Factors influencing winter home ranges and activity patterns of raccoon dogs <Emphasis Type="Italic">Nyctereutes procyonoides</Emphasis> in a high-altitude area of Japan

Between October 2006 and June 2007, we radio-tracked six adult raccoon dogs Nyctereutes procyonoides (Gray, 1834) in a high-altitude area of Japan to determine which factors influenced home ranges, daily movements, and activity patterns of Japanese raccoon dogs, with an emphasis on the winter season. The home-range sizes for the six individuals were smaller in winter than in autumn. In winter, the daily movement distances significantly decreased in response to decreasing temperature and increasing snow depth, suggesting these environmental factors contributed to a decrease in the home-range sizes during this period. Moreover, during daytime, raccoon dogs were more active in winter than in the snow-free periods (autumn and spring), and the proportion of the daytime movement distances to the total daily movement distances significantly increased as the mean daily air temperatures decreased. Therefore, it appears that the raccoon dogs in our study area passed the winter by minimizing their energy expenditure by restricting their movements when temperatures were at their lowest and snow depth was highest, and by moving more during the daytime when temperatures were higher.     

Conspecific reproductive success affects age of recruitment in a great cormorant, Phalacrocorax carbo sinensis, colony.

Few studies have addressed the proximate factors affecting the age at which individuals of long-lived bird species are recruited into the breeding population. We use capture-recapture analysis of resightings of 16 birth cohorts of colour-ringed great cormorants, Phalacrocorax carbo sinensis, in a Danish colony to assess the evidence for two hypotheses: conspecific attraction (earlier recruitment when the colony is large) and conspecific reproductive success (earlier recruitment following years of high breeding success). For both males and females, conspecific reproductive success was the most important covariate explaining the interannual variation in age of recruitment; colony size was also important for females. These covariates explained nearly 60% of the year-to-year variation for both sexes. The age of recruitment increased for cohorts born after 1990, and this increase was correlated with a decline in breeding success in the colony; we interpret this as an indirect and delayed density-dependent effect. Females were recruited earlier than males (mean age of recruitment for cohorts born before 1990: 2.98 years versus 3.53 years); the most plausible reason for this is a skewed sex ratio in favour of males in the adult population. Recruitment of males may thus, to some extent, be constrained by the availability of females. This study provides the first evidence that conspecific reproductive success can affect the age at which individual birds start to breed.     

Effects of hazard-reduction burning on populations of understorey plant species on Hawkesbury sandstone

Density counts were taken for 37 plant species on 12 plots which were assigned to three treatments: spring burn, autumn burn, and control. Post-fire sampling was carried out at intervals of up to 6 years when a second fire treatment was applied, followed by further sampling at 1 year. Most pre-fire species returned by 12 months after fire. Initial recovery was slower on the autumn burn plots and these were surpassed after 12 months by the spring burn plots which remained significantly more diverse until 6 years after fire when the treatments converged. The second fire led to further loss of species, especially on the autumn burn plots. Overall, numbers of individuals were lower in the post-fire community and did not fully recover during the 6 years before the second fire. Of the 37 species followed over the study period, eight had population numbers consistently below pre-fire values, seven showed better recovery on the autumn burn plots, eight showed better recovery on the spring burn plots, and fourteen had population numbers equal to or greater than pre-fire values. Most of the species not fully recovering had relatively poor survival rates. The better recovery rate of species on the autumn burn plots was attributable to better recruitment while the species recovering on the spring burn plots showed better survival and recruitment. The species increasing their numbers after the fire did so through good recruitment or survival, or both. For some species post-fire rainfall or temperature was significantly correlated with greater post-fire recruitment; others were more strongly affected by the treatment itself or more directly by the somewhat different fire intensities in spring and autumn, the differences in soil and litter moisture content, or seasonal variations in soil seed populations.     

An empirically based estimate of home range

The minimum-convex-polygon method for estimating home-range area, in which the outermost points are connected in a particular way, is extremely sensitive to sample size. Existing methods for estimating home-range area that correct for sample size fail to encompass all the important kinds of biological variation in the home-range utilization. (The home-range utilization describes the relative degree to which different units of space are frequented by an animal.) Although previous methods have assumed specific unimodal distributions, such as the bivariate normal, home-range utilizations may resemble funnels or pies as well as hills. A regression method is introduced that uses data from well-sampled individuals whose true home ranges are assumed approximately known to predict home-range areas for less well-sampled individuals. Appendix 5 summarizes this method. Sizes of home ranges estimated by the regression method are half or less than sizes estimated by previous methods in which utilization distributions are assumed to be all of a particular statistical type.     

