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.     

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.     

Evaluating spatially explicit viability of a declining ruffed grouse population

Species associated with early successional habitats have experienced dramatic declines in the eastern United States as a result of land use changes and human disruption of natural disturbance regimes. Consequently, active management is required to create early successional habitat and promote plant and animal communities that depend on periodic forest disturbance. Ruffed grouse (Bonasa umbellus) depend on recently disturbed forest habitat, and have experienced dramatic declines over the last half-century. Although ruffed grouse are extensively studied, little effort has been made to link population dynamics with habitat management at landscape scales. We used stochastic, spatially explicit population models that combined landscape conditions derived from a Geographic Information System with demographic data, and applied the model to a declining ruffed grouse population in Rhode Island, USA. We identified vital rates that influence ruffed grouse population dynamics using baseline models constructed with current demographic rates and landscape conditions, and assessed the effect of landscape-scale forest management alternatives on population persistence by running multiple management simulations. Baseline models typically predicted population decline, and we concluded that vital rates (survival and recruitment) had a greater influence on population persistence than did dispersal capability, carrying capacity, or initial population size. Management simulations predicted greater population persistence under a scenario where high-quality habitat was provided in fewer large blocks as opposed to many small blocks, and the rate at which we allowed ruffed grouse to colonize newly created habitat had a substantial impact on management success. Populations of ruffed grouse in the eastern United States are likely to continue to decline given current disturbance regimes, and our work provides a link between ruffed grouse demography and landscape-scale habitat conditions to support management decisions. © 2011 The Wildlife Society.     

Myoglobin levels and mATPase activity in pectoral muscles of spruce and ruffed grouse (Aves: Tetraoninae)

Myoglobin concentration and myosin ATPase activity were measured in the pectoral muscle of wild spruce grouse (Dendragapus canadensis) and ruffed grouse (Bonasa umbellus), together with the weight of the Mm. pectoralis, supracoracoideus and heart. mATPase activities were similar in both species, but spruce grouse contained 15 times more myoglobin in the pectoralis muscle and the heart was three times heavier than that of the ruffed grouse. The relative mass of the flight muscles and wing loading were similar between species. Characteristics of the pectoral muscle of both grouse species reflect adaptations to predation and advertising displays. The glycolytic nature of the ruffed grouse pectoral muscle and small heart size is an adaptation to a sedentary existence within a small home range. The more oxidative pectoral muscle of spruce grouse together with its larger heart are adaptations to seasonal dispersals requiring more sustained flight.     

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.     

Ruffed Grouse Brood Habitat Use in Mixed Softwood-Hardwood Nordic-Temperate Forests,Quebec, Canada

Abstract: Adequate cover is a critical component of ruffed grouse (Bonasa umbellus) habitat during the brood-rearing period when chick mortality is high. We assessed habitat use by ruffed grouse during the brood-rearing period by comparing characteristics of tree, shrub, and ground layers at ruffed grouse brood and random locations. We captured and radiomarked 29 females with broods in 2 forest settings of the Réserve faunique de Portneuf, Quebec, Canada. We described grouse habitat using ground surveys and forest maps, and we identified the used habitat characteristics using analysis of variance and logistic regression. Females with broods used mixed and regenerated clearcut stands that were 1.5–7 m tall and 11–20 years old. Compared with random locations, grouse locations had higher lateral obstruction (76% vs. 68%), higher small-stem density (29,085 stems/ha vs. 19,340 stems/ha), and were closer to roads and trails. Percentage of coverage by ground vegetation was not higher at grouse locations as often reported in previous studies. Results from this study will help orient ruffed grouse habitat management on Quebec public land and elsewhere in nordic—temperate mixed hardwood—softwood forests to maintain suitable brood habitat after logging operations. Forest management should promote growth of young mixed stands with high horizontal and vertical cover provided by high small-stem density, which offers protection against aerial and terrestrial predation. Edges such as roadsides are also important in brood habitat as they provide food and cover.     

Territorial status and survival in red grouse Lagopus lagopus scoticus: hope for the doomed surplus?

