The 8th International Grouse Symposium - Galliformes-sg.org

The 8th
International Grouse Symposium
Rovaniemi, Finland
September 13 - 17, 1999

ORAL PRESENTATIONS
Alatalo, R.: Lekking in the black grouse
Andreev, A.V.: Life history of the Siberian spruce grouse: a prisoner of evolution prospering in
habitat shadows
Angelstam, P.: Grouse as forest biodiversity management tools
Baines, D., Moss, R. & Summers, R.: Capercaillie breeding success in Scotland in relation to
predator abundance and habitat
Cartwright, K.S. & Robel, R.J.: Effects of landscape and landuse changes on greater prairie-chicken
population indices in eastern Kansas
Cattadori, I. & Hudson, P.: How does the temporal dynamics of grouse populations vary spatially?
Connelly, J.W., Apa, A.D., Smith, R.B. & Reese, K.P.: Hunting and predation as mortality factors
of adult sage grouse in Idaho
Drovetski, S.V.: Habitat, chick survival and density of Caucasian black grouse
Eliassen, S., Wegge, P., Finne, M. & Odden, M.: Daytime space use and social interactions in male
capercaillie during spring
Filacorda, S., Millsa, C., Pascottoa, E. & Perco, F.: Chaos approach to analysis of the black grouse
and rock ptarmigan population dynamics on the Julian and Carnian Alps
Finne, M., Wegge, P., Eliassen, S. & Odden, M.: Roosting habitat preference of capercaillie males
in spring - the importance of forest structure in relation to anti-predator behaviour
Fischer, I. & Storch, I.: Capercaillie and woodpeckers in alpine forests: which is the better indicator
species?
Frilund, G-E. & Pedersen, H.: Effects of hunting on natural winter mortality in willow ptarmigan
Gjerde, I., Wegge, P. & Rolstad, J.: Lost hotspots and passive female preference: the dynamic
process of lek formation in capercaillie grouse
Gregg, M.A., Crawford, J.A. & Coggins, K.A.: Relationship between vegetation changes and
productivity of sage grouse at Hart Mountain National Antelope Refuge, Oregon
Gutiérrez, R.J., Barrowclough, G.F. & Groth, J.G.: Molecular systematics of grouse
Hanowski, J.M. & Niemi, G.J.: Landscape requirements of sharp-tailed grouse in Minnesota, USA
Haysom, S.L. & Baines, D.: The distribution of lekking black grouse in commercial forests in
Scotland
Helle, P.: Assessing effects of changing forest landscape structure on grouse: the Finnish way
Helle, T., Alenius, V. & Naskali, A.: Capercaillie and gross national product - an example on
“reversed” U-curve
Hjeljord, O., Wegge, P., Ivanova, M., Beshkarev, A.B. & Rolstad, J.: Dispersal from leks and
spatial distribution of male capercaillie during summer in relation to dispersion and fragmentation
of habitat
Hjorth, I.: Adaptive radiations in neck anatomy for mating calls in grouse species
Hudson, P.: Population cycles in grouse - the stabilizing and destabilizing effects of parasites

Höglund, J.: The radiation of grouse
Jonzen, N., Ranta, E., Lundberg, P. & Kaitala, V.: Damped dynamics, harvesting and grouse
populations
Kastdalen, L., Pedersen, H.C., Willebrand, T. & Steen, H.: Testing of critical assumptions for the
use of dogs in line transects of willow ptarmigan
Klaus, S., Sun, Y-H., Yun, F. & Scherzinger, W.: Territorial and mating behaviour of Chinese
grouse at Lianhuashan, Gansu province, China
Leonard, K.M., Reese, K.P. & Connelly, J.W.: Distribution, movements and habitats of sage grouse
on the Upper Snake River Plain of Idaho: changes from the 1950’s to the 1990’s
Lindén, H.: The research programme for Finnish grouse
Marjakangas, A., Nikula, A. & Koski, A.: Effect of landscape structure on habitat selection and
survival of black grouse broods
Martin, K. & Wiebe, K.L.: White tailed ptarmigan production in years of extreme conditions in
Colorado: when to opt out, and how to compromise
Ranta, E.: Understanding dynamics of populations: Finnish grouse as an example
Robel, R.J., Jamison, B.E., Salter, G.C. & Walker, T.L. Jr.: Influence of human activity on the
utilization of sandsage habitat by lesser prairie-chickens in Kansas
Rätti, O., Helle, P. & Hollmén, T.: Selective hunting of parasitized grouse
Segelbacher, G. & Storch, I.: Comparing geographic distribution and genetic distance in capercaillie
and black grouse: genetic markers as a tool for large scale conservation
Smedshaug, C.: Increase in grouse hunting yield in early 20th century Norway: what happened
Smith, A.: Grouse populations, management and moorland vegetation
Storaas, T., Kastdalen, L. & Wegge, P.: Renesting in capercaillie
Storch, I.: Status, threats and conservation of grouse worldwide: an overwiev
Storch, I.: Grouse science as a process: where do we stand?
Sun, Y-H.: Distribution and status of Chinese grouse
Svedarsky, D., Westemeier, R.L., Robel, R.J., Gough, S. & Toepfer, J.E.: Status and management
of the greater prairie-chicken in North America
Yun F. & Sun, Y-H: Chick dispersal of Chinese grouse at Lianhuashan
Åberg, J., Jansson, G., Swenson, J. & Mikusinski, G.: Effects of stochasticity and scale on the
occurrence of hazel grouse in a boreal landscape

