Bark salicylates and condensed tannins reduce vole browsing amongst cultivated dark-leaved willows (<Emphasis Type="Italic">Salix myrsinifolia</Emphasis>)

Summary. Vole feeding amongst herbal willows that have a high concentration of salicylates in their bark and leaves, and may therefore be cultivated for use as raw material for herbal medicine was tested in the field and in laboratory conditions. Eight clones of dark-leaved willow (Salix myrsinifolia Salisb.) were cultivated for two years with six different methods combining three fertilisation levels (none, low and high), black plastic mulch applied for suppressing weed competition and unmulched control. Samples for the laboratory feeding trial were taken from the unfertilised plants during willow winter dormancy and twigs were fed to 16 voles as a multi-choice experiment. The bark area removed was calculated from image analysis of the material left by the voles. The diameter and the bark thickness of the twigs were measured. Concentrations of salicin, salicortin, HCH-salicortin, picein, triandrin, triandrin derivative, gallocatechin, (+)-catechin, luteolin-7-glucoside, hyperin, total condensed tannins and total nitrogen were measured from the twigs fed to voles in the laboratory. Browsing by a natural population of voles amongst winter-dormant willows was measured in the field. In the laboratory, voles browsed on 80% of the twigs and in the field voles browsed on 33% of the twigs. Vole feeding followed similar patterns in the field and in the laboratory experiment; feeding was clearly higher amongst the plants grown in unmulched control compared to those in plastic mulch. The same clones, 1, 2 and 6 were preferred in both experiments. Voles preferred thin twigs to thick ones. Feeding correlated negatively with concentrations of salicylates and tannins. As vole feeding seems to be highly affected by willow cultivation method and plant genotype, careful selection of cultivated clones and cultivation methods can enhance the reliability of herbal willow cultivation.     

Different escape tactics of two vole species affect the success of the hunting predator, the least weasel

In the ongoing evolutionary arms race between predators and their prey, successful escape from the predator leads to the evolution of improved escape tactics in prey, but also predators become more effective in following and attacking the prey. Antipredatory behavior of prey is considered to be the strongest towards their most dangerous predators. However, prey species can differ both in vulnerability and efficiency of escape to a shared predator. We studied escape reactions of two vole species, the bank vole (Myodes glareolus) and the field vole (Microtus agrestis), under a simulated predation risk of the least weasel (Mustela nivalis nivalis). We conducted a laboratory experiment where a vole was given a possibility to escape from a weasel by fleeing to a horizontal tunnel or climbing the tree. Subsequently to the vole escape decision, we released a weasel to the same tunnel system to test how the weasel succeeded in following the vole. Weasel presence changed the behavior of voles as especially bank voles escaped by climbing. Instead, the majority of field voles fled into the ground-layer tunnel. The different escape tactics of the voles affected the success of the weasel, because climbing voles were less often successfully followed. We suggest that the difference in escape tactics has evolved as an adaptation to different habitats; meadow-exploiting field voles using ground-level escape while bank voles living in three-dimensional forest habitat frequently use arboreal escape tactics. This is likely to lead to different habitat-dependent vulnerabilities to predation in Microtus and Myodes vole species.     

Leaf ontogeny interacts with Bt modification to affect innate resistance in GM aspens

Bioassays with a non-target slug (Deroceras spp.) and chemical analyses were conducted using leaf tissue from already existing genetically modified insect-resistant aspen trees to examine whether genetic modifications to produce Bacillus thuringiensis (Bt) toxins could affect plant phytochemistry, which in turn might influence plant–herbivore interactions. Three major patterns emerged. First, two independent modifications for Bt resistance affected the phytochemical profiles of leaves such that both were different from the isogenic wild-type (Wt) control leaves, but also different from each other. Among the contributors to these differences are substances with a presumed involvement in resistance, such as salicortin and soluble condensed tannins. Second, bioassays with one Bt line suggest that the modification somehow affected innate resistance (“Innate” is used here in opposition to the “acquired” Bt resistance) in ways such that slugs preferred Bt over Wt leaves. Third, the preference test suggests that the innate resistance in Bt relative to Wt plants may not be uniformly expressed throughout the whole plant and that leaf ontogeny interacts with the modification to affect resistance. This was manifested through an ontogenetic determined increase in leaf consumption that was more than four times higher in Bt compared to Wt leaves. Our result are of principal importance, as these indicate that genetic modifications can affect innate resistance and thus non-target herbivores in ways that may have commercial and/or environmental consequences. The finding of a modification–ontogeny interaction effect on innate resistance may be especially important in assessments of GM plants with a long lifespan such as trees.     

