Predation risk for herbivorous insects on tropical vegetation: A search for enemy-free space and time

Spatial and temporal variability in predation risk for herbivores on 13 rainforest species of Ficus (Moraceae) in Papua New Guinea was studied in order to assess whether predator-free refuges exist on their foliage and if so, whether herbivorous insects concentrate their activity in such refugia. Predation risk from invertebrate predators was measured as the disappearance rate of live termites set up as baits on the foliage. By far the most important predators were ants, accounting for 77% of attacks. No consistent differences in predation rate between Ficus species were found so that tree identity could not be used as an indicator of enemy-free space. Predation risk was highly variable among conspecific trees and also changed rapidly in time, over periods as short as 10 days. Such short-term and unpredictable predator-free refuges may be difficult for herbivores to find and exploit. Predation risk during the day was three times higher than during the night, but abundance of herbivores on the foliage was also higher during the day. Thus, night was confirmed as a relatively enemy-free time which, however, was not exploited by herbivores.     

Predation of codling moth Cydia pomonella L. by the Silvereye Zosterops lateralis (Latham)

Predation of cocooned larvae of codling moth Cydia pomonella by silvereyes Zosterops lateralis was studied in an apple orchard in Nelson, New Zealand. Apple logs with known larval densities were made available to the birds for known periods of time, either in cages or exposed in the apple orchard. The numbers of silvereyes and the natural predation of codling moth were recorded in the same orchard. Predation was density dependent. On caged logs with an initial high density of 32 larvae, 1.1 larvae were consumed per bird‐hour; in contrast, one larva was consumed per 34.5 bird‐hours at three larvae per log. A curvilinear relationship was demonstrated between larval density and the bird‐hours required for predation; this relationship was consistent with the known density dependence of silvereye predation of codling moth. A regression of the total annual winter bird predation of larvae in the orchard on bird numbers was significant. However, the density dependence of predation resulted in declining rates of predation over the winter as larval density declined; the first birds to arrive in the orchard benefitted from particularly high predation rates. As a consequence, fluctuations in bird numbers during the winter had only a secondary influence on predation rates. The numbers of silvereyes in the orchard showed no relationship to the density of the codling moth population present. This study confirmed the importance of silvereyes in the predation of codling moth and a functional, not numerical, rseponse of these birds to codling moth density.     

Influence of adult and egg predation on reproductive success of Epirrita autumnata (Lepidoptera: Geometridae)

We studied the effects of predation and oviposition activity on reproductive success of a late-season moth, Epirrita autumnata by exposing adult females and eggs to predation in their natural habitat in two successive years. Daily survival rates of adult females ranged from 0.4 to 0.8, average being 0.7. Most predation occurred during nights and was caused by harvestmen and other invertebrate predators. Avian predation did not have an effect on adult survival rates, most likely because of the lateness of E. autumnata flight season. Eggs were also preyed upon by invertebrate predators, although a notable proportion of egg mortality was attributable to causes other than predation. Daily survival rates of eggs were more than 0.99. Using modeling based on empirical data on eclosion of female adults, their oviposition behavior and survival rates of adults and eggs, the daily survival rates were translated into population level consequences. Adult predation was estimated to decrease reproductive success of non-outbreaking E. autumnata by 60–85 percent and egg mortality by 20–40 percent. Predation on adult lepidopterans is a mortality factor potentially as relevant as predation in any other life history stage and thus, should not be ignored in studies of population regulation.     

Impact of predation on early stages of the armoured catfish Hoplosternum thoracatum (Siluriformes-Callichthyidae) and implications for the syntopic occurrence with other related catfishes in a neotropical multi-predator swamp