A previous study of survival in territorial and non-territorial red grouse Lagopus lagopus scoticus conducted between 1957 and 1967 found that territorial status in the autumn pre-determined over-winter survival. A very high proportion of territorial birds survived and virtually all non-territorial birds died or emigrated. We tested the hypothesis that over-winter survival was dependent on territorial status within four grouse populations in Scotland between 1986 and 1993. In contrast to the previous study, 66% of non-territorial birds survived over winter compared to approximately 70% of territorial birds. There was no significant effect of territorial status on the survival estimates. Moreover, some of the birds considered to be non-territorial during autumn went on to successfully raise a brood. We suggest that on our study sites, territory ownership in autumn did not greatly influence over-winter survival, and territorial behaviour did not determine breeding density as previously supposed. We postulate differences with other studies may reflect variations in scale and predation pressure.     

Parents, predators, parasites, and the evolution of eggshell colour in open nesting birds

The colourful surface of birds’ eggshells varies dramatically between species, but the selective pressures driving this variation remain poorly understood. We used a large comparative dataset to test several hypotheses proposed to explain the evolution of eggshell colouration. We tested the hypothesis that predation pressure might select for cryptic eggshells by examining the relationship between predation rate and egg colouration. We found that predation rates were positively related to eggshell brightness. The blackmail hypothesis suggests that females lay colourful eggshells to coerce males into providing additional care during incubation to keep colourful eggs covered. According to this hypothesis, conspicuous eggs should be found in situations with high risk of visual detection from predators or brood parasites. In support of this hypothesis, proportional blue-green chroma was positively related to parasitism risk, and eggs with higher proportional blue-green chroma or higher ultraviolet chroma received higher combined parental nest attendance during the incubation period. The sexual signalling hypothesis states that blue-green colour indicates female quality; however, we did not find that blue-green eggshell colour was greater in species where males participate in any form of parental care, and relative male provisioning was unrelated to blue-green eggshell chroma. We found some support for the hypothesis that brood parasitism may select for high inter-clutch variation in eggshell colour to facilitate egg recognition. In our dataset, parasitism risk was negatively related to inter-clutch repeatability of blue-green chroma. Our study highlights the diversity of selection pressures acting on the evolution of eggshell colour in birds and provides suggestions for novel areas of future key research direction.     

Occupancy modeling of ruffed grouse in the Black Hills National Forest

Ruffed grouse (Bonasa umbellus) are a popular game bird and the management indicator species for quaking aspen (Populus tremuloides) in the Black Hills National Forest (BHNF), which requires development of a robust monitoring protocol to evaluate trends in ruffed grouse populations. We used roadside drumming surveys in spring 2007 and 2008 to estimate ruffed grouse occupancy and detection probabilities in the BHNF while simultaneously assessing the influence of sampling and site covariates on these processes. Ruffed grouse occupancy estimates were constant between spring 2007 and 2008 (Ψ = 0.12, SE = 0.03) and were positively influenced by the amount of aspen surrounding the site. Detection probability estimates were constant between spring 2007 and 2008 (p = 0.27, SE = 0.06) and were influenced by survey date in a quadratic form and negatively influenced by wind speed and time of the survey. Collectively, our results demonstrated that ruffed grouse occupancy and detection probabilities in the BHNF were low. Occupancy could be increased by increasing the extent of aspen. To improve monitoring efficiency and maximize probability of detecting ruffed grouse, ruffed grouse monitoring should be conducted during the peak of drumming (mid-May), during favorable weather conditions such as low wind speeds and little precipitation, and during early morning, near sunrise. © 2010 The Wildlife Society.     

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.     