From interference to predation: type and effects of direct interspecific interactions of small mammals

Indirect exploitative competition, direct interference and predation are important interactions affecting species coexistence. These interaction types may overlap and vary with the season and life-history state of individuals. We studied effects of competition and potential nest predation by common shrews (Sorex araneus) on lactating bank voles (Myodes glareolus) in two seasons. The species coexist and may interact aggressively. Additionally, shrews can prey on nestling voles. We studied bank vole mothers’ spatial and temporal adaptations to shrew presence during summer and autumn. Further, we focused on fitness costs, e.g. decreased offspring survival, which bank voles may experience in the presence of shrews. In summer, interference with shrews decreased the voles’ home ranges and they spent more time outside the nest, but there were no effects on offspring survival. In autumn, we found decreased offspring survival in enclosures with shrews, potentially due to nest predation by shrews or by increased competition between species. Our results indicate a shift between interaction types depending on seasonal constraints. In summer, voles and shrews seem to interact mainly by interference, whereas resource competition and/or nest predation by shrews gain importance in autumn. Different food availability, changing environmental conditions and the energetic constraints in voles and shrews later in the year may be the reasons for the varying combinations of interaction types and their increasing effects on the inclusive fitness of bank voles. Our study provides evidence for the need of studies combining life history with behavioural measurements and seasonal constraints.     

Massive and distinctive effects of meadow voles on grassland vegetation

We ask whether vole herbivory in experimental grassland plots is sufficient to create an unpalatable community. In a six-year experiment, meadow voles (Microtus pennsylvanicus) reduced plant standing crop between 30% and 72%, well within the range of ungulate effects. Moreover, meadow voles reduced their available forage species by changing the plant community composition: four grass species and a legume upon which they foraged declined sharply in cover and/or number of individuals, five forbs avoided by voles increased, and two forbs neither declined nor increased with either measure. Reductions of diversity occurred when voles first defoliated the plots in 2000 but disappeared as plant species avoided by voles replaced vulnerable plants. Within six years, meadow voles created plant communities dominated by species that they did not eat.     

Genital morphology linked to social status in the bank vole (<Emphasis Type="Italic">Myodes glareolus</Emphasis>)

Since genital morphology can influence the outcome of post-copulatory sexual selection, differences in the genitalia of dominant and subordinate males could be a factor contributing to the fertilisation advantage of dominant males under sperm competition. Here we investigate for the first time if penile morphology differs according to male social status in a promiscuous mammal, the bank vole (Myodes glareolus). In this species, dominant males typically achieve higher reproductive success than subordinates in post-copulatory sexual selection, and male genital morphology is complex, including both a baculum (os penis) and penile spines. Our results show that despite no difference in body size associated with male social status, baculum width is significantly larger in dominant male bank voles than in subordinates. We also found evidence of positive allometry and a relatively high coefficient of phenotypic variation in the baculum width of male bank voles, consistent with an influence of sexual selection. By contrast, baculum length and three measures of penile spinosity did not differ according to male social status or show evidence of positive allometry. We conclude that dominant male bank voles may benefit from an enlarged baculum under sperm competition and/or cryptic female choice and that differences in penile morphology according to male social status might be important but as yet largely unexplored source of variation in male reproductive success.     