How does food distance influence foraging in the ant Lasius niger: the importance of home-range marking

We study the influence of food distance on the individual foraging behaviour of Lasius niger scouts and we investigate which cue they use to assess their distance from the nest and accordingly tune their recruiting behaviour. Globally, the number of U-turns made by scouts increases with distance resulting in longer travel times and duration of the foraging cycle. However, over familiar areas, home-range marking reduces the frequency and thereby the impact of U-turns on foraging times leading to a quicker exploitation of food sources than over unmarked set-ups. Regarding information transfer, the intensity of the recruitment trail reaching the nest decreases with increasing food distance for all set-ups and is even more reduced in the absence of home-range marking. Hence, the probability of a scout continuing to lay a trail changes along the homeward journey but in a different way according to home-range marking. Over unexplored setups, at a given distance from the food source, the percentage of returning trail-laying ants remains unchanged for all tested nest-feeder distances. Hence, the tuning of the trail recruiting signal by scouts was not influenced by an odometric estimate of the distance already travelled by the ants during their outward journey to the food. By contrast, over previously explored set-ups, a distance-related factor – that is the intensity of home-range marking – strongly influences their recruiting behaviour. In fact, over a home-range marked bridge, the probability of returning ants maintaining their trail-laying behaviour increases with decreasing food distance while the gradient of home-range marks even induces ants which have stopped laying a trail to resume this behaviour in the nest vicinity. We suggest that home-range marking laid passively by walking ants is a relevant cue for scouts to indirectly assess distance from the nest but also local activity level or foraging risks in order to adaptively tune trail recruitment and colony foraging dynamics. Received 13 July 2004; revised 26 January and 20 May 2005; accepted 2 July 2005.     

Elephant spatial use in wet and dry savannas of southern Africa

The influence of elephants on woody vegetation cover varies from place to place. In part this may be due to the way elephants utilize space across landscapes and within their home ranges in response to the availability and distribution of food. We used location data from 18 cows at six study sites across an east to west rainfall gradient in southern Africa to test whether wet- and dry-season home-range sizes, evenness of space use within seasonal home ranges and range overlap between seasons and between years, differed between wet and dry savannas. We then tested whether the quantity, distribution and seasonal stability in vegetation productivity, a coarse measure of food for elephants, explained differences. Elephants in wet savannas had smaller wet- and dry-season home ranges and also returned to a higher proportion of previously visited grid cells between seasons and between years than elephants living in dry savannas. Wet-season home-range sizes were explained by seasonal vegetation productivity while dry-season home-range sizes were explained by heterogeneity in the distribution of vegetation productivity. The influence of the latter on dry-season home ranges differed among structural vegetation classes. Range overlap between seasons and between years was related to inter-seasonal and inter-annual stability in vegetation productivity, respectively. Evenness of elephant spatial use within home ranges did not differ between savanna types, but it was explained by seasonal vegetation productivity and heterogeneity in the distribution of vegetation productivity during the wet season. Differences in elephant spatial use patterns between wet and dry savannas according to vegetation structure and season may need to be included in the development of site-specific objectives and management approaches for African elephants.     

Wolf population dynamics in the U.S. Northern Rocky Mountains are affected by recruitment and human-caused mortality

Reliable analyses can help wildlife managers make good decisions, which are particularly critical for controversial decisions such as wolf (Canis lupus) harvest. Creel and Rotella (2010) recently predicted substantial population declines in Montana wolf populations due to harvest, in contrast to predictions made by Montana Fish, Wildlife and Parks (MFWP). We replicated their analyses considering only those years in which field monitoring was consistent, and we considered the effect of annual variation in recruitment on wolf population growth. Rather than assuming constant rates, we used model selection methods to evaluate and incorporate models of factors driving recruitment and human-caused mortality rates in wolf populations in the Northern Rocky Mountains. Using data from 27 area-years of intensive wolf monitoring, we show that variation in both recruitment and human-caused mortality affect annual wolf population growth rates and that human-caused mortality rates have increased with the sizes of wolf populations. We document that recruitment rates have decreased over time, and we speculate that rates have decreased with increasing population sizes and/or that the ability of current field resources to document recruitment rates has recently become less successful as the number of wolves in the region has increased. Estimates of positive wolf population growth in Montana from our top models are consistent with field observations and estimates previously made by MFWP for 2008–2010, whereas the predictions for declining wolf populations of Creel and Rotella (2010) are not. Familiarity with limitations of raw data, obtained first-hand or through consultation with scientists who collected the data, helps generate more reliable inferences and conclusions in analyses of publicly available datasets. Additionally, development of efficient monitoring methods for wolves is a pressing need, so that analyses such as ours will be possible in future years when fewer resources will be available for monitoring. © 2011 The Wildlife Society.     

