Grouse dynamics and harvesting in Kainuu,northeastern Finland

We study the dynamics of the capercaillie, black grouse, hazel grouse and willow grouse in Kainuu game management district in northeastern Finland in the years 1989–2004. It appears that the 6–7 year periodicity that prevailed in this region from 1960s up to 1980s has now vanished in all species. The grouse data are modelled using a linear autoregressive model with lag terms for population dynamics including grouse harvest as annual bag and an index of winter severity (winter‐time area of Baltic Sea ice cover). We use the Akaike information criterion for selecting the best model for each species; first order lag is forced to the models. It turns out that a term is needed for harvesting (with a negative coefficient) in models for all species. For the capercaillie and the hazel grouse second order lag was included, for the black grouse and the willow grouse first order lag suffices. The willow grouse is the only species where the index of winter strength (with a negative coefficient) is needed in the model.     

How reliable are harvesting data for analyses of spatio‐temporal population dynamics?

Harvest data are commonly used as proxy for count data, especially in studies of long‐term temporal and spatial patterns of population fluctuations. However, usually the concurrence of the conclusions based on different types of data is impossible to verify due to the lack of count data. Here, we use annual (1964–2004) harvest and population census data for capercaillie, black grouse and hazel grouse from 14 game management districts covering Finland, and demonstrate some mismatch in the information that these data sets provide. Overall, linear regressions of annual harvest against population count give a reasonable fit, but the slopes are less than 1 in every species. Harvest bags have been proportionally larger in north and eastern Finland than in southwestern Finland, with marked species‐specific differences. Considering population variation, the CV% in the census data (30–50%) is consistently smaller than it is in the harvest data (60–70%). Most importantly, conclusions on the spatio‐temporal patterns of the population dynamics are different if based on harvest rather than count data. In capercaillie, synchrony decreases faster with distance according to the harvest data, while in black grouse and hazel grouse the census data show the steeper decline. In addition, the autocorrelation coefficients in the census time series are higher in capercaillie and black grouse than in harvest data, but in hazel grouse the opposite is true. Finally, the parameter estimates for a second order autoregressive model using different data sets differ, and these differences are species‐specific. Despite the fact that annual harvest is a positive and linear function of annual grouse population density, the pattern of population dynamics derived from the bag data is different from that shown by the census data. This result urges caution in using wildlife bag data as reliable indices of population dynamics. deceased August 2008.     

Goshawk predation during winter, spring and summer in a boreal forest area of central Sweden

Predation by goshawks was studied in a central Swedish boreal forest area. Data were collected in winter (January–February) 1977-81 by tracking radio-tagged goshawks, and in the breeding season (April–July) by collecting prey remains at the nest. In the breeding season birds dominated the prey, amounting to 86% of prey number and 91% of prey biomass. Wood pigeon Columba palumbus , black grouse Tetrao tetrix , hooded crow Corvus corone cornix and jay Garrulus glandarius accounted for more than 50% of the prey animals, whereas capercaillie Tetrao urogallus and black grouse accounted for more than 50% of prey biomass. There was no functional response to black grouse density fluctuations. Every year goshawks killed significantly more females than males of both capercaillie and black grouse, due to high vulnerability of the grouse hens while laying and incubating. It was estimated that during spring and early summer goshawk predation removed 25% of the female, and 14% of the male black grouse population. In winter squirrel was the dominating prey, both in terms of number (79%) and weight (56%). The proportion of squirrel in the diet was equally high both in winters of low and high squirrel density. The high proportion of squirrel in the winterdiet, as compared to the breeding season, is believed to be due to squirrels having to accept an increased predation risk in winter, in order to feed efficiently enough.     

Nest loss in capercaillie and black grouse in relation to the small rodent cycle in southeast Norway

Summary The relationship between nest loss in boreal forest grouse and the fluctuations in small rodents was studied at Varaldskogen in southeast Norway during 1979–1986, covering two complete rodent cycles. Nest loss in capercaillie (N=174) and black grouse (N=81) was calculated according to Mayfield (1975) based on nests from radio-equipped hens (N=77) and nests found by other methods (N=178). Small rodent density was measured by snap trapping during spring and autumn. Losses varied as predicted by the classical alternative prey hypothesis (Hagen 1952 and Lack 1954, as elaborated by Angelstam et al. 1984): high losses during rodent crash years (85.5% capercaillie, 51% black grouse), and smaller losses during peak years (54.5% capercaillie, 32.5% black grouse). Losses were inversely related to autumn abundance of rodents in capercaillie (P<0.05), but the correlation was not significant for black grouse (0.10<P<0.20). In capercaillie, the only species with an adequate sample for analysis, no relationship was detected between spring density of rodents and nest loss. Losses during the prepeak years were nearly as high as during crash years, a result inconsistent with the model. We conclude that the numerical response of predators to their cyclic main prey (i.e. small rodents) probably play a main role during the low phase and prepeak year, whereas the dietary shift is most important during the peak and crash year of the cycle.     