Behaviour and choice of refuge by voles under predation risk

Animals often show a strong antipredatory response when they are exposed to their most deadly predator. In northern vole populations, the least weasel, Mustela nivalis nivalis, is probably the most important predator of voles. Because of its small size and slender body shape, the least weasel is capable of hunting voles in their burrows. However, small voles can potentially escape weasel predation by selecting holes smaller than those weasels can enter. We studied the choice of nest holes and refuges by two species of voles under simulated predation risk. In a laboratory experiment, voles were provided with four nest boxes as refuges, with individually adjusted entrance sizes. When exposed to the odour of a weasel, voles did not choose the smallest opening; they rather seemed to trade full protection for easy and immediate access by choosing the nest box with an intermediate entrance size. When outside the nest at the time when a weasel entered the arena, voles avoided the refuges with the smallest holes. In addition to using refuges on ground level, voles climbed on top of the boxes as an escape reaction, as well as exhibiting a variety of behavioural responses, such as fast running, freezing and sneaking.Communicated by P.A. Bednekoff     

Ecological differences and coexistence in a guild of microparasites: Bartonella in wild rodents

The study of ecological differences among coexisting microparasites has been largely neglected, but it addresses important and unusual issues because there is no clear distinction in such cases between conventional (resource) and apparent competition. Here patterns in the population dynamics are examined for four species of Bartonella (bacterial parasites) coexisting in two wild rodent hosts, bank voles (Clethrionomys glareolus) and wood mice (Apodemus sylvaticus). Using generalized linear modeling and mixed effects models, we examine, for these four species, seasonal patterns and dependencies on host density (both direct and delayed) and, having accounted for these, any differences in prevalence between the two hosts. Whereas previous studies had failed to uncover species differences, here all four were different. Two, B. doshiae and B. taylorii, were more prevalent in wood mice, and one, B. birtlesii, was more prevalent in bank voles. B. birtlesii, B. grahamii, and B. taylorii peaked in prevalence in the fall, whereas B. doshiae peaked in spring. For B. birtlesii in bank voles, density dependence was direct, but for B. taylorii in wood mice density dependence was delayed. B. birtlesii prevalence in wood mice was related to bank vole density. The implications of these differences for species coexistence are discussed.     

Constitutive or induced defences - how does <Emphasis Type="Italic">Eucalyptus globulus</Emphasis> defend itself from larval feeding?

Summary. Following herbivory, induced responses involving plant secondary metabolites have been reported in a number of tree species. Although a wide range of plant secondary metabolites appear to operate as constitutive plant defences in trees belonging to the Eucalyptus genus, no induced responses have as yet been reported following foliar-chewing insect damage. We empirically tested whether branch defoliation (artificial and larval) of 2-year-old Eucalyptus globulus Labill. trees altered the abundance of specific plant secondary metabolites immediately (3 months after initial larval feeding) and 8 months after the cessation of larval feeding. Metabolites assayed, included essential oils, polyphenolic groups and foliar wax compounds and in all cases their abundance was not significantly altered by defoliation. However, the level of foliar tannins after 3 months of larval feeding did display a trend that suggested elevated levels as the result of defoliation, though this trend was not evident 8 months later, indicating that, if real, the response was a rapid and not a delayed induced response. The level of foliar tannins was also negatively correlated to both average larval survival and average percentage branch defoliation, suggesting that foliar tannins may operate as toxins and/or anti-feedants to M. privata larval feeding.     

Male dominance,female choice and male copulatory behavior in two species of voles (Microtus ochrogaster and Microtus montanus)

Summary Female preferences for dominant males in prairie and montane voles were analyzed in two different test situations. In the first, prairie vole females preferentially spent time in proximity to dominant versus subordinate males which were housed behind a wire mesh screen. In a two-male tether test prairie voles females both preferentially associated and mated with dominant males. Montane voles, on the other hand, showed no preference in either test situation. The baseline copulatory behavior of naive montane vole males which became dominant differed significantly from those which became subordinate; no such differences were evident in the baseline copulatory behavior of naive praire voles. One hour following dominance testing, there was no difference evident between dominant and subordinate males of either species but there was a general facilitation of male copulatory behavior in both species, the effect being of greater magnitude in montane voles.     

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.     