Effects of Sarcoptic Mange on Coyotes at Wind Cave National Park

Abstract: Home-range size and population abundance indices of coyotes (Canis latrans) have not been documented in Wind Cave National Park, South Dakota, USA. In 2003 and 2004, we captured a total of 26 coyotes and radiocollared 22 adults (12 F, 10 M). In 2003 and 2004, 2 of 17 (12%) and 5 of 9 (56%) coyotes, respectively, were infected with sarcoptic mange (Sarcoptes scabiei) at the time of capture. Thus, objectives were modified to document effects of the mange epizootic on the coyote population. In 2003, home-range (adaptive-kernel) sizes for male coyotes with mange and those considered healthy were 8.26 ± 1.63 (SE) km² and 9.67 ± 2.80 km², respectively. In 2004, home-range sizes for those male coyotes with and without mange were 22.69 ± 9.06 km² and 12.51 ± 2.73 km², respectively. Male home-range size did not differ between years (P = 0.14) or by status (with or without mange; P = 0.84). Survival of collared coyotes was 60% at the end of 2003. Results from fecal line transects, an index of relative abundance, indicated that the coyote population decreased by 48% from 2003 to 2004. Continued monitoring of sarcoptic mange epizootics will enable managers to assess the effects of mange on coyote populations.     

Are whooping cranes destined for extinction? Climate change imperils recruitment and population growth

Identifying climatic drivers of an animal population's vital rates and locating where they operate steers conservation efforts to optimize species recovery. The population growth of endangered whooping cranes (Grus americana) hinges on juvenile recruitment. Therefore, we identify climatic drivers (solar activity [sunspots] and weather) of whooping crane recruitment throughout the species’ life cycle (breeding, migration, wintering). Our method uses a repeated cross‐validated absolute shrinkage and selection operator approach to identify drivers of recruitment. We model effects of climate change on those drivers to predict whooping crane population growth given alternative scenarios of climate change and solar activity. Years with fewer sunspots indicated greater recruitment. Increased precipitation during autumn migration signified less recruitment. On the breeding grounds, fewer days below freezing during winter and more precipitation during breeding suggested less recruitment. We predicted whooping crane recruitment and population growth may fall below long‐term averages during all solar cycles when atmospheric CO₂ concentration increases, as expected, to 500 ppm by 2050. Species recovery during a typical solar cycle with 500 ppm may require eight times longer than conditions without climate change and the chance of population decline increases to 31%. Although this whooping crane population is growing and may appear secure, long‐term threats imposed by climate change and increased solar activity may jeopardize its persistence. Weather on the breeding grounds likely affects recruitment through hydrological processes and predation risk, whereas precipitation during autumn migration may influence juvenile mortality. Mitigating threats or abating climate change should occur within ≈30 years or this wild population of whooping cranes may begin declining.     

Growth and carbon allocation of Agropyron desertorum following autumn defoliation

Summary Growth and carbon allocation of a cool season tussock grass, Agropyron desertorum, following defoliation of newly initiated tillers in the autumn of 1988 and 1989 were investigated. Tiller density and mortality, reproductive shoot density, root density, biomass, individual tiller weight, carbon allocation, and soil water depletion were used to evaluate the response of A. desertorum to autumn grazing. Tiller recruitment was lower in the autumn-defoliated treatment in both years compared with the control because of the cessation of tiller development following autumn defoliation. Autumn defoliation also significantly reduced the movement of ¹³C to the roots in 1988 but not in 1989. Soils were cooler and drier in 1989. Other plant growth measurements and soil water depletion rates were not different between treatments. Autumn defoliation in 1988 did not influence tiller recruitment in the following autumn. Two consecutive years of autumn defoliation did not affect tiller overwinter mortality or peak standing crop in 1990.     

