Harvesting a red-listed species: determinant factors for willow ptarmigan harvest rates,bag sizes,and hunting efforts in Norway

A main objective of game management on public lands in Norway is to ensure that harvest levels are sustainable while securing hunting access to the public. The willow ptarmigan is a popular but red-listed small game species, and it is challenging for managers to optimize harvest levels based on uncertain population estimates and limited knowledge of the factors influencing hunting impact. We studied how willow ptarmigan hunting effort, catch per unit effort, and harvest rates were influenced by ptarmigan density, topography, vegetation, and infrastructure. Data were collected during 2013–2016 from 162 hunting blocks covering 26,828 km² state-owned land in the counties Nordland and Troms, northern Norway. Hunting effort averaged 1.27 days/km² (SD?=?2.08), and it was the highest in areas close to roads and cabins. We found an opposite effect on the catch per unit effort (CPUE, mean?=?0.77 ptarmigan/day, SD?=?0.48), which was lower close to roads. There was a marked positive effect of density on CPUE, whereas terrain steepness (slope) had a negative effect. On average, harvest removed <?10% of the autumn population in the hunting areas during the study period. This indicates a relatively low and partially compensatory hunting mortality in most years and areas. Although this study shows a low harvest rate, we recommend managers to survey ptarmigan populations, hunting effort, and harvest, especially in easily accessible areas close to infrastructure and in periods of low population density.     

Successful adult willow grouse are exposed to increased harvest risk

Age and sex ratios in bag records are frequently used as indices of population composition for harvested populations. However, vulnerability to harvest may differ by age and sex thereby producing bias in population estimates. We assessed whether age and sex affected vulnerability to harvest for willow grouse (Lagopus lagopus) where adult density and brood size were known in the harvested populations. We collected bag records during 2 days of controlled hunting in 4 areas in 2 years (2007 and 2008) in Jämtland county, Sweden. We found that vulnerability to harvest was different for chicks and adults, but not between male and female adults. Hunters encountered broods at a higher rate than single birds compared to personnel conducting pre-harvest counts along line transects. Furthermore, the probability of shooting a grouse was higher in encounters of broods than individual grouse. Proportionally, we calculated about a 50% probability of a hunter shooting either a chick or an adult independent of encountering a single bird or broods of 2–10 grouse. Increasing adult density also increased the vulnerability to harvest for adults relative to chicks, independent of the chick to adult ratio in the pre-harvest population. The different vulnerability of adults and chicks to harvest observed in this study will dampen variation in age classes in bag records compared to the population, and we caution against extrapolation of age ratios in bag records to harvested populations. © 2012 The Wildlife Society.     

Big-game,small-game,and gamebird hunting in New Zealand: Hunting effort,harvest, and expenditure in 1988

Abstract

A postal survey of 8639 licensed firearm owners in 1989 indicated that an estimated 117 200 ± 6300 New Zealanders (3.5% of the total population) did some hunting in 1988. An estimated 33 100 former hunters did not hunt in 1988 but thought it likely that they would hunt again in future. The survey provided useful estimates of 1988 national totals for hunting effort (4.4 million hunter days), gross expenditure ($NZ100 million), and harvest (6.5 million animals).

Small-game hunting dominated, involving 81% of hunters, 59% of total hunting effort, and 86% of total numerical harvest: rabbits (Oryctolagus cuniculus) and possums (Trichosurus vulpecula) each comprised 40% of the national bag. Gamebird hunting involved 48% of hunters, 19% of effort, and 11% of total harvest: ducks comprised 73% of the gamebird harvest. Although big-game hunting attracted the fewest hunters (42%) it ranked second for hunting effort (21%). Big game formed 3% of the total numerical harvest: pigs (Sus scrofa), goats (Capra hircus), and red deer (Cervus elaphus scoticus) were the most commonly taken. Big-game were estimated to comprise 49% of total harvest biomass, followed by small-game (47%), and gamebirds (4%). Most deer (nearly 60%) were taken for recreation, with helicopter-based hunting accounting for only one-third the total deer harvest.

A quarter of those people hunting in 1988 hunted on five or fewer days that year, and a relatively small group of mainly professional hunters accounted for a disproportionately large share of the overall harvest Hunters reported spending an average of $851 each on hunting in 1988. Expenditure on big-game comprised 44% of the total, small-game 23%, and gamebirds 33%. Expenditure per animal harvested or per day hunted was greater for big-game animals (other than goats) than for gamebirds, which were more expensive to hunt than small-game. Total hunting effort was inversely related to the average expenditure per animal harvested, regardless of the type of game.     