Nest predation and the evolution of nestling begging calls

Begging by nestling birds can be conspicuous and loud. Such displays are thought to function in signalling nestling condition and securing parental care, but they also may inadvertently attract the attention of predators. We compared the structure of nestling begging calls to the risk of predation among 24 species of birds breeding in a forest community in central Arizona. After controlling for body size and phylogeny, we found that species subject to greater nest predation had calls with higher frequency (pitch) and lower amplitude (loudness) than species subject to lower rates of nest predation. As these acoustic features make it difficult for potential predators to pinpoint the source of a sound, our results suggest that an increased risk of predation has led to the evolution of begging calls that minimize locatability. The relationship between call structure and the risk of predation also supports the hypothesis that attracting predators is a direct cost of begging and that such costs can constrain any evolutionary escalation in the intensity of nestling begging.     

Risk taking by singing males

The distance at which an individual flees from a potential predatorrepresents a measure of risk taking. If individuals are engagedin another activity that might affect fitness, trade-offs betweenthe fitness benefits of flight and the other activity shoulddetermine the nearest distance of approach by a predator. Ina comparative analysis of birds, flight distance representeda reliable measure of risk of predation by the sparrowhawk Accipiternisus that increased with decreasing flight distance acrossspecies. To test the hypothesis that singing males adjustedtheir risk taking to the costs and benefits of early flight,we compared the flight distance of singing and nonsinging birdsto an approaching human observing with a binocular. Singingbirds on average fled at a greater distance than nonsingingbirds, implying that singing birds took small risks. We useda standardized measure of difference in flight distance betweensinging and nonsinging individuals to investigate factors affectinginterspecific variation in risk taking. Species that used moreexposed song posts (sites used for singing) took smaller risksthan species with less exposed song posts. Species that sufferedfrom higher levels of parasitism as reflected by the prevalenceof Plasmodium, but not by 3 other genera of blood parasites,took greater risks during singing compared with nonsinging activities.Likewise, species with high circulating levels of natural antibodies,and hence a history of natural selection caused by bacteriatook relatively greater risks during singing than species withfew natural antibodies. These findings suggest that risks takenby singing birds have been molded by natural and sexual selection,and that risk taking represents a compromise between the costsand benefits of flight from a potential predator.     

Increased nest predation near protected capercaillie leks: a caveat against small reserves

Protecting animal aggregation sites is intuitively an efficient conservation approach, particularly for threatened species in fragmented landscapes. However, the appropriate scale of protection depends on accompanying threats, such as predator attraction to the same aggregation sites, to which the target species may need to respond by moving to alternative habitat patches. We performed experiments using artificial ground nests around protected capercaillie leks in commercially managed forest landscape in Estonia. We considered two scales: the landscape up to 3 km from the lek centre, and among 10–30-ha forest compartments, which differed in predator abundance. We found that nest predation significantly declined with distance from the lek. Nests were depredated by multiple mammalian and avian predator species and total predator abundance explained most of the between-forest variation in predation rate. Our results indicate that, in this hemiboreal area, (i) ground nest predation is largely determined by landscape-scale distribution of predators and (ii) predators can aggregate at capercaillie leks. The implication is that lek-centred habitat protection used for the capercaillie and other grouse may be ineffective unless the peripheral habitat quality and predator abundance are specifically addressed. More generally, predation pressure can be a serious problem in small set-asides within hostile landscapes for threatened ground-nesting birds.     

Response to Predation Risk in Urban and Rural House Sparrows

Habitat urbanization may change the density of predators, and it is often assumed that such changes lead to altered predation risk for urban populations of their prey. Although it is difficult to study predation hazard directly, behavior responses of prey species may be informative in inferring such habitat differences. In this study, we compared the risk‐taking behavior of urban and rural house sparrows (Passer domesticus) after simulated attacks by two of their important predators (sparrowhawk Accipiter nisus and domestic cat Felis catus). The birds were startled by moving dummies of these predators and respective control objects, and their risk taking was estimated as their latency to feed after the startle. We found that sparrows responded more strongly (had longer post‐startle feeding latencies) to sparrowhawk attacks than to the control object, and their responses differed between the habitats. First, risk taking of urban birds strongly decreased with age (older birds had longer latencies than young birds), while there was no such age difference in rural birds. Second, young urban birds responded less strongly, while older urban birds responded more strongly to the sparrowhawk than the same age groups of rural birds, respectively. We did not succeed in evoking antipredatory response by simulated cat attacks, because birds responded similarly to the dummy and the control object. Our results support that predation risk, posed at least by avian predators, is different in urban and rural habitats of house sparrows. The increased wariness of older, hence presumably more experienced, urban birds implies that sparrows may be more exposed to predation in cities.     

