl) Family: Strigidae (Typical Owls) Order: Strigiformes (Owls) Class: Aves (Birds) Fig. 1. Mottled owl, Ciccaba virgata. [http://www.owling.com/mottled13.htm, downloaded 12 November
More informationUse of Agent Based Modeling in an Ecological Conservation Context
28 RIThink, 2012, Vol. 2 From: http://photos.turksandcaicostourism.com/nature/images/tctb_horz_033.jpg Use of Agent Based Modeling in an Ecological Conservation Context Scott B. WOLCOTT 1 *, Michael E.
More informationSupporting Online Material for
www.sciencemag.org/cgi/content/full/314/5802/1111/dc1 Supporting Online Material for Rapid Temporal Reversal in Predator-Driven Natural Selection Jonathan B. Losos,* Thomas W. Schoener, R. Brian Langerhans,
More informationTemperature Gradient in the Egg-Laying Activities of the Queen Bee
The Ohio State University Knowledge Bank kb.osu.edu Ohio Journal of Science (Ohio Academy of Science) Ohio Journal of Science: Volume 30, Issue 6 (November, 1930) 1930-11 Temperature Gradient in the Egg-Laying
More informationGenetic analysis of mentality traits in Rhodesian Ridgeback dogs
Genetic analysis of mentality traits in Rhodesian Ridgeback dogs Kamilla Lysaker, Tormod Ådnøy, Per Arvelius* and Odd Vangen, Department of Animal and Aquacultural Sciences, Ås, Norway *Swedish University
More informationBy Hans Frey ¹ ² & Alex Llopis ²
1/7 By Hans Frey ¹ ² & Alex Llopis ² ¹ Verein EGS-Eulen und Greifvogelschutz, Untere Hauptstraße 34, 2286 Haringsee, Austria. Phone number +43 2214 84014 h.frey@4vultures.org ² Vulture Conservation Foundation
More informationSHEEP SIRE REFERENCING SCHEMES - NEW OPPORTUNITIES FOR PEDIGREE BREEDERS AND LAMB PRODUCERS a. G. Simm and N.R. Wray
SHEEP SIRE REFERENCING SCHEMES - NEW OPPORTUNITIES FOR PEDIGREE BREEDERS AND LAMB PRODUCERS a G. Simm and N.R. Wray The Scottish Agricultural College Edinburgh, Scotland Summary Sire referencing schemes
More informationHomework Case Study Update #3
Homework 7.1 - Name: The graph below summarizes the changes in the size of the two populations you have been studying on Isle Royale. 1996 was the year that there was intense competition for declining
More informationBreeding success of Greylag Geese on the Outer Hebrides, September 2016
Breeding success of Greylag Geese on the Outer Hebrides, September 2016 Wildfowl & Wetlands Trust Report Author Carl Mitchell September 2016 The Wildfowl & Wetlands Trust All rights reserved. No part of
More informationEFFECTS OF FOOD SUPPLEMENTATION ON THE SOCIAL ORGANIZATION OF PRAIRIE VOLES (MICROTUS OCHROGASTER)
Journal of Mammalogy, 81(3):746 757, 2000 EFFECTS OF FOOD SUPPLEMENTATION ON THE SOCIAL ORGANIZATION OF PRAIRIE VOLES (MICROTUS OCHROGASTER) GRAHAM R. COCHRAN AND NANCY G. SOLOMON* Department of Zoology,
More informationGreat Horned Owl (Bubo virginianus) Productivity and Home Range Characteristics in a Shortgrass Prairie. Rosemary A. Frank and R.
Great Horned Owl (Bubo virginianus) Productivity and Home Range Characteristics in a Shortgrass Prairie Rosemary A. Frank and R. Scott Lutz 1 Abstract. We studied movements and breeding success of resident
More informationINHERITANCE OF BODY WEIGHT IN DOMESTIC FOWL. Single Comb White Leghorn breeds of fowl and in their hybrids.