POSTER PRESENTATIONS
Artuso, I.: Black grouse and rock partridge in the Brembana Valley (Italian Central Alps) -
preliminary results
Artuso, I., Sepulcri, A., Frangipane, A., Filacorda, S. & Piasentier, E.: Dietary composition and the
presence of parasites in black grouse shot in the Val Brembana hunting preserve
Baba, Y., Fujimaki, Y., Klaus, S., Butorina, O., Drovetskii, S. & Koike, H.: Population genetics for
the hazel grouse using mithochondrial control region sequences
Bagliacca, M., Calzolari, G., Folliero, M. & Mani. P.: Artificial black grouse breeding in Tuscany
(Apennine region of Italy)
Barnett, J. K., Gregg, M. A. & Roberts, V. A.: Grouse in the National Wildlife Refuge System,
USA
Bevanger, K.: Reindeer fences as a mortality factor for ptarmigan in Finnmark, Norway
Brenot, J. F. & Menoni, E.: Winter densities and spatial evolution of a capercaillie population in
relation to construction of a ski station in the Pyrenees
Breuss, M. & Zeiler, H.: Winter feeding spectrum of hazel grouse in the Austrian Alps (Carinthia)
Brøseth, H. & Pedersen, H.: Hunting effort on temporal and spatial scale: tracking willow
ptarmigan hunters with GPS
Cadamuro, A., de Franceschi, P. F. & Mattedi, S.: Habitat characteristics of nesting and brood
rearing sites of black grouse in the Lanza Plans (Eastern Alps, Friuli-Venezia, Giulia, Italy)
Cartwright, K. S. & Robel, R. J.: Factors affecting greater prairie-chicken lek count data in eastern
Kansas
Cas, M.: The influence of forest changes in Alpine Slovenia on the dynamics of the capercaillie
population density
Ellison, L. & Caizergues, A.: Natal dispersal of black grouse in the French Alps
Fujimaki, Y.: Food habit of hazel grouse in Hokkaido, Japan
Hafner, F., Andreev, A., V. & Gossow, H.: On habitat use and mating system in the Siberian spruce
grouse
Hollmén, T., Franson, J. C., Rätti, O., Helle, P., Väyrynen, E. & Greiner, E.: Occurrence of
Haematozoa in grouse in Northern Finland
Hörnell, M. & Willebrand, T.: Willow grouse population trends from Swedish line transect counts
Kangas, A. & Kurki, S.: Hypotheses about the sex-ratio of capercaillie - the effect of different
assumptions about mortality rates
Keulen, C., Houbart, S. & Ruwet, J.-C.: Black grouse arenas in the Belgian Hautes-Fagnes:
landscape features and management proposals
Kumpu, P., Helle, P. & Vallinkoski, V.-M.: Demography of capercaillie in northern Finnish boreal
forests
Lande, T., Pedersen, H. C. & Pekins, P. J.: The cost of incubation in ptarmigan
Lee, W.-S. & Rhim, S.-J.: Characteristics of distribution and habitat structure of hazel grouse in
Korea
Lindén, H. & Pienmunne, E.: Non-stable age distributions of capercaillie cycles in Finland

Liukkonen-Anttila, T., Saartoala, R., Mäkinen, T. & Hissa, R.: Effects of hand-rearing on the
morphology and physiology of the capercaillie
Loneux, M., Lindsey, J. K. & Ruwet, J. C.: Influence of the climate on the dynamics of the black
grouse population in the Belgian Hautes-Fagnes since 1967
Mani, P. Rossi, G. & Bagliacca, M.: Pathology encountered in captive black grouse bred in the
experimental game bird farm of the national corp of foresters at Orecchiella Natural Park (Tuscany,
Apennine)
Mattedi, S., Perco, F. & Cadamuro, A.: Black grouse breeding and biometric parameters in sample
areas of Eastern Alps (Italy)
Menoni, E., Tautou, L., Magnani, Y., Poirot, J. & Larrieu, L.: Distribution of capercaillie in relation
to age of forest stands
Nielsen, O. K.: Winter mortality of ptarmigan in south-west Iceland
Odden, M., Wegge, P., Finne, M. H. & Eliassen, S.: Foraging behaviour of capercaillie in spring
Perco, F., de Franceschi, P. F. & Filacorda, S.: Influence of meteorological variables on black
grouse and rock ptarmigan populations in the Carnian and Julian Alps
Ploner, R. & Gossow, H.: Habitat suitability characteristics for capercaillie in Central Sweden and
North Italy - a comparison
Porkert, J. & Schindlatz, K-H.: Why is artificial breeding of Tetrao tetrix male x T. urogallus
female hybrids necessary?
Porkert, J., Solheim, R. & Flor, A.: Social relations between capercaillie and hybrid Tetrao tetrix
male x T. urogallus female on one capercaillie display ground in S. Norway
Ruwet, J.-C., Fontaine, S. & Houbart, S.: Development in numbers of lekking black grouse cocks
from 1966 to 1999 in the Belgian Hautes-Fagnes plateau
Ruwet, J.-C. & Hanon, L.: Behavioural ecology and social dynamics of black grouse at the arena
during a long-term study
Sachot, S., Perrin, N. & Neet, C.: Population dynamics of capercaillie in the Jura Mountains
Seiler, C.: Development of anti-predator behaviour of captive reared black grouse chicks
Seiler, C.: Telemetry study of released black grouse in Schleswig-Holstein, Germany
Sieber, U.: The display of black grouse cocks in a mire landscape in eastern Switzerland: influence
of landscape structure and human activities
Storch, I.: Density variation among alpine capercaillie populations at multiple spatial scales: how
much is explained by the habitat?
Strauss, E.: The decline of a black grouse population in a fragmented moor landscape
Sun, Y.-H., Klaus, S., Fang, Y. & Scherzinger, W.: Determination of Chinese grouse habitat at
landscape scale in the Lianhuashan Mountains
Tornberg, R.: Predation impact of the goshawk on grouse
Viht, E.: The density of the hazel grouse and its habitats in the central part of Estonia
Zeiler, H., Breuss, M. & Gossow, H.: Forest development and capercaillie habitats within the
Austrian Alps

LEKKING IN THE BLACK GROUSE
Rauno V. Alatalo, Matti Hovi, Jacob Höglund, Arne Lundberg & Pekka Rintamäki
I will give a summary of our research during 1987-1998 on lekking in black grouse Tetrao tetrix in
central Finland. Our first goal was to study lek evolution comparing the mating success at leks of
different sizes. Males, on average, benefit of joining the leks of larger size, since per capita mating
success reduces strongly in the smallest leks.
Females seem to be actively favouring to mate on larger leks. While matings are highly skewed,
individual males should choose their lekking site according to the relative position in the mating
rank. Playback experiments demonstrated that males are attracted by the presence of other males,
and they also suggested female preference for larger male vocal display. Within leks females favour
central males that display social dominance in interactions with other males. Instead many other
traits, such as male size, lyre length, plumage condition etc. have only marginal effect on male
mating success. In general, the mating system allows the females to mate with the most viable
males. We have also followed the success of individual males over the years, and the most
succesful males in terms of lifetime mating success are among those that survive until the age of 3-4
year. However, annual mortality is high, and thus it seems that males of poor condition make an
attempt to mate already during in their first breeding season, while the future top males are very
passive during their first spring. Once a female has settled o choose a certain male, somewhat
surprisingly, she prefers to mate with the same male in the next year too, even if in general on larger
leks the same male cannot hold his top rank in more than one year.