Guardian or threat: does golden eagle predation risk have cascading effects on forest grouse?

Previous studies on intraguild predation have mainly focused on within-class assemblages, even though avian top predators may also influence mammalian mesopredator prey. By using nation-wide long-term data from Finland, northern Europe, we examined the impacts of golden eagles (Aquila chrysaetos) together with red foxes (Vulpes vulpes) and pine martens (Martes martes) on forest-dwelling herbivores, black grouse (Tetrao tetrix) and hazel grouse (Tetrastes bonasia). We hypothesized that eagles may alleviate the overall predation pressure on grouse by imposing intraguild predation risk on mesopredators. The predation impact of eagle was modelled using eagle density estimates and distance to eagle nest. Wildlife triangle counts were used as predation impact proxies of mammalian mesopredators and as measures of response in grouse. Our results show that eagle density correlated negatively with black grouse abundance indices while being positively associated with the proportion of juveniles in both grouse species, irrespective of the abundance of mesopredators. Yet, foxes and martens alone had a negative effect on the abundance indices and the proportion of young in the two grouse species. This suggests that the possible cascading effects of eagles are not mediated by decreased mesopredator numbers, but instead by fear effects. Alternatively, they may be mediated by other species than fox or marten studied here. In conclusion, we found support for the hypothesis that eagles provide protection for juvenile black and hazel grouse, whereas they are a threat for adult grouse. This important information helps us to better understand the role of avian top predators in terrestrial ecosystems.     

Territory occupancy rate of goshawk and gyrfalcon: no evidence of delayed numerical response to grouse numbers

Two recent studies on territory occupancy rates of goshawk Accipiter gentilis and gyrfalcon Falco rusticolus report a 2–3-year-delayed numerical response to grouse numbers, which is a requirement for a hypothesis of predator-generated grouse cycles. The time lags were assumed to reflect the average age of sexual maturity in the raptor species. In southern Norway, however, subadult (two-year-old) goshawk hens occupied only 18–25% of territories where occupancy was not recorded in the preceding year, and there was no significant relationship between the proportion of subadults among recruits and grouse indices two years earlier. We argue that territory occupancy rates are not appropriate indices of total raptor population levels, but rather reflect the proportion of territorial pairs that attempt to nest. Because this depends on the body condition of the hens, fluctuations in other important winter resident prey species (most important for the goshawk) and winter weather (most important for the gyrfalcon) should also be addressed. During 1988–2006, the annual proportion of goshawk territories with recorded nesting attempts in southern Norway was most closely related to the preceding autumn’s population indices of black grouse Tetrao tetrix and mountain hare Lepus timidus, whereas the annual proportion of gyrfalcon territories with observations of falcons or with confirmed breeding attempts in central Norway were best explained by population indices of willow grouse Lagopus lagopus and ptarmigan L. mutus from the previous autumn, and by December temperatures. Hence, our studies do not support the predation hypothesis for grouse cycles.     

Diet of the golden eagle Aquila chrysaetos during the breeding season in Sweden

During a five-year period, 1975–1979, a total of 2881 prey individuals of 65 prey species were collected at 162 golden eagle nests from northern Sweden and from the island of Gotland. In northern Sweden birds are taken in higher numbers than mammals but calculated as weight the two categories are of equal importance, The main prey during the breeding season are capercaillie, black grouse, willow grouse, ptarmigan, mountain hare and reindeer fawns which together form 91% of the total food biomass. The capercaillie and the black grouse are taken more in the southern part of the coniferous region than in the northern. In contrast, in northern areas, reindeer fawns are more preyed upon than in the South, Ptarmigan and willow grouse are the most commonly captured prey species in mountain areas. The total number of reindeer fawns taken (dead and/or alive) by the Swedish golden eagle population during one summer is estimated at 600 individuals. On Gotland the golden eagles take mammals more often than in its northern distribution area. Rabbit and hedgehog arc the most important species.     