440 GENETICS: N. F. WATERS PROC. N. A. S. and genetical behavior of this form is not incompatible with the segmental interchange theory of circle formation in Oenothera. Summary.-It is impossible for the
More informationContrasting Response to Predator and Brood Parasite Signals in the Song Sparrow (melospiza melodia)
Luke Campillo and Aaron Claus IBS Animal Behavior Prof. Wisenden 6/25/2009 Contrasting Response to Predator and Brood Parasite Signals in the Song Sparrow (melospiza melodia) Abstract: The Song Sparrow
More information[ 144 ] THE GROWTH AND DEVELOPMENT OF MICE IN THREE CLIMATIC ENVIRONMENTS
[ ] THE GROWTH AND DEVELOPMENT OF MICE IN THREE CLIMATIC ENVIRONMENTS BY J. D. BIGGERS, M. R. ASHOUB,* ANNE McLAREN AND DONALD MICHIE Royal Veterinary College, London, N. W. i {Received September 9) INTRODUCTION
More informationBiol 160: Lab 7. Modeling Evolution
Name: Modeling Evolution OBJECTIVES Help you develop an understanding of important factors that affect evolution of a species. Demonstrate important biological and environmental selection factors that
More informationAN APPLIED CASE STUDY of the complexity of ecological systems and process: Why has Lyme disease become an epidemic in the northeastern U.S.
AN APPLIED CASE STUDY of the complexity of ecological systems and process: Why has Lyme disease become an epidemic in the northeastern U.S. over the last few decades? What causes Lyme disease? 1 Frequency
More informationCoyote. Canis latrans. Other common names. Introduction. Physical Description and Anatomy. Eastern Coyote
Coyote Canis latrans Other common names Eastern Coyote Introduction Coyotes are the largest wild canine with breeding populations in New York State. There is plenty of high quality habitat throughout the
More informationHabitats and Field Methods. Friday May 12th 2017
Habitats and Field Methods Friday May 12th 2017 Announcements Project consultations available today after class Project Proposal due today at 5pm Follow guidelines posted for lecture 4 Field notebooks
More informationGreenham Common, Crookham Common and Bowdown Wood Reptile Survey 2010
Greenham Common, Crookham Common and Bowdown Wood Reptile Survey 2010 Rod d Ayala and Martin Burdock, November 2010 CONTENTS Text Executive Summary Page 3 Background Page 4 Survey Methods, Personnel and
More informationRODENTS OF THE GREATER AUCKLAND REGION. by John L. Craig SUMMARY
TANE 29, 1983 RODENTS OF THE GREATER AUCKLAND REGION by John L. Craig Department of Zoology, University of Auckland, Private Bag, Auckland SUMMARY Four rodent species are known in the Greater Auckland
More informationThe effect of weaning weight on subsequent lamb growth rates
Proceedings of the New Zealand Grassland Association 62: 75 79 (2000) 75 The effect of weaning weight on subsequent lamb growth rates T.J. FRASER and D.J. SAVILLE AgResearch, PO Box 60, Lincoln, Canterbury
More informationPROBABLE NON-BREEDERS AMONG FEMALE BLUE GROUSE
Condor, 81:78-82 0 The Cooper Ornithological Society 1979 PROBABLE NON-BREEDERS AMONG FEMALE BLUE GROUSE SUSAN J. HANNON AND FRED C. ZWICKEL Parallel studies on increasing (Zwickel 1972) and decreasing
More informationEffects of prey availability and climate across a decade for a desert-dwelling, ectothermic mesopredator. R. Anderson Western Washington University
Effects of prey availability and climate across a decade for a desert-dwelling, ectothermic mesopredator R. Anderson Western Washington University Trophic interactions in desert systems are presumed to
More informationGenetic parameters and breeding value stability estimated from a joint evaluation of purebred and crossbred sows for litter weight at weaning
Acta Agraria Kaposváriensis (2015) Vol 19 No 1, 1-7. Kaposvári Egyetem, Agrár- és Környezettudományi Kar, Kaposvár Genetic parameters and breeding value stability estimated from a joint evaluation of purebred
More informationHow Does Photostimulation Age Alter the Interaction Between Body Size and a Bonus Feeding Program During Sexual Maturation?
