A major task of the Grouse Specialist Group is to compile an IUCN/SSC Action Plan to assess threats and propose conservation actions for all grouse species worldwide. This paper provides an overview to the conservation status of and major threats to grouse based on information collected during the preparation of the Grouse Action Plan. Three species, Caucasian black grouse Tetrao mlokosiewiczi, Chinese grouse Bonasa sewerzowi and Siberian grouse Falcipennis falcipennis, are listed as globally near-threatened or data deficient, one newly-recognised species, Gunnison sage grouse Centrocercus minimus, is listed as globally endangered and two subspecies, Attwater's prairie chicken Tympanuchus cupido attwateri and Cantabrian capercaillie Tetrao urogallus cantabricus, qualify to be listed as globally threatened according to IUCN criteria. At a national level, 14 of the 18 species are red-listed in at least one country. Populations at the southern edge of a species' range and in densely populated regions are most often red-listed. Based on questionnaire results from 38 countries, habitat degradation, loss and fragmentation due to human land use activities are the major threats to grouse viability. Exploitation, predation and human disturbance were regionally felt to be critical. Major threats and their causes are reviewed based on questionnaire results and the literature. Integrating habitat preservation and human land use practices is concluded to be the major challenge to grouse conservationists worldwide.

We propose a new classification of the grouse that brings their taxonomy into agreement with our molecular phylogenetic studies. Our analyses provide, for the first time, a robust estimate of the evolutionary history of these birds. These analyses are based on aligned sequences of 3,809 basepairs of five complete mitochondrial genes. Our classification does not require novel genera and generally results in the maintenance of accepted generic names. Only two mono- typic genera are required. We recognize the grouse as a subfamily, Tetraoninae, within the family Phasianidae. We recognize three tribes; these include a tribe (Bonasini, a new taxon) for the ruffed grouse Bonasa umbellus, a tribe (Tetrastini, a new rank) for hazel hens in the genus Tetrastes, and a tribe (Tetraonini, a new rank) for all the remaining species. We divide this last, derived tribe into five subtribes that correspond to 1) Falcipennina (a new taxon) for the sharp-winged grouse Falcipennis falcipennis, 2) Canachitina (a new taxon) for the New World spruce grouse in the genus Canachites, 3) Tetraonina (a new rank) for the capercaillies and black grouse in the genera Tetrao and Lyrurus, respectively, 4) Centrocercina (a new taxon) for the New World prairie and forest grouse in the genera Tympanuchus, Centrocercus and Dendragapus, and 5) Lagopodina (a new taxon) for the ptarmigans in the genus Lagopus. All the taxa in our classification, at all ranks, are monophyletic with bootstrap support of 95% or more.

The long-term temporal dynamics of four grouse species in the Italian Alps have been investigated in the attempt to reveal if the populations exhibit the tendency to cycle at the southern edge of their European range. Hunting statistics or count data were collected as total number of individuals shot or counted each year from five provinces. For the province of Trento data were available at the level of mountain groups and a more detailed investigation was carried out. The results from Trento were compared with the more general findings from the other four Italian provinces. Time series analysis was performed to investigate the pattern of cycle and autoregressive models were used to describe the density dependence structure. In general, the populations showed a weak or no tendency to regular fluctuations: rock ptarmigan Lagopus mutus and black grouse Tetrao tetrix populations exhibited the highest tendency to cycle with periods of 5–9 years while capercaillie Tetrao urogallus never showed regular fluctuations. Hazel grouse Bonasa bonasia cyclic dynamics were restricted to a few populations of Trento. When time series from Trento were corrected for hunting effort the detection of cycle among populations sharply increased but no differences in the strength of second order density dependence or in the period length was observed. A linear first order autoregressive model explained better the intrinsic structure of the majority of populations. We compare the findings with studies conducted on populations of northern Europe and suggest possible reasons for the reduced tendency to cycle in the grouse populations of the Italian Alps.

In order to estimate minimum area requirements for viable isolated populations of capercaillie Tetrao urogallus in Central Europe, we developed a stochastic population model. Model parameters were largely based on the results of a field study in the Bavarian Alps. Environmental fluctuations (amount of rainfall in June and July) are taken into account in the early survival of chicks. For the default parameter set, the model predicts a minimum capacity needed for viability, i.e. for the extinction risk not to exceed 1% in 100 years, of about 470 individuals. In the Bavarian Alps, this corresponds to area requirements of an isolated viable population in an order of magnitude of 250 km². These results are, however, sensitive to small changes in model parameters such as female survival, clutch survival and chick survival. We conclude that minimum viable population size and minimum area requirements may vary considerably across the range of the capercaillie. Therefore, we plan to apply our model to different parameter sets from different regions in order to explore the range of conditions under which capercaillie populations may be viable.