Hunters serving the ecosystem: the contribution of recreational hunting to wild boar population control

The extractive nature of recreational hunting may provide a service to both the ecosystem and society, namely the control of problem species. We reviewed the annual wild boar hunting bag data from hunting sites in Asturias (Spain) from 2000/01 to 2013/14, paying particular attention to the evolution on hunting estates after ban periods. We hypothesized that the annual hunting bag after a hunting ban would be larger than that of the pre-ban period, and that this difference could provide an indication of hunters’ relative contribution to wild boar population regulation. The total hunting bag grew during the study period, from 3723 wild boar (0.39ind/km²) in the 2000/01 hunting year to 7593 in that of 2013/14 (0.79ind/km²)—a mean annual increase of 5.63%. Low hunting quotas cannot be blamed for these growing trends, since no more than 50% of the authorized animals are hunted. The growth of the mean annual pre-ban hunting bag on the estates on which hunting bans took place was 8.46%. The hunting bag grew by 40.33% immediately after the hunting ban ended—a growth rate seven times higher than that of the background hunting bag. This constitutes a proxy of the regulatory effect of hunters on wild boar population growth. Following the remarkable increase after the ban, the wild boar hunting bag attained values that were slightly lower than those of the pre-ban period, which indicates that hunters are able to reduce wild boar abundance. Hunting, therefore, provides an important service to both the ecosystem and society by contributing to regulating the growth of problem species such as the wild boar.     

The optimal foraging theory, crowding and Swedish grouse hunters

Hunters that have options to hunt in different areas should evaluate their previous hunting success when they decide where to hunt. Following optimal foraging theory for non-human predators, we investigated if hunting success and density of other hunters on the hunting area will affect the probability of return to the same area, and if such behavioural changes will result in a higher hunting success compared to hunters that change to a new area. For this purpose, we used detailed information about willow grouse (Lagopus lagopus) hunters on state-owned land in Sweden. We found support for the optimal foraging theory application on grouse hunters’ behavioural changes according to hunting success. The return rate increased with increasing hunting success, and hunters that returned to the same area also increased their success compared to hunters that changed to a new area. Only one third of the hunters returned to the same area the subsequent year. We also found a negative effect of density of hunters in an area on hunters’ return rates and their hunting success, suggesting crowding among Swedish grouse hunters.     

Coping with wild boar in a conservation area: impacts of a 10-year management control program in north-eastern Argentina

Wild boar (Sus scrofa) are of serious concern in numerous conservation areas such as El Palmar National Park, Argentina, where their increasing abundance affected the iconic palm tree Butia yatay. We assessed the effectiveness of an innovative management control program on wild boar population dynamics and ground rooting area over 10 years. Park personnel recruited and supervised local recreational hunters who regularly conducted controlled still shooting from widely distributed watchtowers and used trained dogs mainly during the first 2 years post-intervention (YPI). We used the detailed records of harvest and hunting effort to estimate time- and stage-specific catch-per-unit-effort (CPUE) indices and stochastic population growth rates (μ). Catch was linearly related to hunting effort except at large effort levels. CPUE indices declined exponentially at 5–7 % month^?1 over 0–1 YPI and thereafter stabilized with μ indistinguishable from 0. Relative to baseline levels, culling reduced annual pregnancy rates, the fraction of juveniles and older adults, and ground rooting area below target values (<1.3 %). Incipient population recovery followed one of two periods of marginal hunting effort. The program generated few undesirable collateral effects mainly related to dog-hunting. Mesopredator abundance (foxes) steadily increased following a large outbreak of canine distemper at baseline. The combined use of standardized CPUE indices, body-length data and simple population viability analysis models provided reliable metrics for wild boar trend analysis and management. Unlike a preceding plan, a highly structured multi-stakeholder program proved to be sustainable and brought wild boar abundance to a low-density, unstable equilibrium causing minimal damage.     