LIFE HISTORY OF THE SIBERIAN SPRUCE GROUSE: A PRISONER OF EVOLUTION
PROSPERING IN HABITAT SHADOWS
Alexander V. Andreev
A local population of the East-Asian endemic species, the sharp-winged grouse (or the Siberian
spruce grouse) Falcipennis falcipennis was studied in the central part of its range since 1985 with
intensive use of radiotracking in 1994-1997 (49 birds). The study area of 100 sq km, located 100
km north of Komsomolsk, is a dense mosaic of mature and growing stands of the Ajan spruce Picea
ajanensis and larch Larix dahurica mixed with bogs, windfall openings, clearcuts and burns. In
winter, the birds tend to stay in flocks of 2-5 individuals, the hens being more mobile than cocks.
They spend 12-15 hr overnight in the snow burrows under subzero temperatures, but never in the
daytime. As night temperatures are above -20°C, the birds prefer to stay in the spruce or fir canopy.
Their winter diet is exclusively spruce needles. Winter flocks disperse in two steps: by late March -
early April adult cocks move to their lekking sites, and by late April females disperse to look for
mating partner and establish nest territory. Shortly prior to the prebreeding dispersion, the hens are
prone to visit anthills, which provide an important part of the diet at the time.
The mating period starts by the last week of April. The distribution of leks follows the topography
of the study plot. We distinguish the circular pattern (leks follow the bog edges), the linear pattern
(leks follow mountain crests), and the clump pattern (leks surround hilltops).
With the two first mentioned patterns, neighboring cocks display in 400-700 m distance from each
other, and visual and sound contacts between them are minimized. Only subadult cocks’ random
visits diversify these relationships. Under the third pattern, the distance between lekking cocks
varies from 250 to 400 m, and they communicate by ground flatter flights and fairly frequent
aggressive contacts. The distribution of females after mating season follows the distribution of
cocks. Stronger cocks (+3 years) definitely preferred by hens render more copulations than their
neighbors.
The copulation period occurs from 30 April through 9 May and the egg laying starts between 7 and
15 May. During the laying period, females prefer to feed on the outcoming larch needles. Unlike its
congeneric species in North America, the Siberian spruce grouse does not depend on the ground
vegetation at this time. In fact, larch needles perform the principal food from mid-May through
October. The species equally depends on two coniferous species - the Ajan spruce and the Siberian
larch. Only some small ants are taken from the ground. The clutch size varies between 5 and 7 eggs
(5.75 on average, n=13). Older hens begin to incubate one week earlier (12-15 May) than younger
and they also produce larger clutches - 6-7 eggs vs. 5 eggs. The incubation period lasts 23.5 days
(572-580 hr, n=5). Incubating females depart the nest 2-3 times a day for approx. 30 min bouts.
Nest attendance is higher than in other grouse species (0.95-0.97).
The chicks hatch on 6-13 June. During the first two weeks they feed entirely on insects.
The broods begin breaking up by mid-September. At the time of fall dispersion, a good proportion
of young birds are preyed by the owls. The distance of dispersion varies between 1-2 km to 15 km
or more, most commonly within 2-5 km. The overall spring density of the population in the study
area was fairly stable throughout the years: 6-8 birds/sq km.

GROUSE AS FOREST BIODIVERSITY MANAGEMENT TOOLS
Per Angelstam
To secure the long-term maintenance of biodiversity in managed taiga forests, the evolutionary
authentic natural disturbance regimes, and the associated habitats, must be maintained.
Consequently, the composition and structure of habitats as well as processes found in a naturally
dynamic forest ecosystem must be assessed. In Europe there has been some improvements in the
field of practical forest biodiversity assessment. This development has, however, been at the scale
of trees within forest stands, and the scale of forest stands within landscapes, rather than at the scale
of forest landscapes in a region.
To measure the level of naturalness/authenticity at the scales of trees and forest stands, a number of
lichens, fungi and vascular plants are being successfully used in the Nordic countries. However, at
the scale of landscapes, forest species with large area requirements, and which are sensitive to loss
of natural forest components and to changes in land use and forest management should also be used.
This could be made through inventories of species specialising in the different forest types, or to
forest landscape criteria such as stand age, patch size, habitat amount etc., that are defined by the
species´ habitat requirements. Naturally, species´ indicators must scale to the chosen spatial
resolution.
I argue that for the dominating taiga disturbance regimes, the presence and sustained viable
populations of all four forest-living grouse (willow grouse, black grouse, hazel grouse and
capercaillie) in a landscape ensures a sufficiently large variation in age classes ranging from
recently disturbed forest to old forest (but not old-growth). The advantage of grouse over several
other species is that their ecologies are well known by the end-users of environmental monitoring
schemes. This greatly enhances the communication of the status of the forest environment.

BLACK GROUSE AND ROCK PARTRIDGE IN THE BREMBANA VALLEY (ITALIAN
CENTRAL ALPS) — PRELIMINARY RESULTS
Ivano Artuso
This study project aims to investigate the status of black grouse Tetrao tetrix and rock partridge
Alectoris greaca populations in the Brembana valley during the period 1996-2000. There are about
12,000 ha suitable territory for these two species locating in the southern part of the Italian central
Alps.
Censuses of the summer 1998 revealed altogether 426 black grouse (5,145 ha surveyed) and 224
rock partridge (6,450 ha surveyed). Compared to the two previous years, the number of reproducing
breeding adults of both species was lower in the spring 1998, but a good reproductive success was
observed in the summer with a lot of hens with broods and high number of juveniles. About 100
persons carried out these spring/summer yearly counts, which were locally sponsored.

DIET COMPOSITION AND THE PRESENCE OF PARASITES IN BLACK GROUSE SHOT IN
THE VAL BREMBANA HUNTING PRESERVE
Ivano Artuso, Angela Sepulcri, Antonio Frangipane, Stefano Filacorda & Edi Piasentier
The composition of the diet in the crops and gizzards of 50 black grouse Tetrao tetrix tetrix shot in
October and November in the "Valle Brembana (BG)" hunting preserve was studied, together with
the presence of helminth parasites in the intestine. The composition of the gizzards appeared
generally coherent with the observations in the crops, but there was a tendency to underestimate the
fruits and leaves from bilberry Vaccinium myrtillus, shoots from cowberry Vaccinium vitis-idaea
and the grass components. The bilberry shoots appeared to be the most important food items in
terms of frequency and abundance followed by the fruits and buds from rowan Sorbus spp. and
bilberries. In addition to bilberry, rowan and willow Salix spp., larch Larix decidua and juniper
Juniperus communis appeared frequently, apparently as substitute foods. The adult birds also had a
diet rich in grass components, while the young birds appeared to prefer the fruits of rowan and
bilberry, and, to a lesser extent, larch needles and rubigenous rhododendron Rhododendron
ferrugineum. The diet appeared to depend on the shooting area, with sites in which the diet was
composed primarily of juniper and rowan and those in which ericaceus showed low frequencies.
The presence of nematodes such as Ascaridia spp. and Capillaria spp. in the small intestine was
widespread.

POPULATION GENETICS FOR THE HAZEL GROUSE USING MITOCHONDRIAL
CONTROL-REGION SEQUENCES
Yoshiyuki Baba, Yuzo Fujimaki, Siegfried Klaus, Olga Butorina, Serguei Drovetski, Vitalii. A.
Nechav, Sergei. G. Surmach & Hiroko Koike
More than 150 samples of hazel grouse Bonasa bonasia were collected from six locations in
Hokkaido, Sakhalin, Magadan, Siberia, Vostretsovo and Europe. A total of 56 haplotypes were
detected at the 400 base pair in left domain of the mitochondrial control-region. Compared with the
Chinese hazel grouse Bonasa sewerzowi, these two species had separated over 100,000 years ago.
Genetic tree using the 56 haplotypes indicated that most of the haplotypes radiated at 2-3
substitution differences from the root-group, suggesting hazel grouse root was 40,000-30,000 years
ago in East-Asia. Hokkaido was populated before the coldest phase in the last glacial period (about
20,000 years ago), and Magadan, Siberia and Europe were populated after the last glacial period.
Haplotype diversity in Hokkaido was between 0.82 and 0.97, which is higher than that in Sakhalin
and Magadan, suggesting that the southern populations have higher genetic diversity that northern
populations.