Although sage grouse Centrocercus urophasianus have declined throughout their range in North America, little is known about annual mortality patterns of this species. Thus, we summarize a long-term data set on timing and causes of mortality of sage grouse. Predation was the most common cause of death for radio-marked sage grouse. For adult males, 83% of deaths were attributed to predation and 15% to hunting. However, for adult females, 52% of deaths were caused by predation while 42% were attributed to hunting. We rejected the hypothesis that type of mortality (predation vs hunting) was independent of gender of sage grouse. For males, 70% of deaths occurred during spring and summer (March-August) and 28% occurred in September-October. For females, 52% of mortalities occurred during spring and summer and 46% occurred in September-October. We rejected the hypothesis that time of death is independent of the gender of sage grouse. In six of 15 years (40%), harvest rates for adult females may have exceeded 10% while this rate was only exceeded in two of 15 years (13%) for adult males.

We surveyed Caucasian black grouse Tetrao mlokosiewiczi residing on the Lagonakskiy Ridge, northwestern Caucasus, Russia, in June-July 1998, and on the Magisho Ridge in July 1999. Weather affected habitat use by brood hens: on sunny days 18 broods were encountered in meadows, six in ravines, and three in pine forests; on wet days broods moved to the ridge tops (N = 9) and only one was observed in pine forest. Subadult males used the same habitats as females with broods. Seven of eight adult males were encountered in ravines; only one was encountered in a meadow. Habitat use by adult males was not affected by weather and differed from habitat use by females and subadult males. Well-camouflaged females, chicks and subadult males used relatively open, food-rich habitats, whereas black adult males preferred ravines, where nutrition was poor, but where tall grass protected them from aerial predators. One nest with five hatched eggs was found. In broods an average of one chick per 10 days was lost. A goshawk Accipiter gentilis killed one adult female. Our density estimate of 2.3 adults/km² for the Lagonakskiy Ridge was similar to densities reported elsewhere (2.3 ± 1.2; N = 7).

Radio-equipped adult capercaillie Tetrao urogallus males (N = 10) were tracked during April and May 1996–1997 in a coniferous forest in southeastern Norway, which has been subjected to clear-felling and replanting during the last five decades. With the use of radio-telemetry daytime habitat preference was investigated at two levels of scale: within home range and within forest stands inside home ranges. In addition, roosting habitat preference within stands was investigated by locating heaps of intestinal faeces, henceforth called roosting sites. Within forest stands, forest structure and visibility were measured at roosting sites (N = 94) and at random plots (N = 188). Within home ranges the birds used plantations and old naturally regenerated forest according to availability, and open bogs were avoided. Of the roosting sites, 90% were located underneath the low branches of Norway spruce Picea abies trees. For roosting within old naturally regenerated stands (> 70 years old) males preferred sites with higher tree density, higher vertical and horizontal cover and lower visibility than in the surrounding forest. In young and middle-aged plantations, on the contrary, roosting sites tended to have lower tree density (P = 0.118), but still seemed to have better cover than the surrounding forest (P = 0.117). The forest structure preference of roosting sites was reflected in the use of vegetation types within the old naturally regenerated forest. Because dense cover reduces the probability that the birds are detected by a predator, but at the same time increases the risk of being killed once detected, capercaillie males have to compromise between shelter and outlook. Our data indicate that males prefer good cover at the expense of good overview of the surroundings when selecting roosting sites, and that tree density is usually too dense in younger plantations, probably because outlook is reduced and flying obstructed.