This study investigated the role of predators in preventing competitive exclusion among three closely related armoured catfishes (Callichthys callichthys, Hoplosternum littorale and H. thoracatum) that occur synthopically in multi-predator freshwater swamps of Suriname, South America. The potential impact of predation on armoured catfish was determined by combining laboratory measurements of predation rates on five early developmental stages of the armoured catfish H. thoracatum for 24 aquatic predators with field studies of the density of the predators in the swamps. The contribution of a particular predator to the total predation pressure on its prey was determined to a large extent by the density of the predator in the swamp. Seemingly innocuous predators with low or moderate predation rates in the laboratory may be extremely important in the swamps due to their high abundance. Small-sized omnivorous fishes and aquatic invertebrates were major predators of early developmental stages of armoured catfish. Both qualitative and quantitative ontogenetic changes in the predation pressure on armoured catfish were observed. Major predation on eggs, larvae and juveniles of H. thoracatum resulted from a different set of predators in each developmental stage of the prey. In all developmental stages of H. thoracatum the predation pressure involved several predator species and not a single, dominant predator. The potential predation pressure of the 24 predators taken together and the number of predators that were able to prey on H. thoracatum decreased sharply with increasing age (size) of the prey. Even if egg (nest) predation is prevented by the guarding male, the potential impact of the 24 predators on the populations of armoured catfish is large. Predation may account for the high mortality of H. thoracatum observed in the swamps. The high predation pressure on callichthyid catfishes may help to explain the coexistence of three closely related and morphologically quite similar armoured catfishes in Surinamese swamps.     

Temporal variation in nest predation risk along habitat edges between grassland and secondary forest in Central Europe

Habitat fragmentation alters many ecological processes, including trophic cascades. For example, increased predation pressure along habitat edges has often been observed in fragmented landscapes. Here, we studied how nest predation risk varies along the transition zone between grassland and mixed forest in Central Europe. Using artificial nests, we tested the two mechanisms that are expected to underlie higher predation rates along edges: (1) the matrix effect model that supposes predator penetration from a habitat type with higher predator density to one with lower predator density and (2) the ecotonal effect model that assumes specific predator preferences for habitat edges. Although our results do not fully support either of these scenarios, our data show high temporal instability in nest predation along forest–grassland edges. Predation was higher in habitat interiors compared to edges during the first year, whereas the opposite pattern was observed during the subsequent year. In addition, dramatic between-year differences in the species composition of nest predators were observed. Therefore, we hypothesise that the effect of edges on nest predation is difficult to predict in landscapes with high predator diversity. In addition, our data indicate that a high abundance of wild boar considerably increases the risk of predation for ground-nesting birds.     

On the density-dependence of seed predation in<Emphasis Type="Italic"> Dipteryx micrantha</Emphasis>, a bat-dispersed rain forest tree

We studied the effect of seed density on seed predation by following the fate of bat-dispersed Dipteryx micrantha (Leguminosae) seeds deposited under bat feeding roosts. The study was conducted in Cocha Cashu biological station, Amazonian Peru, during the fruiting period of Dipteryx. Predation of Dipteryx seeds in the area is mainly by large to medium-sized rodents. Seed deposits beneath bat feeding roosts were monitored for a 13-week period in an 18-ha study area. A total of 210 seed deposits were found, and on average, seed predators encountered 22% of them during any one week. About one-third of the seed deposits escaped predation, and those deposits that had relatively few seeds were more likely to go unnoticed by rodents than were deposits with many seeds. The mean seed destruction rate was 8% per week; deposits with many seeds tended to lose a smaller proportion of their seeds to seed predators than did deposits with few seeds. Regression tests for the weekly data showed that, at the beginning of the observation period, seed predation was not density-dependent. Later, when the total seed crop beneath roosts was high, the number of seeds predated per deposit was positively density-dependent, while the proportion of seeds predated was negatively density-dependent, indicating predator satiation. Seed deposits that had been visited by seed predators once had a higher probability of being revisited the week after, especially if they contained many seeds when first encountered. This indicates that the foraging behavior of rodents may be affected by their remembering the location of seed-rich patches.     

Predator interference limits fly egg biological control by a guild of ground-active beetles

We examined whether predator interference could prevent effective conservation biological control of Delia spp. flies, important pests of cole crops, by an assemblage of carabid and staphylinid beetles. In laboratory feeding trials we found that the smaller (<1 cm) beetle species common at our site readily ate dipteran eggs, while the most common large carabid species, Pterostichus melanarius, rarely did. However, P. melanarius did eat several of the smaller beetle species. We conducted two field experiments where we manipulated immigration rates of the ground predator guild and then measured predation on fly eggs. Predation rates were consistently higher in cages where predators were added at ambient densities, compared to cages where ground predators were removed. However, in the second field experiment, when we quadrupled predator immigration rates neither beetle activity-density nor predation rate increased. High immigration rate plots had a higher proportion of P. melanarius in the predator community, compared to plots with beetles added at ambient densities, suggesting that P. melanarius was reducing activity-densities of the smaller beetles, perhaps through intraguild predation. Thus, tactics to improve the biological control of Delia spp. by conserving generalist predators, such as providing in- or extra-field refuges, could be thwarted if the primary predators of fly eggs, small carabids and staphylinids, are the targets of intraguild predation by also-conserved larger predators.     