ARTIFICIAL BLACK GROUSE BREEDING IN TUSCANY (APENNINE REGION OF ITALY)
Marco Bagliacca, Giorgio Calzolari, Maurizio Folliero & Paolo Mani
Captive breeding of black grouse Tetrao tetrix is justified because this species is declining in most
of the alpine areas, at the edge of its distribution area. Only reared black grouse originating from the
alpine population, can be used for re-introducing the species in the alpine areas where it has
disappeared or is endangered. For this reason in 1988 the Orecchiella natural park (44.1°N, 1200m
a.s.l.) started the artificial breeding. The experimental breeding has been always based on fixed
pairs; it started with 6 pairs originating from East Alps. In 1993-94, eight young birds, obtained
from eggs coming from Central Alps, were taken for crossbreeding. There are altogether 18 males
and 13 females. The pens built in 1988 were provided with complete net floors and some addtional
pens were built in 1993 and in 1996. Nutrition was based on a three-phase feed (laying, growth, and
rest). Since it was not possible to use merely artificial feed, some natural long-fiber supplementary
feed (Pinus mugo for adults, and Vaccinium myrtillus for growing birds) with siliceous grit was
always given. The laying period in the experimental breeding farm starts in May and ends in July
(mean 5 eggs/female). Eggs are artificially incubated (99.7°F, 47%R.H). The incubation period is
24.5 days. The mean hatchability is 40%. Chicks are reared in commercial heated cages for the first
4 weeks before transferred to the pens. Pairs are formed in the winter.

CAPERCAILLIE BREEDING SUCCESS IN SCOTLAND IN RELATION TO PREDATOR
ABUNDANCE AND HABITAT
David Baines, Robert Moss & Ron W. Summers
Capercaillie Tetrao urogallus breeding success was estimated from fourteen forests in Scotland
during 1991-97. Forest structure, ground vegetation, and predator abundance, were assessed.
Breeding performance differed ten-fold between forests and also between years. Capercaillie bred
more successfully where predators were fewer. Breeding success (chicks per female) and the
proportion of female capercaillie with broods was negatively correlated with carrion crow Corvus
corone abundance and with PCA predator scores. Mean brood size was negatively correlated with
fox abundance.Years when breeding success was low were associated with high temperatures in
May and higher rainfall in mid-June. Weather did not appear to affect brood size, only the
proportion of females with broods.

GROUSE IN THE NATIONAL WILDLIFE REFUGE SYSTEM, USA
Jenny K. Barnett, Michael A. Gregg & Victoria A. Roberts
The National Wildlife Refuge System is the only network of federal lands in the United States of
America set aside solely for the purpose of preserving wildlife and their habitat. Since 1903 over
500 refuges have been established, protecting waterfowl, large ungulates, endangered species and
upland game birds. Species to be managed for by each refuge is set forth in federal law and policy.
Two refuges, Attwater Prairie Chicken NWR and Buenos Aires NWR were established to provide
habitat for endangered ga lliform species. Both refuges manage habitat and are involved in captive
breeding and re-introduction programs. Refuges play a role in grouse research. Concern about
sage grouse is increasing as populations decline across North America. Research has been
conducted on sage grouse Centrocercus urophasianus on the Sheldon/Hart Mountain Refuge
Complex since 1988. Studies have focused on diet, nutrition, and habitat use by nesting, brooding,
and broodless hens. Investigations into vegetation and animal physiology to assess overwinter and
pre-laying nutrition in females have started this year. Livestock grazing was removed from this
342,000 hectare complex in 1991 and replaced with prescribed fire as the primary management tool.
Continued research will investigate the effect of livestock removal and fire on sage grouse.

REINDEER FENCES AS A MORTALITY FACTOR FOR PTARMIGAN IN FINNMARK,
NORWAY
Kjetil Bevanger
Between 1991 and 1994 data were collected to document the importance of reindeer fences as a
mortality factor for birds in the county of Finnmark, northern Norway. The fieldwork was mainly
patrols searching for dead birds and their remains along 12 sections of reindeer fence (totalling 71.1
km) immediately after the snow thawed in early spring (May-June). The fences consisted of steel
wire, steel netting or a combination of these, with heights varying from 100 to 250 cm. A total of
179.9 km was covered and 253 collision victims belonging to at least 20 species were found.
Willow ptarmigan Lagopus lagopus and rock ptarmigan L. mutus comprised 85% of the victims.
Experiments showed that the patrols found approximately 64% of the total number of ptarmigan
that had been killed by the fences (and remained close to them) during the winter. I estimated that at
least two ptarmigan are killed per km of reindeer fence per year in Finnmark. The exact length of
reindeer fences in Finnmark is unknown, but it is probably more than 1000 km.

RESPONSE OF A CAPERCAILLIE POPULATION IN RELATION TO CONSTRUCTION OF A
SKI STATION IN THE PYRENEES.
Jean-François Brenot & Emmanuel Ménoni
A cross country ski station was created in 1988 on the Beille Plateau in the Pyrénées (SW France)
in a habitat of mountain pine Pinus uncinata and grasslands. Between 1989 and 1997, we followed
numbers and distribution of capercaillie in winter, both at the station area and on a nearby
undisturbed control site. The ski station had a strong negative effect on capercaillie, mainly because
ski trails crossed winter ranges of the birds. In fact, capercaillie density in winter was negatively
correlated with density of ski trails. Since 1989, number of capercaillie (about 130 in 1989)
declined at the rate of 9 % per year at the ski station area, but remained stable (about 50) on the
control area.
The decline resulted from birds abandoning wintering sites nearest to the ski trails, representing a
loss of 18% of the total winter range. On the control area, no wintering sites were abandoned
between 1989 and 1997.