The ‘landscape mosaic’ hypothesis predicts that adult male capercaillie Tetrao urogallus track available habitat when making distinct seasonal migrations. We tested this hypothesis by comparing movement from leks to summer range in three geographical regions, viz. northwest Russia (Pechora), southwest Russia (Tver) and southeast Norway (Varaldskogen). All radio-collared males dispersed out of their daytime lek territories to an average distance of 2.3 km (±0.37) from lek centres (N = 52). In Russia, three birds, which were not included in this estimate, moved beyond the 6–7 km detection distance, but returned the following year. Due to large variation among birds at each lek, the only significant difference in movement was found between the birds at the lek in Pechora ( = 1.3 km) and those at one of the two leks in Tver ( = 3.6 km) and at one of the three leks in Norway ( = 3.2 km). At all leks except the one in Tver, the amount of suitable summer habitat, e.g. old Norway spruce Picea abies forest on rich soil, was smaller within a 1–km radius of leks centres than in the outer two 1–km zones. Despite a large proportion of old spruce forest at and near the lek site in Tver, the males at this site moved >2 km from the lek centre in late May/early June. When data were pooled, the dispersal distance did not correlate with the amount of old spruce forest nor with the size of patches (e.g. grain size) with increasing distance from leks. However, at the Norwegian study area, which is highly fragmented with a fine-grained habitat mosaic due to commercial forestry, variation in topography and soil quality, birds tended to move farther away from the leks with increasing patch size and with increasing proportion of old spruce forest. When fitting a second polynomial regression function to the pooled data, there was only a weak relationship between dispersal distance and grain size for leks (P = 0.19). When including all birds, the relationship was highly significant (P = 0.002). The Pechora lek in northwest Russia contributed mainly to the lack of a clear relationship between movement pattern and distribution of old spruce forest. Here large, contiguous bands of old spruce forest were located along the main river courses in a coarse-grained fashion within 2 km of lek sites. Thus, local configuration of preferred summer habitat at the landscape scale probably affects both direction and movement to a larger extent than predicted from the ‘landscape mosaic’ hypothesis.

The prairie sharp-tailed grouse Tympanuchus phasianellus campestris occurs throughout the north central region of North America. It is of management concern because it has decreased in the southeast portion of its range over the past three decades, including marked declines in Minnesota and the Great Lakes region, USA. Although there is general knowledge about the habitat requirements for this species, no quantitative lek site or landscape information has been documented. We quantified landscape composition around active and inactive sharp-tailed grouse lek sites and random points in brush landscapes in northeast Minnesota at multiple scales (200–3,000 m radii circles). Our objective was to compare landscape composition among these sites. We also developed a model to predict the probability of grouse lek site occurrence in the study area. Landscape composition around active and inactive lek sites differed from each other primarily at the 500 m and 1,000 m radii scales. Inactive sites had higher proportions of upland forest and brush cover types and active sites had a higher percentage of native grass than inactive sites. No differences were found in landscape composition between site types at the 200 m radius scale and only one landscape variable (number of cover types) was different at the 3,000 m radius scale. We found non-random distributions of this grouse species at four different scales. Random brush land sites differed from both active and inactive sites having higher percentages of forest and brush cover. In contrast, lek sites had more bare ground, emergent aquatic vegetation, bog brush and roads than the random points. A regression model for the grouse at the 3,000 m scale was used to predict the probability of grouse occurrence in the landscape. The model resulted in a spatial map with about 8% of the area having a probability of grouse occurrence of >80%. This information can be used to locate new lek sites and to guide management activities for this species.

The sage grouse Centrocercus urophasianus population level on the Upper Snake River Plain of Idaho has declined significantly over the past 40 years. We investigated migration patterns and seasonal ranges of these birds to compare to patterns from the 1950s and 1960s. Furthermore, we examined landscape changes that occurred between 1975 and 1992. Migration patterns have not changed since the 1950s. The grouse currently migrate up to 125 km and use an annual population range of at least 2,764 km². The major landscape change since 1975 that occurred in sage grouse habitat was a decline in the total amount of winter range. Between 1975 and 1992,29,762 ha of sagebrush Artemisia spp. rangeland were converted to cropland, a 74% increase in cropland. Regression analysis suggested a relationship between sagebrush habitat loss and grouse population decline (R² = 0.59, P = 0.002). Approximately 1,244 km² of privately-owned sagebrush on the study area could potentially be converted to cropland, which we predict would have serious negative implications for the sage grouse population.

After the first discovery of the endemic Chinese grouse Bonasa sewerzowi in 1876 in the province of Gansu, the species has been found in several areas in China; most recently it was discovered in the county of Jiali in Tibet in 1997. Its distribution area should now be defined as Gansu, Qinghai, Sichuan, Yunnan and Tibet, corresponding to the distribution area of mountain conifer forest in central China between the altitudes of 2,400 and 4,300 m a.s.l. The habitat requirements of the Chinese grouse include two main factors; one is the presence of coniferous trees which are used for cover, and the other is the presence of deciduous trees and shrubs which are used for food. Due to intensive cutting of virgin forests during the past decades, the habitats of the Chinese grouse are heavily isolated and fragmented nowadays. However, due to present and future conservation measures in China, I believe in a better future for the species.