Effects of microhabitat,time of day,and weather on predation of gypsy moth larvae

Summary Wire cages with different-sized meshes were placed on trunks and around leaves at different heights in oak trees and in forest litter. Gypsy moth, Lymantria dispar, instars II–V tethered with threads were placed in each cage (instars II–III only in leaf cages) as well as outside the cages. Predation of larvae decreased from near ground to mid-crown in trees and was highest in litter and very low on leaves. Predation in litter was not strongly related to cage type, suggesting that small, invertebrate predators were active there, but IV–V instars on trunks were mainly killed by relatively large predators, probably forest mice and shrews. Influences of the time of day and weather on predation were evaluated by observing tethered V–VI instars in litter and on tree trunks hourly. Ants, particularly carpenter ants (Camponotus ferrugineus) and Formica sp., and probably vertebrates, were conspicuous predators in the litter. Ants were most active at lower relative humidities, while other predators were apparently not influenced by humidity. No daily activity rhythms of predation were noted. Invertebrates appear to be important predators of larvae only in the litter whereas vertebrates, such as forest mice and shrews, also readily attack larvae on tree trunks.     

Detection of secondary predation by PCR analyses of the gut contents of invertebrate generalist predators

Predation by generalist predators is difficult to study in the field because of the complex effects of positive and negative interactions within and between predator species and guilds. Predation can be monitored by molecular means, through identification of prey DNA within predators. However, polymerase chain reaction (PCR) amplification of prey DNA from predators cannot discriminate between primary and secondary predation (hyperpredation), in which one predator feeds on another that has recently eaten the target prey. Here we quantify, for the first time, the potential error caused by detection of prey DNA following secondary predation, using an aphid-spider-carabid model. First, the aphid Sitobion avenae was fed to the spider Tenuiphantes tenuis and the carabid Pterostichus melanarius, and the postconsumption detection periods, for prey DNA within predators, were calculated. Aphids were then fed to spiders and the spiders to carabids. Aphid DNA was detected in the predators using primers that amplified 245- and 110-bp fragments of the mitochondrial cytochrome oxidase I gene. Fragment size and predator sex had no significant effect on detection periods. Secondary predation could be detected for up to 8 h, when carabids fed on spiders immediately after the latter had consumed aphids. Beetles tested positive up to 4 h after eating spiders that had digested their aphid prey for 4 h. Clearly, the extreme sensitivity of PCR makes detection of secondary predation more likely, and the only reliable answer in future may be to use PCR to identify, in parallel, instances of intraguild predation.     

Two common invertebrate predators show varying predation responses to different types of sentinel prey

Sentinel prey (an artificially manipulated patch of prey) are widely used to assess the level of predation provided by natural enemies in agricultural systems. Whilst a number of different methodologies are currently in use, little is known about how arthropod predators respond to artificially manipulated sentinel prey in comparison with predation on free‐living prey populations. We assessed how attack rates on immobilized (aphids stuck to cards) and artificial (plasticine lepidopteran larvae mimics) sentinel prey differed to predation on free‐moving live prey (aphids). Predation was assessed in response to density of the common invertebrate predators, a foliar‐active ladybird Harmonia axyridis (Coleoptera: Coccinellidae), and a ground‐active beetle Pterostichus madidus (Coleoptera: Carabidae). Significant increases in attack rates were found for the immobilized and artificial prey between the low and high predator density treatments. However, an increased predator density did not significantly reduce numbers of free‐living live aphids included in the mesocosms in addition to the alternate prey. We also found no signs of predation on the artificial prey by the predator H. axyridis. These findings suggest that if our assessment of predation had been based solely on the foliar artificial prey, then no increase in predation would have been found in response to increased predator density. Our results demonstrate that predators differentially respond to sentinel prey items which could affect the level of predation recorded where target pest species are not being used.     