WINTER FEEDING SPECTRUM OF HAZEL GROUSE IN THE AUSTRIAN ALPS
(CARINTHIA)
Monika Breuss & Hubert Zeiler
We identified the winter foraging spectrum of hazel grouse Bonasa bonasia in the Southern
Limestone Alps (Carinthia, Austria) through an analysis of dropping samples (N=83). The study
area consisted of two parts: one characterised by single-tree-selection felling as the main harvesting
method, resulting in a multi-layered forest with a high plant diversity. In the second area harvesting
through clear-cut felling is practised, resulting in a forest with even aged stands and fewer plant
species. Norway spruce Picea abies dominates in both areas.
In the multi-layered forest, hazel grouse feeds on a diverse plant spectrum: mainly on buds, twigs
and catkins of deciduous trees (aspen Populus tremula, grey alder Alnus incana, common beech
Fagus silvatica, rowan Sorbus aucuparia, whitebeam Sorbus aria), shrubs (willow Salix sp., fly
honeysuckle Lonicera xylosteum, hazel Corylus avellana) and dwarf shrubs (blueberry Vaccinium
myrtillus). Within the clear-cut-stands hazel grouse feeds on fewer plant species; mainly buds and
catkins of hazel, grey alder, aspen, stems of blueberry and leaves of wood-sorrel Oxalis acetosella,
also needles of Norway spruce occur in this sample frequently. Because of variation in the snow
conditions in the study period, it could be shown that hazel grouse prefers green parts of ground
vegetation if the snow cover allows it.

HUNTING EFFORT ON TEMPORAL AND SPATIAL SCALE: TRACKING WILLOW
PTARMIGAN HUNTERS WITH GPS
Henrik Brøseth & Hans Chr. Pedersen
With GPS tracking of hunters, radio-marked willow ptarmigan Lagopus lagopus and a GIS, we
studied hunter-game interactions on temporal and spatial scale in central Norway. Ptarmigan
hunters walked on average 16.2 km daily at a speed of 2.8 km/hour, and they hunted for 9 hours
each day, of which almost 6 hours was active hunting time. During 50 hunter-days they had 295
hours of active hunting, covered a distance of 818 km and harvested 20% of the population in the
area. The spatial distribution of hunting pressure was strongly dependent on the starting point of the
hunters, and areas close to the base-cabin received most of the hunting activity. The spatial
distribution of the harvest correlated well with hunting pressure, except in the close proximity of the
cabin, where fewer birds than expected were harvested. Shot radio-marked birds had twice as high
hunting pressure and lived closer to the cabin, compared to surviving marked birds. Marked birds
altered their behaviour in response to hunting activity, by increased daily movement distance. This
method of obtaining quantitative data of human effort should also have application possibilities for
other studies were one need to analyse human effort on temporal and spatial scales.

THE INFLUENCE OF FOREST CHANGES IN ALPINE SLOVENIA ON THE DYNAMICS OF
THE CAPERCAILLIE POPULATION DENSITY
Miran Cas
The original alpine land in the capercaillie Tetrao urogallus area of the Slovenian Mts (about
12,000 km²) was covered 210 years ago by pastures with small areas of beech-fir forests mixed
with planted Norway spruce (coverage about 30%; Josephine’s military maps). Today coniferous
forests (coverage about 80%) are prevailing changing the suitability for the capercaillie as shown by
population densities during the last 120 years.
At all times from the year 1784 the capercaillie were dependent on habitat of suitable structure with
more than 50% mixed mature coniferous forests. We have three well defined periods of population
estimates after 1874 (hunting statistics and counts) in three state regulations of Slovenian regions in
the history. The maximal population densities of the three documented “cycles” (the duration of
each is between 25 and 28 years) were observed in 1880, 1908, 1933 and 1961. The last observed
maximum was in 1961 and the minimum in 1982.
The next predicted (not confirmed) maximum is between 1990 to 1995. Until 1933 the population
density in “cycles” was significantly increasing. We attribute this to the overgrowth of pastures and
to increasing amount of mature forests over 80 years. After 1961 we find a decreasing phase. We
consider this to be due to the overgrowth of the landscape and the negative influences of man in the
forest where the capercaillie lives today.

EFFECTS OF LANDSCAPE AND LANDUSE CHANGES ON GREATER PRAIRIE-CHICKEN
POPULATION INDICES IN EASTERN KANSAS
Kelly S. Cartwright & Robert J. Robel
Historically greater prairie-chickens Tympanuchus cupido were present across the tallgrass prairies
of North America. Their range and numbers declined coincidentally with loss of tallgrass prairie
habitat to agricultural expansion and other human activities. Declining greater prairie-chicken
populations stimulated the development of several census techniques to monitor population status.
Kansas has used lek counts along fixed survey routes to provide an index to greater prairie-chicken
populations since 1963. This population index has declined over the past three decades and our
study was initiated in 1996 to determine if landscape and landuse factors along the routes are
associated with the declining index. Landscape features (tree and shrub coverage, cropland,
grassland, residential development, etc.) and landuse (grazing practices, burning frequency, etc.)
have been determined along the lek survey routes for the 1963-1996 period in seven eastern Kansas
counties. Preliminary analyses show tree/shrub coverage, burning frequency and grazing intensity
increasing along the survey routes and grassland area decreasing. Correlation analyses are being
used to compare landscape and landuse changes with changes in lek survey data. We believe our
results will be relevant to most grouse surveys utilizing fixed transects or routes as their sampling
unit.

FACTORS AFFECTING GREATER PRAIRIE-CHICKEN LEK COUNT DATA IN EASTERN
KANSAS
Kelly S. Cartwright & Robert J. Robel
The greater prairie-chicken Tympanuchus cupido is a grouse of the tallgrass prairies of North
America and its population has declined drastically because of the loss of tallgrass habitat to
intensive agriculture and other human activity. Kansas initiated several census methods to monitor
prairie chicken populations in the late 1950s and adopted lek counts along permanent survey routes
as its primary monitoring technique in 1963. During these counts, biologists record peak numbers
of lekking birds along permanent 16-km routes. The counts are designed to be conducted from 40
minutes before to 90 minutes after sunrise between 20 March and 20 April, and when wind speed is
less than 20 km/h. Starting and finishing times of the counts, as well as observation dates, have
varied widely over the years. One objective of this study is to determine the effect of time-of-day
and day-of-year on numbers of birds counted along routes. Observations of prairie chicken activity
on 3 leks during spring 1997, 1998, and 1999 disclosed that numbers of birds on leks varied with
time-of-day and day-of-year. These data allowed the development of an adjustment model to
remove variability introduced by conducting lek count surveys at different times of the day and
different days of the year. We believe this approach may enable other grouse survey data collected
under different conditions to be adjusted to a standard set of conditions.

HOW DOES THE TEMPORAL DYNAMICS OF GROUSE POPULATIONS VARY
SPATIALLY?
Isabella M. Cattadori & Peter J. Hudson
The tendency of the populations to exhibit cycles as well as the period and amplitude appears to
vary geographically within the species’ distribution. We have been investigated the long term
temporal dynamics of four grouse species in the Italian Alps in an attempt to reveal specific
differences in the spatial pattern of populations at the southern edge of their European range.
Hunting statistics collected at regional scale and referred to different time periods have been used
and time series analysis performed. Rock ptarmigan Lagopus mutus, hazel grouse Bonasa bonasia
and black grouse Tetrao tetrix populations exhibited a weak tendency to cycle, no regular
fluctuations were found in time series of capercaillie Tetrao urogallus. We compare the findings
with studies conducted on populations of other European countries and suggest possible reasons for
the reduced tendency to cycle in the populations of southern Europe.