Demography of alpine red squirrel populations in relation to fluctuations in seed crop size

Vertebrate population dynamics, social organisation and space use often are closely associated with the distribution of critical resources, such as food. Tree squirrels are ideal models to study these relationships, since both key demographic parameters (reproduction, survival and dispersal) and spatio-temporal variation in food supplies (measured as seed-crop size) can be reliably estimated. In this paper we test the following two predictions underlying the association between annual food abundance and demography in six alpine red squirrel populations, both with and without time-lag effects: 1) between-season and between-year fluctuations in survival rate, population density and increase parallel those in food availability; and 2) individuals follow a resource tracking strategy and increase in density mainly the year after a rich seed-crop. Red squirrels occurred at higher densities in Scots pine forest, characterised by stable seed-crops, than in Norway spruce with more abundant but more variable seed crops. Fluctuations in numbers were positively correlated with food availability, measured as annual conifer seed-crop sizes. Overall, adult survival rates were higher than those of subadults, and survival substantially fluctuated between seasons and years. Autumn densities and rates of population increase (summer-autumn) were strongly correlated with the same year's autumn seed-crop, while correlations with the previous year's seed-crop (time-lag models) were either weak (population density) or absent (population increase). Results of this paper show that fluctuations in red squirrel densities in habitats with strong temporal variation in seed production are more closely linked with food availability than in more stable habitats. In addition, in the Alpine conifer forests squirrel population sizes, in autumn, increase in synchrony with food resources, eliminating the population lag normally present when resources are produced in pulses.     

High haplotype diversity with fine-scale structure in a recently established population of an endangered orchid

New populations of endangered plants may establish outside of protected areas if a suitable habitat becomes available. However, it is unclear whether such populations are genetically uniform, as a result of a founder effect, or whether they contain genetic variation resulting from continuous gene flow from source populations, and subsequent recruitment after establishment. We addressed this by examining haplotype variation in the endangered orchid, Anacamptis robusta, which has formed a new population outside of its protected area within the past 20 years. To assess population growth, the number of A. robusta flowering plants were counted every year for 22 years in both the new population and populations within the reserve. Haplotype diversity and fine-scale structure were examined with spatial autocorrelation analysis, both in the new population and representative populations from the protected area. The number of flowering individuals in the new population increased from 9 to 2,277 between 2003 and 2012, whereas within the reserve flowering population sizes varied between years. Seventeen of 23 haplotypes detected for A. robusta were detected in the new population, with seven of these occurring more frequently in the new population than reserve populations. In the new population, there was strong fine-scale spatial structure of haplotypes, similar to patterns found in large populations from the protected area, suggesting multiple colonization events and subsequent local recruitment. This highlights that ongoing demographic and genetic monitoring of plant populations is vital to improve our understanding of population colonization and the conservation of narrow endemics.     

Some population dynamics of the Bank vole, Clethrionomys glareolus and the Wood mouse, Apodemus sylvaticus in mixed woodland

The population dynamics of Clethrionomys glareolus Schr.and Apodemus sylvaticus (L.) in mixed woodland in County Durham, were studied from March 1963 to January 1965. Two areas of 0.9 ha (2.25 acres) each were trapped monthly and information was obtained on population size, reproduction, survival and growth from marked animals.
Clethrionomys populations increased from June to an autumn peak; then declined, at first rapidly, but then more slowly in winter, before reaching a spring trough. The breeding season of Clethrionomys was from May to December; juveniles were caught from mid-June to December. Survival was in general poor during the breeding season but good at other times. Survival of young born early in the summer was particularly good on one of the areas and some individuals lived long enough to breed in two successive years. On both areas young born early in the year matured rapidly and bred in the year of their birth. Young born in late summer and early autumn ceased growing at a weight of about 14.5 g, remained immature, and formed the bulk of the overwintering population. Growth was completed at the time of sexual maturation the following spring.
Trapping failed to provide adequate samples of juvenile Apodemus in summer to account for subsequent recruitment. The possibility that a substantial proportion of the adult population of this species was also either trap shy or had emigrated temporarily is discussed. Breeding occurred from April until the following January, but the numbers trapped remained very low throughout the early months of the breeding season. Large scale recruitment of young fecund animals into the trap-revealed population occurred during the autumn. Apodemus males continued to grow rapidly during the winter.     