Spatial synchrony in red grouse population dynamics

Red grouse Lagopus lagopus scoticus populations exhibit unstable dynamics that are often characterised by regular periodic fluctuations in abundance. Time-series' of grouse harvesting records collected from 287 management units (moors) across Scotland, England and Wales were analysed to investigate the broad scale patterns of synchrony in these fluctuations. Estimation of the spatial autocorrelation of grouse population dynamics across moors indicates relatively high levels of synchrony between populations on adjacent moors, but that this synchrony declines sharply with increasing inter-moor distance. At distances of greater than 100  km, grouse population time-series exhibit only weakly positive cross-correlation coefficients. Twenty-eight geographical, environmental and other candidate variables were examined to construct a general linear model to explain variation in local synchrony. Grouse moor productivity (average size of shooting bag), distance from the Atlantic coast moving in a north-easterly direction, April and June temperatures, and June rainfall significantly increased the explanatory power of this model. An understanding of the processes underlying synchrony in red grouse population dynamics is a prerequisite to anticipating the effects of large-scale environmental change on regional patterns of grouse distribution and abundance.     

Temporal changes in age and sex ratios of forest grouse harvested in northeastern oregon

Ratio of immature (young of the year) grouse to adult birds (I:A) in the harvest of upland game birds is commonly used as an index to annual reproduction; however, I:A ratios can vary as the season progresses producing biased estimates. We analyzed I:A ratios in the daily harvest of dusky grouse (Dendragopus obscurus) and ruffed grouse (Bonasa umbellus) in northeastern Oregon over 28 years (1981–2008) and found that I:A ratios in the harvest declined for both species as the hunting season progressed. We also analyzed ratios of adult female to adult male (AF:AM) grouse to determine if female and male grouse were harvested in equal numbers throughout the harvest season. We found that more males than females of both species were harvested, but that AF:AM ratio of both dusky and ruffed grouse did not change during most of the hunting season. Approximately 50% of the annual harvest occurred during the first 14 days of the hunting season. Therefore, we recommend using the ratios of I:A birds in the first 14 days of the harvest season as the best index to annual reproduction of forest grouse in northeast Oregon. © 2011 The Wildlife Society.     

The role of game management in wildlife populations: uncertainty analysis of expert knowledge

Uncertainties about future states of wildlife populations make it difficult to pre-adapt to possible threats and ensure sustainability of resources and harvesting over the long term. This uncertainty is partly due to the unknown impact and future states of many factors that explain population sizes and variation. In this paper, the effect of local game management activities on the uncertainty of future population sizes of groups of Finnish wildlife species (ungulates, forest grouse, large predators, small predators and mountain hare) was analysed using expert knowledge and the Bayesian belief networks (BBNs) modelling techniques. As a result, the current knowledge and agreement of the relationships between wildlife population sizes and the game management activities explaining their variation as well as trends are evaluated. Information given to hunters and the number of hunters were seen as the most effective factors for the management of game populations. However, there were great uncertainties in the expectations regarding future trends in the management activities, especially in feeding, and there was disagreement in the direction of the trend in the length of the hunting season. The trends in the size of forest grouse populations were viewed as the most uncertain trend among species groups. At the same time, forest grouse were seen as the most regulated species group by local game management. Among interest variables, experts were very uncertain and they disagreed about the direction of the trend in the recreational value of hunting.     

Where is the pulse to have the finger on? A retrospective analysis of two decades of Alpine Galliforms (Aves: Galliformes) census and game bag data in Italy

Information on the abundance of the Italian populations of black grouse (Lyrurus tetrix), Alpine rock ptarmigan (Lagopus muta helvetica) and Alpine rock partridge (Alectoris graeca saxatilis) rely only on extrapolations of local data to the national scale, since there is no national standardized survey. Consequently, their status is virtually unknown. We performed a first-ever assessment of a medium-term (1996–2014) population trend of these species using and comparing post-breeding count and bag data at hunting district scale. These data were collected from various authorities in charge of wildlife management and allowed us to test the influence of hunting policies on the estimated trends. Rock partridge showed a stable trend with numbers fluctuating between years, while there was evidence of a severe decline for rock ptarmigan. No general conclusion could be drawn for the black grouse, as we detected lack of consistency of count and bag data. Counts were greatly overdispersed as a result of an uneven count effort among hunting districts. Adding the game management authority as model covariate resulted in more robust trend estimations, suggesting a significant effect of different policies that emerged also as similar hunting pressure across species within authorities. Hunting effort variation over the time was instead negligible. Species-specific game management bias is discussed. Our results highlight the need for a survey scheme or guidelines to be applied uniformly at a national scale.     