Impact of climate change and prey abundance on nesting success of a top predator, the goshawk

Contemporary research has documented a large number of shifts in spring phenology and changes in distribution range although the average spring temperatures have increased by only 0.3–0.6 °C over the past 100 years. Generally, earlier breeding birds have larger clutch sizes, and the advancing spring could thus potentially increase breeding success. Shifts in spring phenology can, however, be crucial for bird reproduction, and mistiming the breeding event may even have negative consequences for population development. Our aim was to explore how weather and prey abundance relates to the breeding performance of a north European top predator, the northern goshawk Accipiter gentilis. Our nationwide dataset from Finland, spanning the period 1989–2004, shows that ambient weather has a greater impact on the timing and success of breeding than the density of grouse Tetraonidae, the main prey of goshawks. Higher early spring temperatures were associated with advancing hatching date of goshawks. Correspondingly, grouse density and temperature during laying and brooding were positively associated with brood size, while precipitation showed a negative connection. Applying our models to a future scenario of climate warming, combined with a 50 % reduction in grouse density, suggests that average breeding dates will advance only 2.5 days and average breeding success would remain the same. Notably, breeding success was not spatially equal throughout Finland, as northern and eastern populations suffered most from declining grouse densities. The observed pattern is thus the opposite to what is expected from a population situated at the northern edge of its distribution range, and thus may help to understand why populations may not increase at the northern edge of their thermal distribution due to climate change.     

Sex and age influence intestinal parasite burden in three boreal grouse species

Parasite infections are often more common in male vertebrates than in females. Sexual selection leading to dimorphism can cause sexual differences in immune defence, behaviour or body size. Possible proximate explanations for male bias in parasitism are the immunosuppressive effect of male sex hormones and the large body size of males which increases the likelihood of being parasitized. To evaluate these hypotheses, we studied the prevalence and abundance of intestinal helminth parasites in three boreal grouse species, the capercaillie Tetrao urogallus , the black grouse Tetrao tetrix , and the hazel grouse Bonasa bonasia . The first two are sexually dimorphic polygynous species while the latter species is sexually monomorphic and monogamous. We found a male-bias in the prevalence and abundance of the nematode Ascaridia compar . The bias was most pronounced in the polygynous black grouse and capercaillie. In the monogamous hazel grouse, there was a slight male-bias in occurrence of ascarids, but no bias in abundance. In juvenile grouse, the male-bias was larger than in adult grouse. No sexual bias was found in regard to the cestodes ( Skrjabinia cesticillus , Paroniella urogalli and Hymenolepis spp.). However, age was a factor in cestode prevalence: juvenile grouse were more commonly infected than adults. Differences in growth rates and body size are potential factors that may lead to male-biased parasitism in these grouse species, and their impact requires further studies.     

Climate change in Norway: warm summers limit grouse reproduction

Grouse and vole numbers may peak after peaks in the seed crop of bilberry (Vaccinium myrtillus) because of reduced levels of feeding deterrents in bilberry plants. We predicted that grouse reproduction depends also on summer (June–September) temperatures in the 2 previous years, because bilberry plants will be less exhausted after a high seed crop in or after warm summers, and thus rebuild their chemical defence more quickly. After berry peak years, population indices of capercaillie (Tetrao urogallus) and bank vole (Myodes glareolus) in southern Norway were negatively related to summer temperatures in the previous year or previous 2 years. Willow grouse (Lagopus lagopus) chick production in five areas in Norway was negatively related to summer temperatures in the 2 previous years when controlling for vole density. A similar pattern was found for the bilberry-feeding moth (Eulithis populata), an important prey for grouse chicks. In eastern Norway, autumn densities of capercaillie and black grouse (Tetrao tetrix) were more likely to peak in vole peak years at high altitudes, where summer temperatures are low. We conclude that high summer temperatures may limit grouse reproduction through the effect on bilberry plants and that a warm climate thus adversely affects population levels of grouse.     

Vulnerability of black grouse hens to goshawk predation: result of food supply or predation facilitation?

