Retrospective comparisons of competing demographic models give clarity from "messy" management on a Scottish grouse moor.

Retrospective comparison of predictive models that describe competing hypotheses regarding system function can shed light on regulatory mechanisms within the framework of adaptive resource management. We applied this approach to a 28-year study of red grouse (Lagopus lagopus scotica) in Scotland, with the aims of reducing uncertainty regarding important drivers of grouse population dynamics, and of evaluating the efficacy of using seasonal versus annual model assessments. We developed three sets of models that predicted pre-breeding and post-breeding grouse density, matching the timing of grouse counts on the ground. We updated conditions and management through time in the spirit of a real-time, adaptive management program and used a Bayesian model weight updating process to compare model predictions with empirical grouse densities. The first two model sets involved single annual updates from either pre-breeding or post-breeding counts; the third set was updated twice a year. Each model set comprised seven models representing increasingly complex hypotheses regarding potentially important drivers of grouse: the baseline model included weather and parasite effects on productivity, shooting losses and density-dependent overwinter survival; subsequent models incorporated the effect of habitat gain/loss (HAB), control of non-protected predators (NPP) and predation by protected hen harriers (Circus cyaneus, HH) and buzzards (Buteo buteo, BZ). The weight of evidence was consistent across model sets, settling within 10 years on the harrier (NPP + HH), buzzard (NPP + HH + BZ) and buzzard + habitat (NPP + HH + BZ + HAB) models, and downgrading the baseline + habitat, non-protected predator, and non-protected predator + habitat models. By the end of the study only the buzzard and buzzard + habitat models retained substantial weights, emphasizing the dynamical complexity of the system. Habitat inclusion failed to improve model predictions, implying that over the period of this study habitat quantity was unimportant in determining grouse abundance. Comparing annually and biannually assessed model sets, the main difference was in the baseline model, whose weight increased or remained stable when assessed annually, but collapsed when assessed biannually. Our adaptive modeling approach is suitable for many ecological situations in which a complex interplay of factors makes experimental manipulation difficult.

Measures of predator diet alone may underestimate the collective impact on prey: Common buzzard Buteo buteo consumption of economically important red grouse Lagopus lagopus scotica.

Human-wildlife conflicts often centre on economic loss caused by wildlife. Yet despite being a major issue for land-managers, estimating total prey losses to predation can be difficult. Estimating impacts of protected wildlife on economically important prey can also help management decisions to be evidence-led. The recovery in population and range of common buzzards Buteo buteo in Britain has brought them into conflict with some gamebird interests. However, the magnitude of any impact is poorly understood. We used bioenergetics models that combine measures of buzzard abundance from field surveys with diets assessed by using cameras at nests, prey remains and pellet analysis, to estimate their impact on red grouse Lagopus lagopus scotica on a large (115 km2) moor managed for red grouse shooting in Scotland. Whilst grouse consumption by individual buzzards was lower than previous estimates for other raptor species present on our study site, total consumption could be greater given an estimated 55-73 buzzards were present on the study site year-round. Averaging across diet assessment methods, consumption models estimated that during each of three breeding seasons (April-July 2011-2013), the buzzards foraging on our study site consumed 73-141 adult grouse and 77-185 chicks (depending on year). This represented 5-11% of adult grouse present in April (22-67% of estimated adult mortality) and 2-5% of chicks that hatched (3-9% of estimated chick mortality). During two non-breeding seasons (August-March), consumption models using pellet analysis estimated that buzzards ate a total of 242-400 grouse, equivalent to 7-11% of those present at the start of August and 14-33% of estimated grouse mortality during the non-breeding season. Buzzard consumption of grouse has the potential to lead to non-trivial economic loss to grouse managers, but only if buzzards predated the grouse they ate, and if grouse mortality is additive to other causes.

Resource-Area-Dependence Analysis: Inferring animal resource needs from home-range and mapping data.

An animal's home-range can be expected to encompass the resources it requires for surviving or reproducing. Thus, animals inhabiting a heterogeneous landscape, where resource patches vary in size, shape and distribution, will naturally have home-ranges of varied sizes, so that each home-range encompasses a minimum required amount of a resource. Home-range size can be estimated from telemetry data, and often key resources, or proxies for them such as the areas of important habitat types, can be mapped. We propose a new method, Resource-Area-Dependence Analysis (RADA), which uses a sample of tracked animals and a categorical map to i) infer in which map categories important resources are accessible, ii) within which home range cores they are found, and iii) estimate the mean minimum areas of these map categories required for such resource provision. We provide three examples of applying RADA to datasets of radio-tracked animals from southern England: 15 red squirrels Sciurus vulgaris, 17 gray squirrels S. carolinensis and 114 common buzzards Buteo buteo. The analyses showed that each red squirrel required a mean (95% CL) of 0.48 ha (0.24--0.97) of pine wood within the outermost home-range, each gray squirrel needed 0.34 ha (0.11-1.12) ha of mature deciduous woodland and 0.035-0.046 ha of wheat, also within the outermost home-range, while each buzzard required 0.54 ha (0.35-0.82) of rough ground close to the home-range center and 14 ha (11-17) of meadow within an intermediate core, with 52% of them also relying on 0.41 ha (0.29-0.59) of suburban land near the home-range center. RADA thus provides a useful tool to infer key animal resource requirements during studies of animal movement and habitat use.