Spatial segregation of two vole species (Arvicola sapidus and Microtus cabrerae) within habitat patches in a highly fragmented farmland landscape

Spatial segregation is one of the common mechanisms allowing the co-existence of similar interacting species in heterogeneous environments. Analysing spatial segregation requires information on individual home-range sizes and their degree of spatial overlap. In this study, we used radio-tracking to report for the first time the home-range and core-area sizes of sympatric Cabrera and water voles and to analyse intra- and inter-specific space sharing within habitat patches in a highly fragmented landscape. Results indicated that both species exhibited strong fine-scale site fidelity and reduced variation in range size across sexes and seasons. Monogamous mating system seemed to prevail for both species, although water voles may also exhibit polygynous breeding strategies. Mean home-range and core-area sizes of water voles (946.3 and 156.6 m²) were about twice that of Cabrera voles (418.2 and 55.1 m²). Within habitat patches, individuals of both species often overlapped their home ranges, particularly during the dry season (May–September), though intra-specific home-range overlap was generally higher than inter-specific overlap. Inter-specific space sharing was restricted to areas outside the centre of activity of animals, as no core-area overlap was ever recorded between Cabrera and water voles. Taken together, results support the view that co-existence of Cabrera and water voles in Mediterranean patchy habitats may in part result from spatial segregation among individuals, which may reflect competitive displacement or small-scale habitat partitioning. Results highlight the need to account for species interactions when designing conservation management strategies for sympatric Cabrera and water voles in fragmented landscapes.     

Culling Versus Density Effects in Management of a Deer Population

Abstract: Wildlife managers often manipulate hunting regulations to control deer populations. However, few empirical studies have examined the level of hunting effort (hunter-days) required to limit population growth and demographic effects through harvesting of females. Moreover, the relative importance of density effects on population growth has not been quantified. We reconstructed a sika deer [Cervus nippon] population over a period of 12 years (1990–2001) using age- and sex-specific harvest data. Using cohort analysis, we analyzed population dynamics, focusing on 1) the relationship between hunting effort and hunting-induced mortality rate, 2) relative contributions of hunting mortality and recruitment of yearlings to annual changes in population growth rate, and 3) annual variation in recruitment rate. Population size increased until 1998 and declined thereafter. The population growth rate changed more in response to annual changes in recruitment rate than hunting mortality rate. Temporal variation in recruitment rate was not controlled by birth rate alone; direct density dependence, intensities of hunting mortality for fawns, and for females (≥2 yr of age), which accounted for the fawn survival rate, were required as factors to explain temporal variation. Density effects on the recruitment rate were not strong enough to regulate the population within the study period; high hunting mortality, with intensive female harvesting, was necessary to prevent population growth. Hunting effort was a good predictor of the hunting mortality rate, and female harvest had a negative effect on the recruitment rate through fawn survival. We suggest that >3,500 hunter-days and prioritization of female harvesting are required to prevent increases in this deer population.     

Effects of farming practices on spatial behaviour of common voles

This study aimed to reveal changes in spatial behaviour of common voles (Microtus arvalis) after alteration of their habitat by farming practices. Radio-collared common voles were tracked before and after mulching, mowing, harvesting wheat, and ploughing in the flood plain of the river Unstrut in central Germany. Voles undisturbed by agricultural practices were tracked on a mulchland, an abandoned pasture, and a cattle pasture. There was a large decrease in home-range size after harvesting wheat (96%, P<0.001). Changes after mowing (–74%, P=0.06) were almost significant whereas changes after mulching were not (+14%, P=0.60). On the cattle pasture we found a decrease in home-range size (42%, P=0.03) possibly due to increased spatial activity of cattle in autumn. There was a positive correlation of home-range size and vegetation height for plots with and without farming activity but no correlation with vegetation cover, population density, and breeding. Radio-collared common voles did not show evasive movements and farming practices did not cause a shift of centres of activity. Common voles clearly reacted to sudden changes in vegetation height, which may indicate an immediate response to predation risk. Electronic Publication     

Factors influencing home range and activity of wild boar (Sus scrofa) in a Mediterranean coastal area