Greater prairie-chickens Tympanuchus cupido pinnatus are grouse of the tallgrass prairie of North America. Their range expanded greatly following the spread of early European agriculture into the grasslands and logging in forested areas. When the optimum mix of cropland and grass was exceeded, their range generally contracted to the regions where climatic and/or soil factors favoured the retention of grassland. Historically they probably occurred in 20 states of the United States and four Canadian provinces, but presently they only occur in 11 states and no longer in Canada. Their current status throughout the range varies considerably depending on habitat conditions, population levels, management capabilities and local land-use economic factors. A variety of conservation efforts, including translocation, are underway in the states where they occur, the intensity of which is generally inverse to numbers remaining. Noteworthy, is the Conservation Reserve Program (CRP) which has increased grassland cover on private land through incentive payments.

The paper sketches the state of grouse science as reflected by the 8th International Grouse Symposium held at Rovaniemi, Finland, in September 1999, by contrasting the representation of species, topics and trends in the literature and at the Symposium. The analysis was based on 5,349 grouse papers published between 1930 and 1998 and 75 abstracts submitted for the Symposium. In the past, grouse research has focused on the species with the greatest economic and cultural importance. During the 1990s, population dynamics, habitat and behaviour have become the major research topics. At the Symposium, a number of trends became apparent: increasing importance of genetic and landscape ecological studies, integration of disciplines, approaches and explanations, as well as increasing cooperation. However, a significant gap seems to exist in the documentation and communication of experience in grouse conservation and management.

The evolutionary processes behind the polygynous mating system known as leks are difficult to document. One approach is to study the behaviour that drives the formation of new leks today. Several hypotheses have been put forward to explain the formation of leks, and they can roughly be divided into two groups; one which advocates that the males are the driving force and one which argues that the females are the driving force. In this study we use data from a long-term study (1979–1998) of a capercaillie population at Varaldskogen in southeast Norway to develop a model describing how new leks are formed in this species. By using data on spacing pattern and behaviour of radio-marked young and adult birds of both sexes we demonstrate how the situation develops from winter towards mating in spring. Furthermore, we report on a few cases of new leks that have arisen in the area during the 20 years of study. We argue that both female mate choice, male territoriality and male attraction to locations with high densities of females are involved in a dynamic process of lek formation. We present results which indicate that new leks are mainly founded by young birds. Finally, we show that when new leks are formed the spacing patterns of the individuals involved change. This questions the method of using comparisons between the position of female home ranges and established leks to infer how leks are formed.

In a population study during 1979–1988 at Varaldskogen in southeastern Norway, 234 capercaillie Tetrao urogallus nests and broods were classified as first nests or renests. Of the females that had their first nest depredated, 9–87% (mean 36%) renested. Over a 6-year period, autumn brood production increased from 30 to 38% due to renesting. Renesting is physically demanding for the females; the eggs in renests are fewer and smaller, and the females take more and longer recesses than when incubating first nests. All the females incubated their first nests till the eggs hatched or the nest was depredated. Two of the renesting females took more and longer recesses until they gave up their nests. The ability to renest seems to be weight-related, as yearling females, which weigh less than adult females, did not renest, and the weight of adult females on leks was highest in the two years when most renesting occurred. The female will renest if the nest is depredated during the first three days of incubation. Each of the following 19 days, all years combined, a mean of 26% of females who lost their nests renested. Capercaillie renesting was related to the vole cycle; it was highest in the year before the small rodents peaked and decreased through the vole crash and the year after.

Capercaillie Tetrao urogallus and black grouse T. tetrix are threatened species in central Europe. Their habitats are fragmented both at a continental and a regional scale, and spatial connectivity may play an important role for their (meta-)population dynamics and persistence. In order to identify conservation priorities, it is important to know if and to what extent exchange between local populations occurs. In this paper, we present the rationale and techniques of an ongoing project into the spatial structure of capercaillie and black grouse populations in central Europe using non-invasive genetic methods. In this project, we assess the genetic differentiation of spatially distinct populations using microsatellite analysis based on DNA extracted from feathers. This approach will allow us to identify critical geographic distances beyond which demographic connectivity between populations is not assured. We expect to find a correlation between geographic and genetic distance.