RELATIONSHIP BETWEEN VEGETATION CHANGES AND PRODUCTIVITY OF SAGE
GROUSE AT HART MOUNTAIN NATIONAL ANTELOPE REFUGE, OREGON
Kreg A. Coggins, John A. Crawford & Michael A. Gregg
Vegetation and sage grouse Centrocercus urophasianus productivity relationships were investigated
during 1989-1997 at Hart Mountain National Antelope Refuge, Oregon, USA. Initial studies
indicated sage grouse habitat preference and reproductive success were related to herbaceous
vegetation. Livestock grazing was eliminated from Hart Mountain in 1991 and during 1989-1997
there was an upward trend in growing season precipitation. Because grazing and precipitation
influence herbaceous vegetation, we investigated the combined effects of these factors on sage
grouse nest and brood habitat preference and productivity. Canopy cover of herbaceous vegetation
increased during the 8 years of study. Nest initiation and success was greater during 1995-1997
compared with 1989-1991. Increasing nest initiation rates were concordant with increases in spring
forb cover. Greater availability of forbs probably provided hens more opportunity to become
physiologically prepared to attempt nesting. Increased residual grass cover likely resulted in a
higher probability of hens initiating nests in cover that provided greater nest concealment and
resulted in higher nest success. Brood success was slightly greater during 1995-1997 compared
with 1989-1991. Greater forb availability, perhaps, reduced home range size and movements of
hens with broods, which are related to the nutritional requirements of chicks. As a result chick
survival was somewhat improved.

HUNTING AND PREDATION AS MORTALITY FACTORS OF ADULT SAGE GROUSE IN
IDAHO
John W. Connelly, Anthony D. Apa, Randall B. Smith & Kerry P. Reese
Sage grouse Centrocercus urophasianus populations have been studied in Idaho from 1977 to
1998, using banding and radiotelemetry to obtain mortality data. Here we assess mortality in these
populations and test the hypotheses that type of mortality is independent of gender and losses are
equally distributed over the year. Predation was the most common cause of death for grouse. For
adult males, 81% of deaths were attributed to predation and 19% to hunting. For adult females,
53% died from predation and 41% from hunting. Thus we rejected the hypothesis that type of
mortality is independent of gender (G = 5.75, P < 0.025) and concluded that females were more
likely to die from hunting than males. Thirty-nine percent of all deaths occurred in the months of
September and October, 47% from May through August. Of the September/October deaths, >95%
were from hunting. Only 2% of the deaths occurred during winter. In 6 of 15 years, harvest rates
for adult females may have exceeded 10%, while this rate was only exceeded in 2 of 15 years for
adult males. Although exploitation rates for grouse were relatively low, harvest of females may
have been excessive in some years. Hunting seasons for this species should be relatively
conservative compared to seasons for other gamebirds.

HABITAT, CHICK SURVIVAL AND DENSITY OF CAUCASIAN BLACK GROUSE
Sergei V. Drovetski
We surveyed Caucasian Black Grouse residing on Lagonakskiy Ridge (NW Caucasus, Russia) in
June-July 1998, and Magisho Ridge in July 1999. Weather affected habitat use by brood hens: on
sunny days 18 broods were encountered in meadows, 6 in ravines, and 3 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 encountered were in
ravines; just one was in a meadow. Habitat use by adult males was not affected by weather and
differed from habitat use by females and subadult males. Cryptically colored 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. Broods lost an average of one chick per ten days. An Accipiter gentilis
killed one adult female. Our density estimation for Lagonakskiy Ridge (2.3 adults per 1 km 2 ) was
similar to those reported elsewhere (2.3 ± 1.2; n = 7).

DAYTIME SPACE USE AND SOCIAL INTERACTIONS IN MALE CAPERCAILLIE DURING
SPRING.
Sigrunn Eliassen, Per Wegge, Mats H. Finne & Morten Odden
During the breeding season adult male capercaillie Tetrao urogallus occupy relatively exclusive
day-ranges extending radially from the lek centre. In this study, 14 males were radiotracked in a
coniferous forest in southeastern Norway during March-May 1996-1997. The daytime spacing
behaviour of adult males was examined, both in relation to neighbouring males, and with reference
to fragmentation of the forest habitat. Playback experiments were performed to check for territorial
behaviour at daytime. In general, males did not respond aggressively to any of two different
announcement sounds presented when in the core areas or at the periphery of their daytime ranges.
The spatial overlap of 95% kernel home ranges between neighbouring males averaged 18%
(SD=25%), but there was no temporal overlap. The males seemed to change between core areas
inside their home ranges in a 3-7 day successive manner. The space use patterns varied with the
social status of males and with the degree of fragmentation of habitat, but the analysis is at present
not conclusive. Additional tracking data will be included for further analysis on spatial
arrangements and social interactions.

NATAL DISPERSAL OF BLACK GROUSE IN THE FRENCH ALPS
Laurence Ellison & Alain Caizergues
Data on dispersal distances of juveniles are important for understanding the genetic structure of
populations, population regulation processes and the effects of landscape ecology on
metapopulation dynamics. We studied natal dispersal in juvenile black grouse Tetrao tetrix in the
French Alps between 1990 and 1997, by radio-tracking 40 young grouse captured in August or
early autumn on a 836-ha area. Natal dispersal distance for females was the straight-line distance
from the point of capture to the first nest, and for males the distance from point of capture to the
median of the x and y coordinates of the first summer range. Natal dispersal occurred in two
phases, in autumn (October and November) and in spring (April). Mean distance travelled by
females exceeded that of males in autumn (6.35 km vs 1.56 km, P < 0.0005 ) but not in spring (4.61
km vs 2.72 km, P < 0.37). Natal dispersal distance of females was greater than that of males (7.96
km vs 1.45 km, P < 0.0005). Natal dispersal resulted in most females (14/16) leaving the 836-ha
study area to nest 5 to 29 km from their site of capture. Males were more philopatric. By the next
summer after capture, only 1 of 11 had emigrated from the study area.

CHICK DISPERSAL OF CHINESE GROUSE AT LIANHUASHAN
Yun Fang & Yue-Hua Sun
In brood period, female Chinese grouse Bonasa sewerzovi and their chicks leave the spring home
ranges, whereas males still stay in their territories. The time for brood dissolution varied in 1995-
1998, from mid-September to early October. Six chicks were radio-tracked in 1995 and 1997 at
Lianhuashan Natural Reserve in Gansu Province, China. The birds were captured by nets with the
aid of radio-equipped hens before brood dissolution. Two chicks were captured on 20 September
1995. In early October, one moved in a winter flock 2.8 km away, the other one took a round trip of
6.0 km, and then stayed in a winter flock 500 m from her mother. Her nests in 1996 and 1997 were
0.60 km and 0.78 km from her birth site. In 1997, four chicks of our radio-equipped hens were
radio-tracked, and the nests of two female chicks in 1998 were 0.43 km and 0.81 km away from
their birth site, and one used the same brood range as her mother. We conclude that the chicks do
not live in the same flock with their mother in winter, and at least female chicks do not move far
from their birth site.

