Smaller Microtus vole species competitively superior in the absence of predators

Interspecific competition is assumed to generate negative effects on coexisting species, possibly including slower population growth and lower survival. The field vole ( Microtus agrestis ) and the sibling vole ( M. rossiaemeridionalis ) are sympatric close relatives which compete for similar resources. Previous non-experimental studies suggest that the smaller sibling vole is a superior competitor, yet more vulnerable to predation than the larger field vole. We studied the effects of coexistence on population densities, reproductive parameters, and survival in these two species by means of experimentation in large, predator-free outdoor enclosures. While populations of both species reached higher densities in the absence of the other, field voles appeared to suffer more from interspecific competition than sibling voles. The proportion of young individuals in the population was higher in the sibling vole than in the field vole at the end of the experiment. The presence of a coexisting species reduced the survival of field voles. Sibling voles, on the other hand, appeared to suffer more from intraspecific competition than interspecific competition. On a population level, the sibling vole seems to be a superior competitor in the absence of predators due to better survival and possibly a higher reproductive capacity. However, predation probably has a profound influence on the interspecific dynamics of these two species indicating that in natural surroundings apparent competition (i.e. competition via shared predators) is stronger than direct competition.     

Juvenile prey induce antipredator behaviour in adult predators

It is generally assumed that the choice of oviposition sites in arthropods is affected by the presence of food for the offspring on the one hand and by predation risk on the other hand. But where should females oviposit when the food itself poses a predation risk for their offspring? Here, we address this question by studying the oviposition behaviour of the predatory mite Amblyseius swirskii in reaction to the presence of its counterattacking prey, the western flower thrips Frankliniella occidentalis. We offered the mites a choice between two potential oviposition sites, one with and one without food. We used two types of food: thrips larvae, which are predators of eggs of predatory mite but are consumed by older predator stages, and pollen, a food source that poses no risk to the predators. With pollen as food, the predators preferred ovipositing on the site with food. This might facilitate the foraging for food by the immature offspring that will emerge from the eggs. With thrips as food, female predators preferred ovipositing on the site without thrips. Predators that oviposited more on the site with thrips larvae killed more thrips larvae than females that oviposited on the site without food, but this did not result in higher oviposition. This suggests that the females killed thrips to protect their offspring. Our results show that predators display complex anti-predator behaviour in response to the presence of counter-attacking prey.     

Seasonal changes in the numerical responses of predators to cyclic vole populations

Theoretical models predict that a delayed density-dependent mortality factor with a time lag of ca 9 months is able to drive 3–5-yr population cycles of northern voles. We studied numerical responses of predators in western Finland during 1986–92, in an area with 3-yr population cycles of voles. Abundances of small mammals were monitored in several farmland areas (each 3 km²) by snap-trapping in April, June, August, and October (only in 1986–90), and the abundances of avian, mammalian, and reptilian predators by visual censuses during trapping occasions. The 3-yr cycle studied was a cycle of Microtus voles (field vole M. agrestis and sibling vole M. rossiaemeridionalis ) and their small-sized predators (small mustelids and vole-eating birds of prey). The numerical responses of both migratory avian predators and small mustelids to changes in vole densities were more alike than different. In late summer (August), the time lag in the numerical response of all main predators was short (0–4 months), whereas longer time lags prevailed from spring to early summer. The length of the time lag in spring appeared to be related to the length of the winter, which indicates that strong seasonality may create longer time lags to the numerical response of predators at northern latitudes than at more southern latitudes. Our results suggest that, from spring to early summer, predation by migratory avian predators may act in concordance with mustelid predation to produce the long time lag necessary to drive the 3-yr cycle of voles, whereas almost direct density-dependent predation by all major predators in late summer may dampen spatial variation in prey densities.     

Behaviour and resource use of two competing vole species under shared predation risk

Indirect interaction between two competing species via a shared predator may be an important determinant of population and community dynamics. We studied the effect of predation risk imposed by the least weasel Mustela nivalis nivalis on space use, foraging and activity of two competing vole species, the grey-sided vole Myodes rufocanus, and the bank vole Myodes glareolus. The experiment was conducted in a large indoor arena, consisting of microhabitat structures providing food, shelter, trees for refuge and separated areas with high and low predation risk. Voles were followed for 5 days: 2 days before, 1 day during and 2 days after the presence of weasel. Our results suggest an effect of weasel presence on the vole community. Voles of both species shifted their activity from risky to less risky areas, climbed trees more often and were less active. Seed consumption was not affected by weasel presence. The time spent in the risky and less risky area did not differ between species, but bank voles spent more time in trees than grey-sided voles. Males of both species were more exposed to predation risk than females, i.e. generally spent more time in the risky area. Proportion of time spent in the risky area, the use of area, trees and food stations were sex dependent. Activity and use of trees were species dependent. We found no evidence for despotic distribution between our two species, although bank voles seemed to be more affected by coexistence, since they lost weight during the experiment. Based on our results we conclude that predator response was largely similar between species, while the sex-specific responses dominated. Besides a stronger escape response in the bank vole, the strongest individual differences were sex specific, i.e. males were more prone to take risks in space use and activity.     