The plant cycle hypothesis says that poor-quality food affects both herbivorous voles (Microtinae spp.) and grouse (Tetraonidae spp.) in vole decline years, leading to increased foraging effort in female grouse and thus a higher risk of predation by the goshawk Accipiter gentilis. Poor-quality food (mainly the bilberry Vaccinium myrtillus) for these herbivores is induced by seed masting failure in the previous year, when the bilberry is able to allocate resources for chemical defence (the mast depression hypothesis; MDH). The predation facilitation hypothesis (PFH) in turn states that increased searching activity of vole-eating predators during or after the decline year of voles disturbs incubating and brooding grouse females. The behaviours used by grouse to avoid these terrestrial predators make them more vulnerable to predation by goshawks. We tested the main predictions of the MDH and PFH by collecting long-term (21-year) data from black grouse Tetrao tetrix hens and cocks killed by breeding goshawks supplemented with indices of bilberry crop, vole abundance and small carnivores in the vicinity of Oulu, northern Finland. We did not find obvious support for the prediction of the MDH that there is a negative correlation of bilberry crop in year t with vole abundance and with predation index of black grouse hens in year t + 1. We did find obvious support for the prediction of the PFH that there is a positive correlation between predator abundance and predation index of grouse hens, because the stoat Mustela erminea abundance index was positively related to the predation index of black grouse hens. We suggest that changes in vulnerability of grouse hens may mainly be caused by the guild of vole-eating predators, who shift to alternative prey in the decline phase of the vole cycle, and thus chase grouse hens and chicks to the talons of goshawks and other avian predators.     

Spatial relationships and mechanisms of coexistence between dominant and subordinate top predators

Most forest ecosystems contain a diverse community of top‐level predators. How these predator species interact, and how their interactions influence their spatial distribution is still poorly understood. Here we studied interactions among top predators in a guild of diurnal forest raptors in order to test the hypothesis that predation among competing predators (intraguild predation) significantly affects the spatial distribution of predator species, causing subordinate species to nest farther away from the dominant ones. The study analyzed a guild in southwestern Europe comprising three raptor species. For 8 years we studied the spatial distribution of used nests, breeding phenology, intraguild predation, territory occupancy, and nest‐builder species and subsequent nest‐user species. The subordinate species (sparrowhawk Accipiter nisus) nested farther away from the dominant species (goshawk A. gentilis), which preyed on sparrowhawks but not on buzzards Buteo buteo, and closer to buzzards, with which sparrowhawks do not share many common prey. This presumably reflects an effort to seek protection from goshawks. This potential positive effect of buzzards on sparrowhawks may be reciprocal, because buzzards benefit from old sparrowhawk nests, which buzzards used as a base for their nests, and from used sparrowhawk nests, from which buzzards stole prey. Buzzards occasionally occupied old goshawk nests. These results support our initial hypothesis that interspecific interactions within the raptor guild influence the spatial distribution of predator species in forest ecosystems, with intraguild predation as a key driver. We discuss several mechanisms that may promote the coexistence of subordinate and dominant predators and the spatial assembly of this raptor guild: spatial refuges, different breeding phenology, spatial avoidance, low territory occupancy between neighboring nesting territories, nest concealment and protection, and diet segregation.     

Reproductive Responses of Northern Goshawks to Variable Prey Populations

ABSTRACT Developing comprehensive conservation strategies requires knowledge of factors influencing population growth and persistence. Although variable prey resources are often associated with fluctuations in raptor demographic parameters, the mechanisms of food limitation are poorly understood, especially for a generalist predator like the northern goshawk (Accipiter gentilis). To determine the reproductive responses of goshawks to variable prey populations, we evaluated 823 goshawk breeding opportunities on the Kaibab Plateau, Arizona, USA, during 1994–2002. Concurrently, density was estimated for 4 prey species (2 avian, 2 mammalian). We explored the relationship between goshawk reproduction and prey density at one temporal scale (year) and 2 spatial scales (study area, forest type). Prey density for all 4 species combined accounted for 89% of the variation in goshawk reproduction within the entire study area (P < 0.001), 74% in mixed conifer forest (P = 0.003) and 85% in ponderosa pine (Pinus ponderosa) forest (P < 0.001). We found that an incremental increase in prey density resulted in a greater increase in goshawk reproduction in ponderosa pine forest than in mixed conifer forest, suggesting that the denser structural conditions of mixed conifer forest may have reduced prey availability. Red squirrel (Tamiasciurus hudsonicus) density explained more annual variation in goshawk reproduction within the study area (r² = 0.87, P < 0.001), mixed conifer forest (r² = 0.80, P = 0.001), and ponderosa pine forest (r² = 0.85, P < 0.001) than did any other individual species. Although certain prey species were more strongly correlated with fluctuations in goshawk reproduction than were others, the high model selection uncertainty and the strong relationship between total prey density and number of goshawk fledglings produced indicated that alternate prey species were readily substituted for one another. Therefore, conservation strategies concerned with the status of goshawk populations should incorporate forest management practices that increase the abundance, diversity, and availability of prey resources.     