CHAOS APPROACH TO ANALYSIS OF THE BLACK GROUSE AND ROCK PTARMIGAN
POPULATION DYNAMICS IN THE JULIAN AND CARNIAN ALPS
Stefano Filacorda, Colin Mills, Ernesto Pascotto & Fabio Perco
The black grouse Tetrao tetrix tetrix and rock ptarmigan Lagopus mutus helveticus populations in
the Carnian and Julian Alps show a general dramatic decrease. In order to compare these species the
'Osservatorio Faunistico del Friuli-Venezia Giulia-Udine' has analysed their population dynamics
through a density-dependent model and a chaotic approach. For the analysis we have used the
hunting bags (HB) and population censuses (CE) collected for 26-36 years in the Julian and
Carnian Alps. Parameter r’ has been considered a measure of density-dependence and Lyapunov
exponent, r, of chaotic (r=positive), periodic (r=negative) or bifurcation (r=0) responses of system.
The apparent natural time scales, t, have been studied. The ptarmigan-CE (r'=0.91) and grouse-CE
(r'=0.71) show a stronger density-dependence than ptarmigan-HB (r'=0.45) and grouse-HB
(r'=0.59). The results for t are: 4.8 year for ptarmigan-CE, 1.3 ptarmigan-HB, 2.9 grouse-CE, and
1.9 years for grouse-HB. The ptarmigan-CE (r=-0.21) moves rapidly toward bifurcation response
and grouse-CE (r=-0.35) slowly to periodic; the ptarmigan-HB (r=-0.80) and grouse-HB (r=-0.51)
show the tendency to change from periodic to bifurcation response. The t, r, and r' seem to be useful
parameters to describe the population fluctuations. The abrupt shifting of population dynamics to
bifurcation response, as observed for rock ptarmigan, should correspond to increasing likelihood of
local extinction.

ROOSTING HABITAT PREFERENCE OF CAPERCAILLIE MALES IN SPRING - THE
IMPORTANCE OF FOREST STRUCTURE IN RELATION TO ANTI-PREDATOR
BEHAVIOUR
Mats H. Finne, Per Wegge, Sigrunn Eliassen & Morten Odden
Radioequipped capercaillie Tetrao urogallus males (n=14) were monitored during April and May
1996-1997 in a coniferous forest in southeastern Norway. Daytime roosting habitat preference was
investigated on two levels of scale: forest type and within stands. Radiolocations when the birds
were not active were used to examine forest type preference (n=127). For habitat preference within
stands, roosting sites (n=94) were located by searching for heaps of intestinal droppings.
When roosting, males selected forest types with high tree density, high horizontal and vertical
cover, and low visibility, such as spruce-dominated old forest and plantations >3 m. Ninety percent
of the roosting sites were located underneath the low branches of a spruce tree. Within old forest
stands, these sites had higher tree density, higher cover, and shorter visibility than the nearby
surrounding forest. In plantations, roosting sites tended to have lower stem density (2,400 trees/ha
vs 2,938 trees/ha), but higher vertical and horizontal cover and shorter visibility than random
locations within the same stands.
Because dense cover decreases the probablility of being detected by a predator, but at the same time
increases the chance of being killed once detected, the birds have to compromise between shelter
and outlook. The data indicate that males select good cover at the expense of good overview of the
surroundings when selecting roosting habitat during daytime.

CAPERCAILLIE AND WOODPECKERS IN ALPINE FORESTS: WHICH IS THE BETTER
INDICATOR SPECIES?
Irene Fischer & Ilse Storch
In Central Europe the capercaillie Tetrao urogallus is widely regarded as an indicator species of an
intact mountain forest community. Conservation efforts to improve capercaillie habitat are believed
to benefit other mountain forest species as well. The objective of the paper is to assess the
applicability and limitations of this concept. We compared habitat relationships of the capercaillie
in the Bavarian Alps with that of two other species commonly used as indicators of mountain
forests, the black woodpecker Dryocopus martius and the three-toed woodpecker Picoides
tridactylus. All three species prefer late successional stages, but use different resources within their
habitat; e.g. capercaillie prefer feeding on bilberry, while woodpeckers require insect-infested dead
wood and large trees for their nest cavities. Good capercaillie habitat was correlated with high
woodpecker use, but the presence of woodpeckers did not indicate habitat suitability for
capercaillie. Our results suggest that habitat management for capercaillie is likely to also provide
for the habitat needs of woodpeckers, but not vice versa. We conclude that the use of the
capercaillie as an indicator species is justifiable as long as management measures meet speciesspecific
requirements of all target species, e.g. standing dead wood for woodpeckers.

EFFECTS OF HUNTING ON NATURAL WINTER MORTALITY IN WILLOW PTARMIGAN
Gunn E. Frilund & Hans Chr. Pedersen
Here we examine if hunting mortality is additive or compensatory to natural winter mortality in
willow ptarmigan Lagopus lagopus. In willow ptarmigan, winter mortality is assumed to be about
60%, so this species might have a potential for compensation. Hunting mortality was studied in a
120 km² area, divided into five sub-areas, in Central Norway. The hunting season was between
10th of September and 1st of November. Over a three-year-period, all sub-areas had 0%, 15% and
30% harvest rate, estimated from line-transect censuses of the population in August. A total of 215
birds were radiomarked in all sub-areas, using radios with mortality sensors. Most of the birds were
marked in August shortly before hunting started, and some were marked in March-April and while
nesting. Birds have been tracked from ground and from aeroplane throughout the year. The tracking
has been most effective during summer and in the hunting period, but more sporadic during winter
and spring. Unfortunately it was not possible to cover all the areas and birds with the same
intensity. In addition to discussing the effects of hunting mortality in relation to natural mortality,
we will discuss findings of mortality rates in an inland population of willow ptarmigan in relation to
earlier findings often used in population dynamic calculations.

FOOD HABIT OF HAZEL GROUSE IN HOKKAIDO, JAPAN
Yuzo Fujimaki
The food habit of hazel grouse Bonasa bonasia was analyzed basing on crop contents of 229 birds
shot from early October to late January of 1994/95 and 1995/96 and from April to September 1996
in Hokkaido, Japan. Main foods were buds and catkins of deciduous broad-leaved trees from
November to January, fruits from October to December, and herbs during snow-free season. In
addition to these foods, arthropods were eaten in June and July. Comparing food habits in Europe,
in Hokkaido buds of many kinds of tree species were eaten in winter and fruits of liana such as Vitis
coignetiae and Actinidia arguta were important foods, although the food habit is fundamentally
similar to that in Europe.