Response of Red-Backed Voles to Recent Patch Cutting in Subalpine Forest

We examined the response of southern red-backed voles ( Clethrionomys gapperi ) to patch cutting in a forested landscape in which 23% of the forest cover had been removed by timber harvest. We live trapped voles in and around 18 patchcuts in one watershed of southern Wyoming. Although we found a significant difference in capture rates between patchcut interior and forest habitats in 1 of 2 years, voles did not strongly avoid the interior of patchcuts. This result contrasts with results from most studies of voles in clearcut ecosystems, which report that red-backed voles are generally rare or absent from clearcuts. Capture rates were highest on both sides of the patchcut edge, which also contrasts with studies of voles at the edges of forest remnants. The use of patchcut interior and edge habitats could not be explained as a consequence of juvenile voles dispersing to those habitats or males moving through the habitat in search of mates. We suggest that, despite similar physiognomy in patchcut and clearcut sites, the differences in landscape structure in perforated versus fragmented landscapes lead to very different patterns of vole movement. Understanding the scales at which voles perceive landscapes as coarse- or fine-grained will be key to developing predictive models to aid managers in designing timber sales that maintain high vole populations. Our results emphasize the importance of the spatial pattern and scale of disturbance in determining the response of vertebrates to landscape change and the need for more refined investigations of the consequences of deforestation.     

Unintentional Wildlife Poisoning and Proposals for Sustainable Management of Rodents

In Europe, bromadiolone, an anticoagulant rodenticide authorized for plant protection, may be applied intensively in fields to control rodents. The high level of poisoning of wildlife that follows such treatments over large areas has been frequently reported. In France, bromadiolone has been used to control water voles (Arvicola terrestris) since the 1980s. Both regulation and practices of rodent control have evolved during the last 15 years to restrict the quantity of poisoned bait used by farmers. This has led to a drastic reduction of the number of cases of poisoned wildlife reported by the French surveillance network SAGIR. During the autumn and winter 2011, favorable weather conditions and high vole densities led to the staging of several hundreds of Red Kites (Milvus milvus) in the Puy‐de‐Dôme department (central France). At the same time, intensive treatments with bromadiolone were performed in this area. Although no misuse has been mentioned by the authorities following controls, 28 Red Kites and 16 Common Buzzards (Buteo buteo) were found dead during surveys in November and December 2011. For all these birds, poisoning by bromadiolone as the main cause of death was either confirmed or highly suspected. Other observations suggest a possible impact of bromadiolone on the breeding population of Red Kites in this area during the spring 2011. French regulation of vole control for plant protection is currently under revision, and we believe this event calls for more sustainable management of rodent outbreaks. Based on large‐scale experiments undertaken in eastern France, we propose that direct control of voles at low density (with trapping or limited chemical treatments) and mechanical destruction of vole tunnels, mole control, landscape management, and predator fostering be included in future regulation because such practices could help resolve conservation and agricultural issues. Envenenamiento No Intencional de Fauna Silvestre y Propuestas para un Manejo Sustentable de Roedores.     

Edge Effects and Isolation: Red-Backed Voles on Forest Remnants

Negative effects of habitat edge have been advanced as an important proximate cause of extinction, and a growing literature calls attention to the matrix surrounding habitat remnants as a critical factor determining population persistence. I examined spatial distribution of California red-backed voles ( Clethrionomys californicus ) on 13 forest remnants and five control sites in southwestern Oregon. The species was virtually isolated on remnants, making little use of the regenerating clearcuts surrounding the remnants. The effects of the clearcut also impinged on the remnants as edge effects: six times more voles were captured per trap in the interior of remnants than on the edge. Consequently, the density of voles per unit area on remnants increased with remnant size, despite the potential buildup of population density in small isolates due to limited emigration. I explored potential mechanisms of the negative edge effect on voles and found that the biomass of coarse woody debris, per se, did not explain the vole distribution because both number and volume of logs increased from the interior to the edge of remnants. However, the distribution of the vole's primary food item, hypogeous sporocarps of mycorrhizal fungi, did correspond to the vole edge effect     

Patterns of chemical defences in plants: an analysis of the vascular flora of Chile