16 How Does Photostimulation Age Alter the Interaction Between Body Size and a Bonus Feeding Program During Sexual Maturation? R A Renema*, F E Robinson*, and J A Proudman** *Alberta Poultry Research Centre,
More informationWeaver Dunes, Minnesota
Hatchling Orientation During Dispersal from Nests Experimental analyses of an early life stage comparing orientation and dispersal patterns of hatchlings that emerge from nests close to and far from wetlands
More informationREPORT ON SCOTTISH EID TRIALS
REPORT ON SCOTTISH EID TRIALS PREPARED FOR: SEERAD PREPARED BY: SAOS Ltd Rural Centre West Mains Ingliston, EH28 8NZ January 2007 CONTENTS 1. Introduction 2 Page 2. Trial Objectives. 2 3. Methodology..
More informationManagement of bold wolves
Policy Support Statements of the Large Carnivore Initiative for Europe (LCIE). Policy support statements are intended to provide a short indication of what the LCIE regards as being good management practice
More informationAggressive Ural owl mothers recruit more offspring
Behavioral Ecology doi:10.1093/beheco/arp062 Advance Access publication 6 May 2009 Aggressive Ural owl mothers recruit more offspring Pekka Kontiainen, a Hannu Pietiäinen, a Kalle Huttunen, a Patrik Karell,
More informationLEVEL 2 AWARD IN THE SAFE USE OF RODENTICIDES
LEVEL 2 AWARD IN THE SAFE USE OF RODENTICIDES Specimen Paper PAPER NUMBER: 0000 INSTRUCTIONS TO CANDIDATES 1 Attempt to answer every question. 2. Enter your answers on the accompanying answer sheet. Each
More informationBio4009 : Projet de recherche/research project
Bio4009 : Projet de recherche/research project Is emergence after hibernation of the black ratsnake (Elaphe obsoleta) triggered by a thermal gradient reversal? By Isabelle Ceillier 4522350 Supervisor :
More informationThe Effects of Meso-mammal Removal on Northern Bobwhite Populations
The Effects of Meso-mammal Removal on Northern Bobwhite Populations Alexander L. Jackson William E. Palmer D. Clay Sisson Theron M. Terhune II John M. Yeiser James A. Martin Predation Predation is the
More informationSomatic Cell Count as an Indicator of Subclinical Mastitis. Genetic Parameters and Correlations with Clinical Mastitis
Somatic Cell Count as an Indicator of Subclinical Mastitis. Genetic Parameters and Correlations with Clinical Mastitis Morten Svendsen 1 and Bjørg Heringstad 1,2 1 GENO Breeding and A.I. Association, P.O
More informationDoes supplementary feeding reduce predation of red grouse by hen harriers?
Ecology 2001 38, Blackwell Oxford, JPE Journal 0021-8901 British December 38 6000 Ecological of UK Science 2001 Applied Ltd Society, Ecology2001 PRIORITY CONTRIBUTION Supplementary S.M. Redpath, S.J. feeding
More informationABSTRACT. Ashmore Reef
ABSTRACT The life cycle of sea turtles is complex and is not yet fully understood. For most species, it involves at least three habitats: the pelagic, the demersal foraging and the nesting habitats. This
More informationRemoval of Alaskan Bald Eagles for Translocation to Other States Michael J. Jacobson U.S Fish and Wildlife Service, Juneau, AK
Removal of Alaskan Bald Eagles for Translocation to Other States Michael J. Jacobson U.S Fish and Wildlife Service, Juneau, AK Bald Eagles (Haliaeetus leucocephalus) were first captured and relocated from
More informationThe Arctic fox in Scandinavia yesterday, today and tomorrow.