Effects of Earn-a-Buck and Special Antlerless-Only Seasons on Wisconsin's Deer Harvests

ABSTRACT Effective management of wildlife populations often requires motivating hunters to harvest sufficient numbers of animals of prescribed sex and age classes to meet management goals. For cervids, it is convenient to design harvest regulations relative to presence (male) or absence (young and female) of antlers because harvest of females has a larger effect on population growth. We used regression techniques to evaluate effects of 2 supplemental hunting programs based on additional days of hunting opportunity and an additional incentive used to complement additional days on harvest of antlered and antlerless deer in Wisconsin, USA. Earn-a-buck regulations, an incentive-based program that requires hunters to register an antlerless deer before being authorized to harvest an antlered deer, were associated with an average increase of 2.04 deer/km² in antlerless harvest and a 0.60 deer/km² decrease in harvest of antlered deer. Providing more opportunity for hunting of antlerless deer in the form of 4- and 8-day supplemental firearm seasons was associated with 1.10 deer/km² and 1.32 deer/km² increases, respectively, in antlerless harvest with trivial (0.02 deer/km² and 0.09 deer/km²) decreases in harvests of antlered deer. Our analysis suggests that extra days of hunting opportunity coupled with the earn-a-buck incentive was 56–88% more effective at increasing antlerless harvest relative to additional days of hunting without the incentive. Use of the earn-a-buck incentive resulted in decreased harvest of antlered deer and was disliked by many hunters. Quantifying these relationships is important for helping managers predict the costs and benefits of various hunting programs.     

Are indirect measures of abundance a useful index of population density? The case of red grouse harvesting

Indirect measures of population abundance, such as harvest data, are often used to make inference on long term population dynamics when direct data are either not available or are logistically difficult to obtain. However, when harvesting records are used, a common concern is that they may not reflect actual population abundance. We investigated the extent to which harvest data reflected changes in population density of the red grouse Lagopus lagopus scoticus in Great Britain. We used 92 independently managed populations over the period 1977–2000 and examined the temporal and spatial variability of the hunting records and independently obtained count data from each of these managed estates. Three different analyses support the conclusion that grouse hunting records are a reliable indicator of grouse abundance: 1) the number of red grouse shot in autumn showed a tendency to be linearly related to the density of individuals counted in the summer prior to the harvesting, 2) the relationships between the variance and the mean in the harvesting and corresponding count data, calculated over different populations at the same time, or the same locations at different times, were not statistically distinguishable, 3) similar direct and delayed density dependence patterns were observed in hunting records and count data. Our results suggest that red grouse hunting time series are a good proxy for population abundance.     

Distribution of mountain hares Lepus timidus in Scotland: results from a questionnaire

A questionnaire survey of land owners, managers and gamekeepers was conducted in order to assess the distribution of mountain hares in Scotland, assess their current management, collate numbers harvested in 2006–07 and estimate distribution change by comparing with similar data collected in 1995–96. The land area covered by returned questionnaires was 71098km² (90% of Scotland). Mountain hares were reported as present on 34359km² (48%) and absent from 36739km² (52%). Mountain hare presence was strongly associated with heather moorland managed for red grouse shooting. Moorland managed for driven grouse shooting had the highest percentage area of mountain hare presence (median 64%) followed by moorland managed for walked‐up grouse shooting (median 9%) and moorland with no grouse shooting (median 0%). Approximately 25000 mountain hares were harvested in 2006–07. Based on the estimated UK population in 1995 of 350000 (range ±50%), this represents around 7% of the population (range 5–14%). Reasons given by respondents for harvesting hares were tick control (50%), sport (40%) and forestry or crop protection (10%). Comparison of the estates surveyed in both 2006–07 and 1995–96 (a total area of 20462km²) indicated no net gain or loss in hare distribution. Furthermore, there was no evidence that levels of harvest had reduced the range of mountain hares in this area. It is not possible to comment on any distribution change outside this area (58737km²). Similarly, as no data were collected on abundance, it is not possible to draw conclusions on changes in density. Regular monitoring of mountain hare distribution within Scotland is required to identify any distribution changes. Measures of abundance throughout the range are necessary to estimate the population size, investigate the relationship between harvest intensity and changes in abundance and further assess the conservation status of this UK Biodiversity Action Plan species.     