Saprospira species—Natural predators

SomeSaprospira spp. from fresh waters have been shown to be predatory on other aquatic bacteria, particularly gliding bacteria. Some can be grown as well axenically but others so far have been cultured only on the host bacteria for which they show a degree of specificity. A marine strain resemblingSaprospira grandis, although capable of growth axenically, is also a predator on a marineCytophaga sp.S. grandis ATCC 23119 failed to grow on the base media or media overgrown with host cells of otherSaprospira spp.     

Timing of vole hunting in aerial predators: RAPTOR GROUP RUG/RIJP*

In a short but intense field investigation, surface activity of common voles in a cropped lucerne field was assayed by live-trapping while, simultaneously, hunting activity and yield of three species of raptors were recorded by continuous observation. Pronounced short-term rhythms in trappability of the vole population ran parallel with fluctuations in yield per hour of flight-hunt of hen harriers and kestrels. These raptors, as well as Rough-legged buzzards, hunted more at times of increased vole surface activity; hen harriers saved c. 15 hours of flight-hunt per day by such temporal adjustment, corresponding to about 12% of their daily energy intake. Voles suffered a predation rate of an estimated 0 2% per day; under such heavy predation and temporal concentration of raptor hunting at times of increased vole activity, surface feeding in synchrony with the vole majority was associated with increased risk of predation.     

Pike predators induce morphological changes in young perch and roach

The morphological response of two 1 year-old prey species, perch Perca fluviatilis and roach Rutilus rutilus to the presence of predatory pike Esox lucius , was tested in a 6 week aquarium experiment. The growth of perch was higher than that of roach but there was no difference in growth of prey between predator and control treatments. The presence of pike caused morphological changes in both perch and roach. Perch showed an increase in body depth during the experiment whereas the morphological response of roach to pike was more related to a displacement of the dorsal and pelvic fins and width of the anal fin. Differences in morphological responses of perch and roach to pike can be interpreted by their overall differences in antipredator strategies.     

Population‐level manipulations of field vole densities induce subsequent changes in plant quality but no impacts on vole demography

Grazing‐induced changes in plant quality have been suggested to drive the negative delayed density dependence exhibited by many herbivore species, but little field evidence exists to support this hypothesis. We tested a key premise of the hypothesis that reciprocal feedback between vole grazing pressure and the induction of anti‐herbivore silicon defenses in grasses drives observed population cycles in a large‐scale field experiment in northern England. We repeatedly reduced population densities of field voles (Microtus agrestis) on replicated 1‐ha grassland plots at Kielder Forest, northern England, over a period of 1 year. Subsequently, we tested for the impact of past density on vole life history traits in spring, and whether these effects were driven by induced silicon defenses in the voles’ major over‐winter food, the grass Deschampsia caespitosa. After several months of density manipulation, leaf silicon concentrations diverged and averaged 22% lower on sites where vole density had been reduced, but this difference did not persist beyond the period of the density manipulations. There were no significant effects of our density manipulations on vole body mass, spring population growth rate, or mean date for the onset of spring reproduction the following year. These findings show that grazing by field voles does induce increased silicon defenses in grasses at a landscape scale. However, at the vole densities encountered, levels of plant damage appear to be below those needed to induce changes in silicon levels large and persistent enough to affect vole performance, confirming the threshold effects we have previously observed in laboratory‐based studies. Our findings do not support the plant quality hypothesis for observed vole population cycles in northern England, at least over the range of vole densities that now prevail here.     

Population changes of different predators during a water vole cycle in a central European mountainous habitat

Population changes in long-eared owls Asio otus , polecats Mustela putorius , red foxes Vulpes vulpes , stone martens Martes foina and badgers Meles meles were monitored during a water vole Arvicola terrestris scherman cycle in western Switzerland. Long-eared owls confirmed their status of highly mobile specialist predators in responding strongly and without time lag to water vole population changes. Even though polecats are considered generalists, they exhibited also a strong response to water vole fluctuations. Their numbers tracked water vole densities with a 1-yr time lag. Marked population changes were also recorded in red foxes and stone martens, but these changes were not related to water vole densities. Lastly, badgers did not show any significant population changes during the water vole cycle. We discuss the possible reasons for these differences and conclude that multi-factorial approach is clearly required to understand population processes in predator-prey systems.     