This paper analyses the temporal variation in the size of home range and activity of adult wild boar ( Sus scrofa L.). Eighteen boar were radiotracked between 1991 and 1993 in the Maremma Natural Park (Central Italy). Variations of home-range size and activity were related to sex, winter food availability and population density. The hypotheses that home-range size was inversely related to food availability and population density, and that the amount of activity was directly related to population density and inversely related to food abundance, were tested. The size of the acorn crop was used as a measure of winter resource level. No differences were found in the size of ranges, core area, and activity between males and females. In 1991–92 home-range size was larger than in 1992–93; conversely, activity increased from 1991–92 to 1992–93. Winter food availability was high in 1991–92 and low in the following year. Population density increased greatly in spring 1992 but crashed in spring and summer 1993, when a high mortality of wild boar occurred due to starvation.
The decrease in home-range size during the food shortage was explained as a possible strategy adopted by wild boar to cope with starvation. The difference in home-range size and amount of activity found between the first and the second year of study were probably influenced by differences in population density, although the individual effects of food availability and population density were not clearly separable.     

Diet and condition of wild boar, Sus scrofu scrofu, without supplementary feeding

From 1987 to 1992 diet and condition of two populations of wild boar which received no supplementary feeding were studied in the Veluwe area, a large area of heathlands and forests in The Netherlands, and were compared with those obtained in a previous study (1974–1976), when supplementary food was provided. Composition of stomach contents depended mainly on season, mast availability, and area-specific factors, whereas sex and age were of little or no importance. Density dependence was found for the decrease in mast (tree seed) consumption from autumn to winter. In autumn, and in winters of rich mast years, mast was the main constituent of the diet. In winters of poor mast years this was replaced by broadleaved grasses in one area and by broadleaved grasses, wavy hairgrass, and roots in the other. We found no important differences between the stomach contents of animals receiving no supplementation, and the natural fraction during a period of supplementary feeding. Variation in body weight was related mainly to age and sex, but also to mast availability. Judged by relative loss of body weight and decrease of bone marrow fat, juveniles seemed to suffer more from poor mast availability than adults. The decrease in body weight from autumn to winter was greater when population density was high. In poor mast years, recruitment into the population receiving no supplementation depended on the availability of broadleaved grasses; in rich years, recruitment was still lower than in populations receiving supplementary feeding. In populations receiving supplementary feeding, recruitment seemed independent of mast availability.     

Density dependent processes and the importance of periodic germination in the winter annual plant Saxifraga osloensis

Saxifraga osloensis is an allopolyploid plant, endemic to Scandinavia A field study of Its population biology was performed during two years, 1989-90 and 1990-91 The timing of different life stages in spring was correlated with weather conditions, mainly temperature Density effects were found on survival of rosettes in autumn and on capsule production in spring It seemed as if the density effects occurred only when the initial density was very high or when there was shortage of water Most plants emerged in autumn, but between 10% and 15% emerged m spring Spring gemunators were exposed to higher mortality risks in spring than autumn germinators Fitness, estimated as life-time capsule production, was lower for spring cohorts both years Most plants in the spring cohorts produced no seeds Thus, spring germination should not be selected for in this population     

Patterns in vendace recruitment in Lake Pyhäjärvi, south-west Finland

A vendace Coregonus albula population has long formed the basis of a flourishing fishery in Lake Pyhäjärvi, south-west Finland. Between 1971 and 1990, it exhibited variations in year-class strength generally characterized by a 2-year cycle of alternating strong and weak year classes, which implied density-dependent regulation probably due to asymmetrical food competition between age groups. The abundance of piscivorous predators in the lake and the wanning of water after the hatching of larvae in spring together determined the final year-class strength of vendace. Since 1990, year-class sizes have remained very low and the 2-year cycle has been disrupted. The continuously low levels of recruitment could not be explained by an increased larval mortality, although the decline started with 2 successive years with very high mortality of this life stage which reduced the whole spawning stock to a fraction of earlier levels, and consequently led to low larval numbers such that between 1988 and 1996 larval abundance was positively correlated with the spawning stock size. The importance of the spawning stock size to recruitment could also be seen in the whole 26-year time series as a significant correlation between year-class sizes with a 2-year lag. The observed recruitment patterns of the vendace stock in Lake Pyhäjärvi thus show the importance of spawning stock size, intraspecific competition, predator effects and abiotic environmental variation.