Mark–recapture using tetracycline and genetics reveal record-high bear density

We used tetracycline biomarking, augmented with genetic methods to estimate the size of an American black bear (Ursus americanus) population on an island in Southeast Alaska. We marked 132 and 189 bears that consumed remote, tetracycline-laced baits in 2 different years, respectively, and observed 39 marks in 692 bone samples subsequently collected from hunters. We genetically analyzed hair samples from bait sites to determine the sex of marked bears, facilitating derivation of sex-specific population estimates. We obtained harvest samples from beyond the study area to correct for emigration. We estimated a density of 155 independent bears/100 km², which is equivalent to the highest recorded for this species. This high density appears to be maintained by abundant, accessible natural food. Our population estimate (approx. 1,000 bears) could be used as a baseline and to set hunting quotas. The refined biomarking method for abundance estimation is a useful alternative where physical captures or DNA-based estimates are precluded by cost or logistics. © 2011 The Wildlife Society.     

Temporal dynamics of grouse populations at the southern edge of their distribution

Studies of grouse conducted at northern latitudes have shown that tetraonids frequently exhibit cyclic fluctuations in abundance but little is known about the dynamics of grouse species at the southerly edge of their range. Hunting statistics from four species of grouse based on 30 yr of data collected from 210 hunting areas were examined from the Dolomitic Alps in the province of Trentino. These data were summed to represent 18 time series from discrete mountain groups. Analyses identified cycles of ca 5 yr in the minority of rock ptarmigan Lagopus mutus and hazel grouse Bonasa bonasia populations. These cycles only showed significant negative autocorrelation at half the cycle period and were classified as phase-forgetting quasi-cycles. Cycles were not found in time series of black grouse Tetrao terix or capereaillie Tetrao urogallus. Correcting time series for hunting effort or hunting restrictions tended to increase the proportion of populations that exhibited cycles but no difference in the strength of second order density dependence, A linear first order density-dependent autoregressive model described the dynamics of most of the populations with the exception of a proportion of rock ptarmigan and black grouse populations where a non linear first order model provided the best tit. We compare the findings with studies conducted in Finland and suggest possible reasons for the reduced tendency to cycle in the populations of southern Europe.     

Censusing rainforest game species with communal net hunts

This paper presents a method to address two wildlife management problems in central African rainforests: the need for local communities to take responsibility for wildlife management, and the lack of simple and appropriate wildlife monitoring techniques. The method uses encounters of game species during net hunts to calculate abundance indices as well as to estimate population densities for the four principal game species in the Dzanga–Sangha region: the duikers Cephalophus monticola (10.7–20.4 km^?2), C. dorsalis (1.2–2.0 km^?2), and C. callipygus (0.9–1.2 km^?2), and the brush-tailed porcupine Atherurus africanus (2.7–5.3 km^?2). Game species behaviour, the hunting practice, and comparisons with results from other research across central Africa suggest that the method can provide valid density estimates for C. monticola and C. dorsalis, but only abundance indices for C. callipygus and A. africanus. Nevertheless, the method can be applied by hunters in the course of their normal activities, and is adapted to the local habitat types and game species. As such, it can be an important tool for local communities in developing sustainable wildlife management programmes.     

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.     