Use of hunters in wildlife surveys: does hunter and forest grouse habitat selection coincide?

Wildlife monitoring performed by volunteer personnel may suffer from bias with regard to their habitat use. Such errors can lead to erroneous population estimates, evidently influencing both management programmes and research that are based on the monitoring. We used a dataset on hunters’ habitat use in forest while searching for black grouse (Tetrao tetrix) and capercaillie (Tetrao urogallus) to test whether hunters’ utilisation of the habitat was independent of that of grouse or if it corresponded to the grouse habitat preference. Twenty volunteer hunters with dogs registered their tracks and all observations of capercaillie and black grouse in Østfold County, Norway, during August 2003 and 2004. We performed an ecological niche factor analysis (ENFA) and a K-select analysis with respect to hunters’ selection for habitat, described with ecogeographical variables related to forest stand characteristics, as well as the habitat preference of the observed grouse, conditional on the habitat utilisation of the hunters. Individual ENFA on the hunter's tracks revealed large variation in the habitat preferences of the hunters. The K-select indicated few overall patterns in the habitat characteristics of grouse observations, conditional on the hunters selected habitat. Accordingly, the results indicate that hunters’ observation of grouse prior to the hunt may give indicators of changes in grouse density unbiased by habitat preference due to the large between-hunter variation in habitat preference, given that a sufficient number of hunters is used. This suggests that such monitoring programmes can provide information about fluctuations in grouse population sizes valuable for both the management and research of forest grouse species.     

Delayed numerical response of goshawks to population fluctuations of forest grouse

Delayed density-dependent mortality induced by delayed numerical response of predators can drive prey populations to fluctuate in high-amplitude cycles. We studied numerical response of goshawks Accipiter gentilis to varying densities of their main prey (forest grouse) in western Finland during 1979–1996. Occupancy rate of goshawk territories tracked grouse numbers with a two year lag. Occupancy rate of goshawk territories and pooled number of adult and young goshawks correlated negatively with a 1–2 year lag to the chick production of grouse. Goshawk to grouse ratio was negatively related to grouse density. This suggests that goshawk predation on grouse is inversely dependent on grouse density. We conclude that in northern Europe with few alternative preys, goshawk predation might contribute to the generation of multiannual cycles of forest grouse. This should be tested experimentally.     

The impact of predation on the viburnum whitefly, (Aleurotrachelus jelinekii)

Summary In association with a detailed study of the population dynamics of the viburnum whitefly, Aleurotrachelus jelinekii, observations were made on the predators including spiders and five species of insect, notably the mirid Campyloneura virgula and the neuropteran Conwentzia psociformis. Although the species richness of the predator guild was positively correlated with prey population density, at the individual level there was little evidence over twelve years of a density dependent relationship. In these populations natural enemies do not seem to play an important role in the dynamics.     

Cycles in voles and small game in relation to variations in plant production indices in Northern Sweden

Population dynamics for voles (Cricetidae), Tengmalm's owl (Aegolius funereus (L.)), red fox (Vulpes vulpes (L.)) willow grouse (Lagopus lagopus (L.)), black grouse (Lyrurus tetrix (L.)), capercaillie (Tetrao urogallus L.), hazel hen (Tetrastes bonasia (L.)), mountain hare (Lepus timidus L.) and tularemia (Francisella tularensis (McCoy & Chapin)) and game bird recruitment were studied by index methods in northern Sweden. In addition contemporary temperature records and spruce (Picea abies (L.) Karst.) and pine (Pinus silvestris L.) cone crops (as indices for plant production) and the occurrence of forest damage, caused by voles eating bark, were studied.During 1970–80 two synchronous 4-year cycles were observed for voles, predators (Tengmalm's owl and red fox) and their alternative prey species (grouse and mountain hare). In grouse the change of numbers was correlated with that of recruitment. Autumn vole numbers peaked about a year before the other species and extensive forest damage occurred at winter peak densities of voles. These population fluctuations are consistent with a predator-prey model for their regulation. In short the model suggests that vole-food plant interactions trigger the cycle of voles, that voles generate the cycle of predators and that these in turn synchronize alternative prey populations to the others at vole declines.For voles, grouse and red fox the amplitude was higher in the first cycle compared to the second one whilst the opposite was true for the mountain hare. Although temperature and cone crops showed large interannual variations they still implied that herbivore food conditions were better during the former cycle. Hence, the reduction of the amplitude of the vole cycle may be explained by inter-cyclic differences in plant food conditions, implying food shortage (as indicated by bark-eating) at different population levels. The similar decrease of grouse and red fox populations may also be explained by deteriorated food conditions and/or for the fox by an outbreak of sarcoptic mange (Sarcoptes scabiae var. vulpes). The increased amplitude of the mountain hare cycle was part of a long-term rise in numbers after a tularemia epidemic in 1967. This is interpreted as a recovery, probably towards the generally higher pre-epidemic population level.     