The Arctic fox in Scandinavia yesterday, today and tomorrow. The biology of the Arctic fox The Arctic fox is a small fox that is found in Arctic and subarctic areas around the northern hemisphere in Siberia,
More informationPrey refuges as predator hotspots: ocelot (Leopardus pardalis) attraction to agouti (Dasyprocta punctata) dens
Acta Theriol (2014) 59:257 262 DOI 10.1007/s13364-013-0159-4 ORIGINAL PAPER Prey refuges as predator hotspots: ocelot (Leopardus pardalis) attraction to agouti (Dasyprocta punctata) dens Willem-Jan Emsens
More informationPROGRESS REPORT for COOPERATIVE BOBCAT RESEARCH PROJECT. Period Covered: 1 April 30 June Prepared by
PROGRESS REPORT for COOPERATIVE BOBCAT RESEARCH PROJECT Period Covered: 1 April 30 June 2014 Prepared by John A. Litvaitis, Tyler Mahard, Rory Carroll, and Marian K. Litvaitis Department of Natural Resources
More informationReintroducing bettongs to the ACT: issues relating to genetic diversity and population dynamics The guest speaker at NPA s November meeting was April
Reintroducing bettongs to the ACT: issues relating to genetic diversity and population dynamics The guest speaker at NPA s November meeting was April Suen, holder of NPA s 2015 scholarship for honours
More informationANIMAL BEHAVIOR. Laboratory: a Manual to Accompany Biology. Saunders College Publishing: Philadelphia.
PRESENTED BY KEN Yasukawa at the 2007 ABS Annual Meeting Education Workshop Burlington VT ANIMAL BEHAVIOR Humans have always been interested in animals and how they behave because animals are a source
More informationTECHNICAL BULLETIN Claude Toudic Broiler Specialist June 2006
Evaluating uniformity in broilers factors affecting variation During a technical visit to a broiler farm the topic of uniformity is generally assessed visually and subjectively, as to do the job properly
More informationSTATISTICAL REPORT. Preliminary Analysis of the Second Collaborative Study of the Hard Surface Carrier Test
STATISTICAL REPORT To: From: Subject: Diane Boesenberg, Reckitt Benckiser Emily Mitchell, Product Science Branch, Antimicrobials Division/Office of Pesticide Programs/US EPA Martin Hamilton, Statistician
More informationSlide 1. Slide 2. Slide 3 Population Size 450. Slide 4
Slide 1 Slide 2 The science behind management of game birds, predators, and landscapes of the Midwest: the ups and downs of pheasant populations William R. Clark Iowa State University Iowa DNR, DU- IWWR,
More informationIntraspecific relationships extra questions and answers (Extension material for Level 3 Biology Study Guide, ISBN , page 153)
i Intraspecific relationships extra questions and answers (Extension material for Level 3 Biology Study Guide, ISBN 978-1-927194-58-4, page 153) Activity 9: Intraspecific relationships extra questions
More informationBelow, we present the methods used to address these objectives, our preliminary results and next steps in this multi-year project.
Background Final Report to the Nova Scotia Habitat Conservation Fund: Determining the role of food availability on swallow population declines Project Supervisor: Tara Imlay, tara.imlay@dal.ca In the past
More informationLizard Surveying and Monitoring in Biodiversity Sanctuaries
Lizard Surveying and Monitoring in Biodiversity Sanctuaries Trent Bell (EcoGecko Consultants) Alison Pickett (DOC North Island Skink Recovery Group) First things first I am profoundly deaf I have a Deaf
More informationNATAL DISPERSAL OF SNOWSHOE HARES DURING A CYCLIC POPULATION INCREASE
NATAL DISPERSAL OF SNOWSHOE HARES DURING A CYCLIC POPULATION INCREASE ELIZABETH A. GILLIS AND CHARLES J. KREBS Department of Zoology, University of British Columbia, 6270 University Boulevard, Vancouver,
More informationAnalysis of Sampling Technique Used to Investigate Matching of Dorsal Coloration of Pacific Tree Frogs Hyla regilla with Substrate Color
Analysis of Sampling Technique Used to Investigate Matching of Dorsal Coloration of Pacific Tree Frogs Hyla regilla with Substrate Color Madeleine van der Heyden, Kimberly Debriansky, and Randall Clarke
More information