Summary. The plant apparency hypothesis predicts that apparent plants invest in broadly effective defences such as tannins while unapparent plants invest in specific toxins such as alkaloids. The stress hypothesis states that plants invest in cheaper defences if they have evolved in habitats that impose abiotic limitations to plant fitness. We tested these hypotheses by determining the concentrations of alkaloids and tannins in a representative sample of the vascular plants of continental Chile (with exclusion of Pteridophyta, Cactaceae, and Poaceae) consisting of 396 species. In a subsample of 166 species which contained both alkaloids and tannins, we constructed the A/T index (A/T = [alkaloids]/ [tannins]). We discarded the presumed effect of phylogeny (as estimated by taxonomy) on the variation observed in the data because no correlation of A/T with taxonomic relationships among species either at family or genus levels was found in a nested ANOVA with genera nested in families. Concentration of alkaloids was negatively correlated with that of tannins. We compared the value of A/T among species differing in life form (herbs, shrubs or trees), herb longevity (annual or perennial), leaf-shedding manner of woody plants (deciduous or evergreen), latitudinal range, and level of water stress typical in their natural habitat. Unapparent plants (herbs, annual) exhibited higher mean A/T index than apparent plants (shrubs and trees, perennial). A/T did not correlate with latitudinal range. Mean A/T values decreased from deserts to deciduous forests. The comparisons were not always significant due to the inevitable unbalance of the data set which lowers the power of the statistical tests employed. The results suggest that chemical defences are indeed distributed in a non-random manner among plants, and that to a large extent the predictions derived from the apparency and stress hypotheses are sustained.     

Space use,longevity, and reproductive success in meadow voles

Summary We addressed the question of how reproductive success (RS) was limited in the shortlived but highly fecund meadow vole, Microtus pennsylvanicus. In so doing, we asked how differential space use patterns could affect longevity and hence RS in each sex. The sample comprised all voles achieving sexual competency over the course of a 40-week breeding season in a live-trapped population in Manomet, MA USA. Matrilineal families were determined using a radionuclide labelling technique; paternity was estimated using a maximum likelihood model. Individual RS was defined as the number of offspring successfully recruited into the trappable population per adult. We found that the variance in RS among female meadow voles was greater than the variance among males. In an attempt to explain this pattern, reproductively successful individuals were compared to reproductively unsuccessful individuals with regard to survivorship, maximum body weight achieved, and spatial mobility. The only difference between fathers and reproductively unsuccessful males was that fathers were heavier. In contrast, mothers differed from unsuccessful females in every measurement. Females lived longer than males, and mothers lived longer than either fathers or reproductively unsuccessful females. The observed differences in longevity may have been largely the result of differences in levels of mobility, assuming more mobile voles were more susceptible to predation. Mothers were significantly more site tenacious than were either males or unsuccessful females. These patterns explain the distribution of RS in our population if predation differentially affects male and female meadow voles. The meadow vole is the only non-polyandrous vertebrate reported to date in which the variance in RS among females exceeds the variance in RS among males.     

A field test of the Bruce effect in the monogamous prairie vole (Microtus ochrogaster)

The Bruce effect is a widely studied reproductive phenomenon in rodents in which exposure of pregnant females to unknown males causes termination of the current pregnancy. The Bruce effect has been reported from numerous studies in the laboratory, and one field study with the promiscuous gray-tailed vole, Microtus canicaudus, failed to support it. We conducted a field study with the monogamous prairie vole, M. ochrogaster, to determine if complete replacement of the male population every 10-14 days affected pregnancy and juvenile recruitment. The mean days to first parturition for control and treatment females were 36.8 and 44.4 days. Fifty-five percent of control females and 33% of treatment females conceived within the first 14 days of the study. All control females and 79% of treatment females successfully delivered at least one litter. These differences between treatment and control populations provide minimal support for the Bruce effect when compared with results from laboratory studies. Nulliparous females may have experienced some pregnancy disruption, but not parous females. Removal of mates, rather than exposure to strange males, may have contributed more to the lower reproductive success of treatment females than exposure to strange males. Treatment females, however, had lower juvenile recruitment than controls, which may have been due to infanticide from strange males. Our results are more similar to those of the field study of the gray-tailed vole than predicted, based on laboratory studies of prairie voles.     