Multiple predator effects result in risk reduction for prey across multiple prey densities

Investigating how prey density influences a prey’s combined predation risk from multiple predator species is critical for understanding the widespread importance of multiple predator effects. We conducted experiments that crossed six treatments consisting of zero, one, or two predator species (hellgrammites, greenside darters, and creek chubs) with three treatments in which we varied the density of mayfly prey. None of the multiple predator effects in our system were independent, and instead, the presence of multiple predator species resulted in risk reduction for the prey across both multiple predator combinations and all three levels of prey density. Risk reduction is likely to have population-level consequences for the prey, resulting in larger prey populations than would be predicted if the effects of multiple predator species were independent. For one of the two multiple predator combinations, the magnitude of risk reduction marginally increased with prey density. As a result, models predicting the combined risk from multiple predator species in this system will sometimes need to account for prey density as a factor influencing per-capita prey death rates.     

Can prey behaviour induce spatially synchronic aggregation of solitary predators?

Spatio-temporal patterns of predator aggregations within their settlement areas (i.e. temporary settling zones used during dispersal or, more generally, foraging patches) were studied. By integrating the main behavioural rules of juveniles of Spanish imperial eagles Aquila adalberti during dispersal with the temporal availability of the eagles' main prey, we have developed several individual-based models under different simulation scenarios. The results suggest that the spatially synchrony of the aggregation patterns that we observed were derived from a combination of behavioural responses of the prey and the distances between available settlement areas. These results are discussed in the context of predator–prey relationships, optimal foraging and self-organizing processes.     

The role of the southern water vole Arvicola sapidus in the diet of predators: a review

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Population-level declines in Australian predators caused by an invasive species

The cane toad Bufo marinus has been migrating westward across northern Australia since its introduction as a biological control agent in 1935. It has been implicated in the widespread decline of many native frog-eating predators. To investigate the impacts of this invasive species on native predatory reptiles, annual surveys were conducted from 2001 to 2007 to document variation in the relative abundances of three varanid lizards ( Varanus mertensi, Varanus mitchelli and Varanus panoptes ) and one crocodile Crocodylus johnstoni species known to consume toads. In addition, the indirect effects of this variation on one agamid lizard Amphibolurus gilberti , a known prey item of V. panoptes , were also examined. Surveys were performed at two sites in northern Australia before and after the arrival of B. marinus . Significant declines in the relative abundances of all three species of varanid lizard were observed following toad arrival. Declines in the abundance of V. panoptes, V. mitchelli and V. mertensi at the two sites ranged 83–96, 71–97 and 87–93%, respectively. In contrast, A. gilberti increased by 23–26%; whereas there were no significant population-level declines in C. johnstoni despite observations of individual effects (i.e. several dead crocodiles with B. marinus in their stomachs). These findings suggest population-level changes in Australian lizards caused by an invasive species.     

Predator-induced synchrony in population oscillations of coexisting small mammal species

Comprehensive analyses of long-term (1977-2003) small-mammal abundance data from western Finland showed that populations of Microtus voles (field voles M. agrestis and sibling voles M. rossiaemeridionalis) voles, bank (Clethrionomys glareolus) and common shrews (Sorex araneus) fluctuated synchronously in 3 year population cycles. Time-series analyses indicated that interspecific synchrony is influenced strongly by density-dependent processes. Synchrony among Microtus and bank voles appeared additionally to be influenced by density-independent processes. To test whether interspecific synchronization through density-dependent processes is caused by predation, we experimentally reduced the densities of the main predators of small mammals in four large agricultural areas, and compared small mammal abundances in these to those in four control areas (2.5-3 km(2)) through a 3 year small-mammal population cycle. Predator reduction increased densities of the main prey species, Microtus voles, in all phases of the population cycle, while bank voles, the most important alternative prey of predators, responded positively only in the low and the increase phase. Manipulation also increased the autumn densities of water voles (Arvicola terrestris) in the increase phase of the cycle. No treatment effects were detected for common shrews or mice. Our results are in accordance with the alternative prey hypothesis, by which predators successively reduce the densities of both main and alternative prey species after the peak phase of small-mammal population cycles, thus inducing a synchronous low phase.     

Host plant specificity in several species of generalist mite predators

1. Species in the genus Neoseiulus are considered to be generalist predators, with some species used in biological control programmes against phytophagous mites and insects. 2. A general survey of Neoseiulus species in inland Australia indicated that different species are associated with particular tree species. This pattern of host plant use was investigated for four Neoseiulus species (N. buxeus, N. cappari, N. brigarinus, N. eremitus) by means of a sampling programme through time and across space. 3. Each species of Neoseiulus was collected entirely or mostly from one species of tree; little or no overlap was detected despite the tree species growing in well‐mixed stands. Host plant specificity thus appears to be strong in this genus. 4. Species in two other genera (Pholaseius and Australiseiulus), also considered to be predatory, showed a similar association with particular tree species. 5. The implications for the use of these predators in biological control are considerable. In particular, phytoseiid species with specific needs in terms of host plants may not be suitable for use as general purpose predators. Meeting the needs of phytoseiids through the modification of host plant attributes may be a step towards enhancing their efficacy as biological control agents.