Can polyphagous predators control the bird cherry-oat aphid (Rhopalosiphum padi) in spring cereals?: A simulation study

Predation by two carabid species,Bembidion lampros (Herbst.) andPterostichus cupreus L., on the cereal aphid,Rhopalsiphum padi L., was described using a discrete event simulation model. The results were incorporated into an aphid population growth model. The aphid model was run using actual temperatures,R. padi emigration rates, field densities and distributions from three years with distinctly different aphid population levels. Predation was incorporated at different time periods. Predation during the arrival of emigrants and establishment of aphid populations had approximately the same effect as predation over the entire aphid period, whereas predation during the 20 days prior to peak had very little effect on resulting peak populations. We conclude that a) yearly levels ofR. padi infestation are mainly determined by the volume and intensity of emigrants; b) peak levels are sensitive to changes in predator density; and c) in some cases an outbreak ofR. padi may be prevented by the action of spring active polyphagous predators alone.     

Experimental assessment of predation risk for juvenile green sturgeon,Acipenser medirostris,by two predatory fishes

Predation is a common cause of early life stage mortality in fishes, with reduced risk as individuals grow and become too large to be consumed by gape-limited predatory fishes. Large-bodied species, such as sturgeon, may reach this size-refuge within the first year. However, there is limited understanding of what this size threshold is despite the value of this information for conservation management. We conducted laboratory-based predation experiments on juvenile green sturgeon, Acipenser medirostris, to estimate vulnerability to predation during outmigration from their natal reaches in California to the Pacific Ocean. Two highly abundant and non-native predatory fish species (largemouth bass, Micropterus salmoides, and striped bass, Morone saxatilis) were captured in the wild to be tested with developing juvenile green sturgeon from the UC Davis Green Sturgeon Broodstock Program. Experimental tanks, each containing five predators, received thirty prey for 24-hr exposures. Between sturgeon prey trials, predators were exposed to alternative prey species to confirm predators were exhibiting normal feeding behaviors. In addition to green sturgeon mortality data, trials were video recorded and predatory behaviors were quantified. Overall, these predator species displayed much lower rates of predation on juvenile green sturgeon than alternate prey. Predation decreased with green sturgeon size, and predation risk diminished to zero once sturgeon reached a length threshold of roughly 20–22 cm total length, or between 38% and 58% of predator total length. Behavioral analyses showed low motivation to feed on green sturgeon, with both predators attempting predation less frequently as sturgeon grew. Results of this study imply that optimizing growth rates for larval and juvenile sturgeon would shorten the time in which they are vulnerable to predation. Future experiments should assess predation risk of juvenile green sturgeon by additional predator species common to the Sacramento-San Joaquin watershed.     

Predator‐mediated interactions between preferred,alternative and incidental prey in the arctic tundra

Apparent competition between prey is hypothesized to occur more frequently in environments with low densities of preferred prey, where predators are forced to forage for multiple prey items. In the arctic tundra, numerical and functional responses of predators to preferred prey (lemmings) affect the predation pressure on alternative prey (goose eggs) and predators aggregate in areas of high alternative prey density. Therefore, we hypothesized that predation risk on incidental prey (shorebird eggs) would increase in patches of high goose nest density when lemmings were scarce. To test this hypothesis, we measured predation risk on artificial shorebird nests in quadrats varying in goose nest density on Bylot Island (Nunavut, Canada) across three summers with variable lemming abundance. Predation risk on artificial shorebird nests was positively related to goose nest density, and this relationship was strongest at low lemming abundance when predation risk increased by 600% as goose nest density increased from 0 to 12 nests ha^?1. Camera monitoring showed that activity of arctic foxes, the most important predator, increased with goose nest density. Our data support our incidental prey hypothesis; when preferred prey decrease in abundance, predator mediated apparent competition via aggregative response occurs between the alternative and incidental prey items.     