Interactive effects of prey and p,p'-DDE on burrowing owl population dynamics.

We used population models to explore the effects of the organochlorine contaminant p,p'-DDE and fluctuations in vole availability on the population dynamics of Burrowing Owls (Athene cunicularia). Previous work indicated an interaction between low biomass of voles in the diet and moderate levels of p,p'-DDE in Burrowing Owl eggs that led to reproductive impairment. We constructed periodic and stochastic matrix models that incorporated three vole population states observed in the field: average, peak, and crash years. We modeled varying frequencies of vole crash years and a range of impairment of owl demographic rates in vole crash years. Vole availability had a greater impact on owl population growth rate than did reproductive impairment if vole populations peaked and crashed frequently. However, this difference disappeared as the frequency of vole crash years declined to once per decade. Fecundity, the demographic rate most affected by p,p'-DDE, had less impact on population growth rate than adult or juvenile survival. A life table response experiment of time-invariant matrices for average, peak, and crash vole conditions showed that low population growth under vole crash conditions was due to low adult and juvenile survival rates, whereas the extremely high population growth under vole peak conditions was due to increased fecundity. Our results suggest that even simple models can provide useful insights into complex ecological interactions. This is particularly valuable when temporal or spatial scales preclude manipulative experimental work in the field or laboratory.     

Effects of early management of Pinus massoniana plantation target trees on soil physicochemical properties and plant diversity北大核心CSCD

In order to provide a theoretical reference for the early management of target trees in the low mountain region of eastern Sichuan, and a theoretical basis for the sustainable management of masson pine (Pinus massoniana) plantations, we chose three different kinds of densities (100 target trees per hectare, 150 target trees per hectare, and 200 target trees per hectare) in a 33-year-old masson pine plantation in the low mountain region of eastern Sichuan. We investigated the change patterns of soil physicochemical properties and plant diversity during the early stage of the management of the target trees. The results showed that compared to the control (CK), the early stage of management of P. massoniana plantation target trees dramatically improved the soil physical properties, the pH value, contents of organic matter, and total phosphorus (P < 0.05); however, the available phosphorus and the available nitrogen varied slightly (P > 0.05). Compared to the control (CK), significant (P < 0.05) differences in the richness index were observed between shrubs and herbs (P < 0.05), and in the shrub layer, the dominant position was replaced by others. Simpson, Shannon, and Pielou indexes of shrubs showed no significant difference in the study (P > 0.05). On the contrary, Simpson, Shannon, and Pielou i ndexes of herbs were significantly lower than those for CK (P < 0.05). There were significant correlations between plant species diversity and soil physicochemical properties such as soil pH, contents of total phosphorus, and the available nitrogen. The early stage of management of P. massoniana plantation target trees significantly improved the plant diversity and soil physicochemical properties. Among all the three different treatments, the density of 150 target trees per hectare had the best effect on the soil physicochemical properties and plant diversity. © 2018 Science Press. All rights reserved.     

Endemic hantavirus infection impairs the winter survival of its rodent host

The influence of pathogens on host fitness is one of the key questions in infection ecology. Hantaviruses have coevolved with their hosts and are generally thought to have little or no effect on host survival or reproduction. We examined the effect of Puumala virus (PUUV) infection on the winter survival of bank voles (Myodes glareolus), the host of this virus. The data were collected by monitoring 22 islands over three consecutive winters (a total of 55 island populations) in an endemic area of central Finland. We show that PUUV infected bank voles had a significantly lower overwinter survival probability than antibody negative bank voles. Antibody negative female bank voles from low-density populations living on large islands had the highest survival. The results were similar at the population level as the spring population size and density were negatively correlated with PUUV prevalence in the autumn. Our results provide the first evidence for a significant effect of PUUV on host survival suggesting that hantaviruses, and endemic pathogens in general, deserve even more attention in studies of host population dynamics.