Compensation for predator-induced reduction in nestling feeding rate in the Great Spotted Woodpecker

Parent birds make efforts to prevent the immediate costs of predation through plastic behavioural responses to the actual predation risk, but this may incur future costs for offspring due to reduced parental care. However, the temporary nature of predator encounters suggests that nestling feeding reduced during the risky periods may be later compensated for by an increased feeding effort (the predation risk allocation hypothesis). We tested this prediction in the Great Spotted Woodpecker (Dendrocopos major) confronted with its major nest predator/competitor, the European Starling (Sturnus vulgaris). A brief encounter with a live starling was followed by a reduction in the nestling feeding rate, but the lost feedings were subsequently compensated for by an increased feeding rate. This compensatory effect was higher in older nestlings that are highly demanding in terms of energy requirements and fitness value from the parents’ perspective. Thus, birds are potentially able to respond not only to the immediate risk of nest predation by nest defence but also to compensate for the potential costs of nest defence in terms of unfulfilled nestling demands. However, data on the amount of delivered food are necessary to distinguish whether parents truly compensate for lost feeding or whether the increased feeding frequency represents a nest guarding strategy.     

Evaluation of Potential Factors Predisposing Livestock to Predation by Jaguars

ABSTRACT Depredation of livestock by large carnivores is an important but poorly understood source of human-carnivore conflict. We examined patterns of livestock depredation by jaguars (Panthera onca) and pumas (Puma concolor) on a ranch-wildlife reserve in western Brazil to assess factors contributing to prey mortality. We predicted jaguars would kill a greater proportion of calves than yearling and adult cattle and that proximity to suitable habitat would increase mortality risk. We further speculated that exposure to predation risk would promote livestock grouping and increased movement distance. We recorded 169 cattle mortality incidents during 2003–2004, of which 19% were due to predation by jaguars and pumas. This level of mortality represented 0.2–0.3% of the total livestock holdings on the ranch. Jaguars caused most (69%) cattle predation events, and survival in allotments was lower for calves than for other age classes. Forest proximity was the only variable we found to explain patterns of livestock mortality, with predation risk increasing as distance to forest cover declined. Due to low predation risk, cattle movement patterns and grouping behavior did not vary relative to level of spatial overlap with radiocollared jaguars. The overall effect of predation on cattle was low and livestock likely constituted an alternative prey for large cats in our study area. However, selection of calves over other age cohorts and higher predation risk among cattle in proximity to forest cover is suggestive of selection of substandard individuals. Cattle ranchers in the Pantanal region may reduce cattle mortality rates by concentrating on losses due to nonpredation causes that could be more easily controlled.     

Social context affects risk taking by a satellite species in a mixed-species foraging group

Mixed-species flocks of birds form during winter in the easterndeciduous forests of North America. These flocks consist oftwo flock-leading nuclear species, tufted titmouse (Baeolophusbicolor) and Carolina chickadee (Poecile carolinensis), andseveral follower, or satellite, species, including downy woodpecker(Picoides pubescens) and white-breasted nuthatch (Sitta carolinensis).Hypotheses explaining the adaptiveness of participation in suchmixed-species foraging groups have focused on increased foragingsuccess and/or decreased predation risk. We tested the predictionthat if nuthatches join nuclear species to reduce predationrisk, they should be more reluctant to visit an exposed feederin the absence of titmice than in their presence. When the feederwas positioned 16 m from forest cover, latency to visit thefeeder was greater for both male and female nuthatches whentitmice were absent. Removal of titmice had no effect on latencyat 8 m. In the absence of titmice, nuthatches visited the feederless frequently at both distances. These results indicate thatreduced predation risk is a benefit that satellite species gainby flocking with nuclear species.     

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.