Predator effects on prey population dynamics in open systems

Animal population dynamics in open systems are affected not only by agents of mortality and the influence of species interactions on behavior and life histories, but also by dispersal and recruitment. We used an extensive data set to compare natural loss rates of two mayfly species that co-occur in high-elevation streams varying in predation risk, and experience different abiotic conditions during larval development. Our goals were to generate hypotheses relating predation to variation in prey population dynamics and to evaluate alternative mechanisms to explain such variation. While neither loss rates nor abundance of the species that develops during snowmelt (Baetis bicaudatus) varied systematically with fish, loss rates of the species that develops during baseflow (Baetis B) were higher in streams containing brook trout than streams without fish; and surprisingly, larvae of this species were most abundant in trout streams. This counter-intuitive pattern could not be explained by a trophic cascade, because densities of intermediate predators (stoneflies) did not differ between fish and fishless streams and predation by trout on stoneflies was negligible. A statistical model estimated that higher recruitment and accelerated development enables Baetis B to maintain larger populations in trout streams despite higher mortality from predation. Experimental estimates suggested that predation by trout potentially accounts for natural losses of Baetis B, but not Baetis bicaudatus. Predation by stoneflies on Baetis is negligible in fish streams, but could make an important contribution to observed losses of both species in fishless streams. Non-predatory sources of loss were higher for B. bicaudatus in trout streams, and for Baetis B in fishless streams. We conclude that predation alone cannot explain variation in population dynamics of either species; and the relative importance of predation is species- and environment-specific compared to non-predatory losses, such as other agents of mortality and non-consumptive effects of predators. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Experimental Evidence that Sexual Displays are Costly for Nest Survival

Nest predation is one of the most significant limitations for successful breeding of tropical passerines. Thus, parental strategies may include choosing appropriate nest sites and behaving in ways that minimize predation. Habitat characteristics that may influence nest success include degree of nest concealment, proximity to habitat edge, plant architecture as well as several others cited in the literature. However, few studies have examined display behavior as a factor that could also influence nest survival. We experimentally tested whether sexual motor displays served as a cue for visually oriented predators to locate artificial nests in a population of blue‐black grassquits Volatinia jacarina, a Neotropical passerine that exhibits a complex sexual display and is subjected to elevated rates of nest predation. We also evaluated the effect of nest substrate on survival. Predation rate was higher for nests within territories of displaying males relative to areas without displaying males and for nests placed in shrubs relative to grasses. Predation increased sharply in the third experimental replicate, at the end of the breeding season, which suggests that predators may develop a search image for nests or may become more abundant during specific periods of the season. Avian predators appear to be the most important nest predators. Results suggest that there may be a trade‐off between the increase in fitness derived from sexual displays of males to attract potential mates and the decrease owing to predation of active nests within their territories.     

Snake predation on North American bird nests: culprits,patterns and future directions

Predation is the leading cause of nest failure for most birds. Thus, for ornithologists interested in the causes and consequences of variation in nest success, knowing the identity and understanding the behavior of dominant nest predators is likely to be important. Video documentation of nests has shown that snakes are frequent predators. Here we reviewed 53 North American studies that used nest cameras and used these data to identify broad patterns in snake predation. Snakes accounted for 26% (range: 0–90%) of recorded predation events, with values exceeding 35% in a third of studies. Snakes were more frequent nest predators at lower latitudes and less frequent in forested habitat relative to other nest predators. Although 12 species of snakes have been identified as nest predators, ratsnakes Elaphe obsoleta, corn snakes E. guttata and fox snakes E. vulpina were the most frequent, accounting for > 70% of all recorded nest predation events by snakes and have been documented preying on nests in 30–65% of studies conducted within their geographic ranges. Endotherm‐specialist snakes (Elaphe and Pituophis genera) were more likely to depredate nests in forests and the canopy relative to other snakes, due to their affinity for edge habitat. Predation by only ratsnakes and corn snakes was predominantly nocturnal and only ratsnakes were more likely to prey on nests during the nestling stage. Snakes were not identified to species in over 30% of predation events, underlining the need for more complete reporting of results. A review of research to date suggests the best approach to investigating factors that bring snakes and nests into contact involves combining nesting studies with radio tracking of locally important snake nest predators.     

Potential cotton aphid,Aphis gossypii,population suppression by arthropod predators in upland cotton