Duck hunter activity,success, and satisfaction on public hunting areas

Because hunting disturbance can influence local distribution of ducks and their availability to hunters, managers often limit access to hunting areas to improve hunting success and satisfaction. Few studies have quantified the effectiveness of public area access restrictions on duck hunter activity, harvest, or hunters' satisfaction with their hunting experience. We used a cross-over design over 6 consecutive hunting seasons (2008–2009 through 2013–2014) on State Wildlife Areas (SWAs) in northeastern Colorado, USA, to compare the effects of restricted hunting access regulations and regulations without these restrictions on duck hunter activity, harvest success, harvest levels, and satisfaction. We also considered effects of SWA types, duck abundance, temperature, precipitation, use of equipment by duck hunting parties, and, for hunter satisfaction, hunting success, hunting parties' satisfaction with ducks seen, habitat conditions, crowding from other hunters, and SWA regulations. The number of days when duck hunters had access to restricted properties was about half that on unrestricted properties, and unrestricted properties were used by about twice as many duck hunting parties, but the mean number of hunting parties per available hunting day and mean party size were similar under the 2 types of regulations. Most (56%) duck hunting parties did not bag any ducks; hunting success (harvest of ≥1 duck by a hunting party) was best explained by a model that included a regulation type × hunting season interaction, a SWA type × month interaction, hunter density the previous day, an index of hunter investment (number of decoys used and whether dogs and calls were used), and temperature. Successful hunting parties harvested 1.92 ± 1.60 (SD) ducks/hunter/day (range = 0.1–7.0); the best model predicting the number of ducks harvested per hunter in successful parties included a regulation type × hunting season interaction, a SWA type × month interaction, hunter density the previous day, an index of hunter investment, temperature, and precipitation. Overall satisfaction of duck hunting parties with a day's hunt averaged 3.62 ± 1.20 based on a rank scale of 1 (very unsatisfied) to 5 (very satisfied), and was best explained by a model that included hunting party success; hunter investment; temperature; and satisfaction with duck numbers, habitat conditions, hunting regulations, and crowding. While greatly reducing days available for hunting, access restrictions were associated with improved chances of hunting parties successfully harvesting ≥1 duck in 5 of 6 hunting seasons, and substantially greater numbers of ducks harvested by hunters in successful parties in 1 of 6 hunting seasons. Restrictions did not have a strong direct effect on hunting parties' satisfaction with a day's hunt. Uncontrolled factors, including weather and use of equipment by hunters, had important influences on hunter success, harvest, and satisfaction. Managers should carefully assess hunting activity, hunter expectations and desires, and hunting area characteristics when considering access restrictions on public hunting areas.     

Patterns in grouse and Woodcock Scolopax rusticola hunting yields from central Norway 1901–24 do not support the alternative prey hypothesis for grouse cycles

According to the alternative prey hypothesis, autumn populations of ground-nesting game birds fluctuate in synchrony with vole numbers because generalist predators that mainly eat voles switch to alternative prey, such as eggs and chicks, when vole numbers decline. In hunting statistics from Nord-Trøndelag, central Norway, 1901–24, annual fluctuations in the number of Willow Grouse Lagopus lagopus and Western Capercaillie Tetrao urogallus , but not of Woodcock Scolopax rusticola , were positively related to vole numbers in the current year. Both Woodcock and grouse indices were related to hunting indices of Goshawk Accipiter gentilis and to weather variables assumed to influence the birds' survival or reproduction, suggesting that the indices actually reflected local population levels. Synchronous vole and grouse fluctuations are consistent with the alternative prey hypothesis (although predator densities were low in the early 1900s), but the asynchronous Woodcock fluctuations refute the hypothesis. Rather, because the Woodcock does not feed on plants utilized by voles and grouse, I suggest that food quality is the ultimate factor for the synchrony in vole and grouse numbers in Norway.     

Impact of unintentional selective harvesting on the population dynamics of red grouse

1.?The effect of selective exploitation of certain age, stage or sex classes (e.g., trophy hunting) on population dynamics is relatively well studied in fisheries and sexually dimorphic mammals. 2.?Harvesting of terrestrial species with no morphological differences visible between the different age and sex classes (monomorphic species) is usually assumed to be nonselective because monomorphicity makes intentionally selective harvesting pointless and impractical. But harvesting of the red grouse (Lagopus lagopus scoticus), a monomorphic species, was recently shown to be unintentionally selective. This study uses a sex- and age-specific model to explore the previously unresearched effects of unintentional harvesting selectivity. 3.?We examine the effects of selectivity on red grouse dynamics by considering models with and without selectivity. Our models include territoriality and parasitism, two mechanisms known to be important for grouse dynamics. 4.?We show that the unintentional selectivity of harvesting that occurs in red grouse decreases population yield compared with unselective harvesting at high harvest rates. Selectivity also dramatically increases extinction risk at high harvest rates. 5.?Selective harvesting strengthens the 3- to 13-year red grouse population cycle, suggesting that the selectivity of harvesting is a previously unappreciated factor contributing to the cycle. 6.?The additional extinction risk introduced by harvesting selectivity provides a quantitative justification for typically implemented 20-40% harvest rates, which are below the maximum sustainable yield that could be taken, given the observed population growth rates of red grouse. 7.?This study shows the possible broad importance of investigating in future research whether unintentionally selective harvesting occurs on other species.