Decomposing Predation: Testing for Parameters that Correlate with Predatory Performance by a Social Bacterium

Predator–prey interactions presumably play major roles in shaping the composition and dynamics of microbial communities. However, little is understood about the population biology of such interactions or how predation-related parameters vary or correlate across prey environments. Myxococcus xanthus is a motile soil bacterium that feeds on a broad range of other soil microbes that vary greatly in the degree to which they support M. xanthus growth. In order to decompose predator–prey interactions at the population level, we quantified five predation-related parameters during M. xanthus growth on nine phylogenetically diverse bacterial prey species. The horizontal expansion rate of swarming predator colonies fueled by prey lawns served as our measure of overall predatory performance, as it incorporates both the searching (motility) and handling (killing and consumption of prey) components of predation. Four other parameters—predator population growth rate, maximum predator yield, maximum prey kill, and overall rate of prey death—were measured from homogeneously mixed predator–prey lawns from which predator populations were not allowed to expand horizontally by swarming motility. All prey species fueled predator population growth. For some prey, predator-specific prey death was detected contemporaneously with predator population growth, whereas killing of other prey species was detected only after cessation of predator growth. All four of the alternative parameters were found to correlate significantly with predator swarm expansion rate to varying degrees, suggesting causal interrelationships among these diverse predation measures. More broadly, our results highlight the importance of examining multiple parameters for thoroughly understanding the population biology of microbial predation.     

Spatial dynamics in breeding performance of a predator: the connection to prey availability

Using nationwide long-term data on goshawk and grouse populations in Finland we study the spatial dynamics of the numbers of breeding northern goshawk ( Accipiter gentilis ) pairs, goshawk brood size and offspring sex ratio and their connection to the abundance of grouse. Our first large-scale data comprise of observations on goshawk nests during 1986–2001 pooled to 21 different regions. The second set are annual (1989–1998) observations of brood size and offspring sex ratio (females over the sum of females and males) in goshawk nests all over the country, aggregated to 50 km grid level (n=28 grid units). The third set comprises counts (1989–2001) of four species of woodland grouse, split to adults and juveniles, also given in the same 50 km grid units. Using these data, we show that the annual numbers of northern goshawk nests in the different regions fluctuate in synchrony. Synchrony is also found in long-term fluctuations of northern goshawk brood size and offspring sex ratio. Moreover, synchrony is found in annual numbers of grouse juveniles and adults, the main prey for the northern goshawk. In the brood size and offspring sex ratio of the goshawk, as well as in the annual numbers of grouse juveniles and adults the degree of synchrony falls off with increasing distance. However, only in sex ratios and in grouse dynamics are the slopes of synchrony vs distance roughly matching. We also found that sex ratio either vs grouse juveniles or grouse adults has a more matching spatial dimension (50 km radius) that sex ratio vs brood size. These observation lend support to the hypothesis that goshawk offspring sex ratio and grouse abundance are interconnected. Despite the reason, consequences of spatial coupling in sex ratio could have repercussions on other life history events.     

Variation in foraging success among predators and its implications for population dynamics

The effects of the expected predation rate on population dynamics have been studied intensively, but little is known about the effects of predation rate variability (i.e., predator individuals having variable foraging success) on population dynamics. In this study, variation in foraging success among predators was quantified by observing the predation of the wolf spider Pardosa pseudoannulata on the cricket Gryllus bimaculatus in the laboratory. A population model was then developed, and the effect of foraging variability on predator–prey dynamics was examined by incorporating levels of variation comparable to those quantified in the experiment. The variability in the foraging success among spiders was greater than would be expected by chance (i.e., the random allocation of prey to predators). The foraging variation was density‐dependent; it became higher as the predator density increased. A population model that incorporates foraging variation shows that the variation influences population dynamics by affecting the numerical response of predators. In particular, the variation induces negative density‐dependent effects among predators and stabilizes predator–prey dynamics.