The cotton aphid, Aphis gossypii Glover, predation rate of convergent lady beetle, Hippodamia convergens Guerin‐Meneville, was determined by assigning a single predator randomly to each of four prey density treatments in the laboratory. Prey densities included 25, 50, 100, and 200 aphids per Petri dish arena. Predation response was recorded at 1, 4, 8, 16, 24, and 48 h after assigning predators to their prey treatments. Rate of consumption increased through time, with all 25 aphids consumed during the first 4 h of the experiment. At the highest density, adult lady beetle consumed on average 49, 99, 131, 163, 183, and 200 aphids within 1, 4, 8, 16, 24 and 48 h, respectively. Predators showed a curvilinear feeding response in relation to total available time, indicating that convergent lady beetles have the potential to suppress larger populations of aphids through continuous feeding by regulating their predation efficiency during feeding. The analysis of age‐specific mortality in absence of prey revealed that lady beetles could survive for an extended period of time (more than 2 weeks) without prey. The ability of a predator to survive without prey delays or prevents the rebound of pest populations that is a significant factor in natural biological control. A two‐year field sampling of 10 cotton arthropod predator species showed that spiders (27%) were the most dominant foliage dwelling predators in the Texas High Plains cotton followed by convergent lady beetles (23.5%), hooded beetles (13.5%), minute pirate bugs (11%), green lacewings (9.5%), bigeyed bugs (7.5%), scymnus beetles (3%), soft‐winged flower beetles (2%), damsel bugs (1.5%), and assassin bugs (1.5%). A field cage study showed that one H. convergens adult per plant released at prey density of one aphid per leaf kept the aphid population below economic threshold for the entire growing season.     

The effect of sand and light on predation of juvenile plaice (Pleuronectes platessa) by fishes and crustaceans

Rates of predation on 0-group plaice, Pleuronectes platessa . in aquaria were compared under four different combinations of conditions to test the hypothesis that the presence of sand in which they may bury affords a refuge from predators. The effect of light and darkness on predation rate was also examined, Two crustaceans, the shrimp, Crangon crangon , and the portunid crab, Liocarcinus holatus , and two fishes, cod, Gadus morhua , and pollack, Pollachius pollachius , were used as predators. Predaton rates were significantly higher in the dark for all predators except pollack. Predation rates in the absence of sand were signifcantly greater only for pollack. The results suggest that predation rates on plaice during their juvenile nursery stage on sandy beaches will be significantly greater during darkness than during the day. Burying in sand appears to provide only a partial refuge from predation, perhaps because natural predators have evolved effective methods of foraging for buried prey.     

The relationship between predator density, community composition, and field predation of Colorado potato beetle eggs

Conservation biological control programs seek to increase natural enemy densities through the adoption of more benign farming practices, under the assumption that higher predator densities will lead to more effective pest suppression. However, predator–predator interference may lead to diminishing returns in improved pest control as predator densities increase. We examined the relationship between predator density and predation rates on Colorado potato beetle eggs in production potato fields. These potato fields naturally spanned a 10-fold range in predator density, due to differences in management practices. Periodically through the growing season we simultaneously measured predator densities and subjected sentinel eggs masses to predation, allowing us to correlate predator density and egg predation for each field on each sample date. Egg predation rates were significantly positively correlated with total predator densities, a correlation that was not improved when predator densities were scaled to reflect differences in feeding rates on potato beetle eggs of the constituent predator taxa. There was no correlation between per-capita egg predation rates and predator density, and so no evidence that predator interference increased with increasing predator density. We divided predators into six dominant taxa—dwarf spiders, crab spiders, minute pirate bugs, big-eyed bugs, damsel bugs, and Lygus bugs (together constituting 93% of all predators collected), and a seventh group, “other predators” that included all other, less common, taxa—and examined correlations between all predator combinations and egg predation rates. The highest correlation was between combined densities of the six most common predator taxa, excluding only the “other predators” grouping. This suggests that predators may be largely equivalent in their impact on Colorado potato beetle eggs, and that field scouts might be able to ignore uncommon predator taxa when sampling for natural enemies.     

Can predation cause the 10-year hare cycle?

Summary We relate causes of mortality of snowshoe hares to density of hares over an 8-year period that included a peak in numbers. We then use simulation modeling to examine whether these density-dependent relationships could produce changes in hare density similar to those observed in our study are in Yukon, Canada.Predation during winter was the largest source of mortality for snowshoe hares at Kluane, Yukon during 1978–84. There was a one-year lag in the response of winter predation mortality rate to hare density. There was a two-year lag in the response of winter mortality not caused by predators to hare density.A simple simulation model with density-dependent predation produced 8–11 year cycles only within a narrow range of parameters that are inconsistent with data from the Kluane region. However, a simulation model that predicted winter mortality rates using a delayed density-dependent numerical response and a Type II functional response by predators, produced 8–11 year cycles within the range of parameter values measured in our study. Yet another simulation model that predicted both summer and winter mortality rates using a delayed density-dependent numerical response and a Type II functional response by predators, did not produce 8–11 year cycles within the range of parameter values measured in our study. Lack of data on juvenile mortality may be one reason for this result.