Influences of extreme weather, climate and pesticide use on invertebrates in cereal fields over 42 years.

Cereal fields are central to balancing food production and environmental health in the face of climate change. Within them, invertebrates provide key ecosystem services. Using 42 years of monitoring data collected in southern England, we investigated the sensitivity and resilience of invertebrates in cereal fields to extreme weather events and examined the effect of long-term changes in temperature, rainfall and pesticide use on invertebrate abundance. Of the 26 invertebrate groups examined, eleven proved sensitive to extreme weather events. Average abundance increased in hot/dry years and decreased in cold/wet years for Araneae, Cicadellidae, adult Heteroptera, Thysanoptera, Braconidae, Enicmus and Lathridiidae. The average abundance of Delphacidae, Cryptophagidae and Mycetophilidae increased in both hot/dry and cold/wet years relative to other years. The abundance of all 10 groups usually returned to their long-term trend within a year after the extreme event. For five of them, sensitivity to cold/wet events was lowest (translating into higher abundances) at locations with a westerly aspect. Some long-term trends in invertebrate abundance correlated with temperature and rainfall, indicating that climate change may affect them. However, pesticide use was more important in explaining the trends, suggesting that reduced pesticide use would mitigate the effects of climate change.

Factors associated with shooting accuracy and wounding rate of four managed wild deer species in the UK, based on anonymous field records from deer stalkers.

The amount of wounding during routine culling is an important factor in the welfare of wild deer. Little information exists on factors determining shooting accuracy and wounding rates under field conditions in the UK. In this study, 102 anonymous stalkers collected data on the outcomes and circumstances of 2281 shots. Using hot-deck imputation and generalised linear mixed modelling, we related the probability that a shot hit its target, and the probability that the shot killed the deer if it was hit, to 28 variables describing the circumstances of the shot. Overall, 96% of deer were hit, of which 93% were killed outright. A reduced probability of hitting the target was associated with an uncomfortable firing position, too little time available, shooting off elbows or freehand, taking the head or upper neck as point of aim, a heavily obscured target, a distant target, shooting at females, lack of shooting practice and a basic (or no) stalker qualification. An increase in the likelihood of wounding was associated with an uncomfortable firing position, shooting with insufficient time, a distant target (only when time was not sufficient), a bullet weight below 75 grains, a target concealed in thicket or on the move and an area rarely stalked. To maximise stalking success and deer welfare, we recommend that stalkers ensure a comfortable firing position, use a gun rest, aim at the chest, use bullets heavier than 75 grains, avoid taking a rushed shot, shoot a distant animal only if there is plenty of time, fire only when the target is stationary, avoid shooting at an obscured animal, take care when the ground is unfamiliar, and do shooting practice at least once a month. The high miss rate of basic-level stalkers suggests that training should include additional firing practice under realistic shooting conditions.

Vigilance and fitness in grey partridges Perdix perdix: the effects of group size and foraging-vigilance trade-offs on predation mortality.

1. Vigilance increases fitness by improving predator detection but at the expense of increasing starvation risk. We related variation in vigilance among 122 radio-tagged overwintering grey partridges Perdix perdix (L.) across 20 independent farmland sites in England to predation risk (sparrowhawk Accipiter nisus L., kill rate), use of alternative antipredation behaviours (grouping and use of cover) and survival. 2. Vigilance was significantly higher when individuals fed in smaller groups and in taller vegetation. In the covey period (in early winter when partridges are in flocks), vigilance and use of taller vegetation was significantly higher at sites with higher sparrowhawk predation risk, but tall vegetation was used less by larger groups. Individuals were constrained in reducing individual vigilance by group size and habitat choice because maximum group size was determined by overall density in the area during the covey period and by the formation of pairs at the end of the winter (pair period), when there was also a significant twofold increase in the use of tall cover. 3. Over the whole winter individual survival was higher in larger groups and was lower in the pair period. However, when controlling for group size, mean survival decreased as vigilance increased in the covey period. This result, along with vigilance being higher at sites with increasing with raptor risk, suggests individual vigilance increases arose to reduce short-term predation risk from raptors but led to long-term fitness decreases probably because high individual vigilance increased starvation risk or indicated longer exposure to predation. The effect of raptors on survival was less when there were large groups in open habitats, where individual partridges can probably both detect predators and feed efficiently. 4. Our study suggests that increasing partridge density and modifying habitat to remove the need for high individual vigilance may decrease partridge mortality. It demonstrates the general principle that antipredation behaviours may reduce fitness long-term via their effects on the starvation-predation risk trade-off, even though they decrease predation risk short-term, and that it may be ecological constraints, such as poor habitat (that lead to an antipredation behaviour compromising foraging), that cause mortality, rather than the proximate effect of an antipredation behaviour such as vigilance.