LOST HOTSPOTS AND PASSIVE FEMALE PREFERENCE: THE DYNAMIC PROCESS OF
LEK FORMATION IN CAPERCAILLIE GROUSE
Ivar Gjerde, Per Wegge & Jørund Rolstad
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 leks today. Several
hypotheses have been put forward to explain the formation of leks, and they can roughly be divided
in those that advocate that the males are the driving force and those that argue that the females are.
In this study we use data from a long-term study (1979-1998) of a capercaillie population at
Varaldskogen in south-east 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 arised 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
density of females are involved in a dynamic process of lek formation. We present results that
indicate that new leks are founded by young birds. Finally, we show that when new leks are formed
the spacing pattern of the individuals involved changes. This questions the method of using
comparisons between the position of female home ranges and established leks to infer how leks are
formed.

MODELLING FOR GROUSE CONSERVATION: ASSESSING POPULATION VIABILITY
AND MINIMUM VIABLE POPULATION SIZE
Volker Grimm & Ilse Storch
During the past few decades, grouse numbers have been declining throughout central Europe. Many
populations have disappeared, most remaining ones are small and scattered, and there is major
concern about their future. In this workshop, we aim to discuss the potential of population
modelling as a conservation tool. We first introduce the concept of population viability analysis
(PVA), and present a stochastic simulation model to assess the risk of extinction of grouse
populations and metapopulations in fragmented habitats. We deduce estimates of minimum viable
population (MVP) size and demonstrate how stochastic processes and deterministic factors such as
habitat quality, fragmentation, and predation pressure, may affect population persistence. Then, we
would like to initiate a round-table discussion on the potential and limitations of PVAs to support
management decisions in general and with regard to grouse in particular. Finally, we want to invite
workshop participants to feed our model developed for capercaillie Tetrao urogallus in the
Bavarian Alps with their own parameter estimates to explore outputs for populations in different
parts of the range.

ON HABITAT USE AND MATING SYSTEM IN THE SIBERIAN SPRUCE GROUSE
Franz Hafner, Alexander V. Andreev & Hartmut Gossow
The endemic Siberian spruce grouse Falcipennis falcipennis appears comparatively monotypic.
There is a considerable and interesting overlap of the spruce grouse range in Far-East Siberia with
the range of the Ajan spruce P. jezoensis: the needles of the spruce are the main winter food of the
bird. Even a small amount of spruce, only 2-5 % of all trees, will enable winter suvival. The grouse
use seasonally different kinds of habitat: in winter mainly dying spruce forests, or light larch stands
with spruce and fir in the mid layer are used, whereas in the vegetation period various types of larch
forest and open spruce stands with a well developed layer of herbs, grasses, and dwarf shrubs are
preferred. The main driving factor for habitat suitability are storms, the regular dying of spruce
stands, and forest fires.
Display grounds are mostly associated with topographical charateristics. They are distributed along
bogs, creeks, and crests, and they are seldom isolated but often located close to each other. Hill-tops
appear to be good arena-like sites. This grouse seems to prefer some openness in lekking as well as
in the initial form of a pair-bond system.

LANDSCAPE REQUIREMENTS OF SHARP-TAILED GROUSE IN MINNESOTA, USA
JoAnn M. Hanowski & Gerald J. Niemi
The sharp-tailed grouse Tympanuchus phasianellus occurs throughout the north central region of
North America. The species is associated with extensive open grass and brush landscapes and has
decreased in the southeast portion of its range over the past three decades, including marked
declines in the Great Lakes region. Habitat requirements for this species are well known and it is
easy to predict that habitat changes that include replacement of grass with brush, increase in brush
height, and increasing tree cover would result in a population decrease. Changes in these habitat
types and configurations are due to suppression of wild fire. This species has a large home range
and habitat requirements that change seasonally. Therefore, to properly manage for this species it is
important that landscape requirements are identified. We identified landscape characteristics within
a 1000m radius of active and defunct lekking sites. A regression model for the grouse was used to
predict the probability of occurrence on a landscape level in a geographic information system
platform.

THE DISTRIBUTION OF LEKKING BLACK GROUSE IN COMMERCIAL FORESTS IN
SCOTLAND
Susan L. Haysom & David Baines
The British population of black grouse Tetrao tetrix is declining probably as a result of habitat
degradation and loss. In Scotland, black grouse are primarily birds of heather moorland and birch
scrub habitats but the pre-thicket stages of commercial plantation forests are becoming an
increasingly important habitat.
The aim of this project was to identify the habitat and area requirements of black grouse inhabiting
afforested areas and to determine the relative importance of first and second (“restock”) rotation
components of forest structure.
Lek surveys were performed between 1996 and 1998 and habitat data were extracted from forest
stock maps. Associations between the distribution of black grouse leks and habitat features at
“patch” and “home range” scales are discussed. Our results show that patch size and the amount of
pre-thicket habitat within a home range are the predominant factors in the distribution of leks. This
has implications for forest management prescriptions.

ASSESSING EFFECTS OF CHANGING FOREST LANDSCAPE STRUCTURE ON GROUSE:
THE FINNISH WAY
Pekka Helle
There is a great demand of relevant information on the effects of landscape structure on wildlife in
boreal regions in order to minimize unfavourable effects of increasingly efficient land-use. Since
1989, the Finnish monitoring of grouse has produced exact locations of observations done along
transects totalling about 10,000 km annually. These data can be linked with geo-referenced National
Forest Inventory data with the aid of Geographic Information Systems, which allows one to run
complicated, multi-scaled spatial analyses. Basic landscape concepts and features measured include
level of fragmentation, isolation, connectivity, edge effect, landscape diversity, among others. We
are especially interested in the importance of the amount of old or mature forest in the landscape
with respect to the density and breeding success of grouse. Starting from habitat level (10-20 ha) the
scope has been widened up to tens of thousands of hectares in area. In many cases landscape effects
are more important in explaining habitat-level phenomena than habitat itself. For example,
landscape area best explaining variation in breeding success of grouse is much larger (100 km 2 )
than that used by a grouse individual (a few km 2 ) demonstrating that the relevant scale to be used in
the analyses should be that of predators, not that of the prey. Coming years will improve our
understanding ofthe dynamics of landscape-wildlife relationships. We shall be able, using a 10-15year
time window for example, to compare changes in densities and breeding success of grouse in
landscapes with different silvicultural treatment.

CAPERCAILLIE AND GROSS NATIONAL PRODUCT - AN EXAMPLE ON "REVERSED" U-
CURVE ?
Timo Helle, Virpi Alenius & Arto Naskali
According to the Finnish Game and Fisheries Research Institute, the relative density of the Finnish
capercaillie Tetrao urogallus population decreased between the years of 1964 and 1996 by 63
percent, and that of the black grouse Tetrao terix and hazel grouse Bonasa bonasia some less. It is
a common assumption that the reasons for the decline of different grouse species are the same, and
they are related to the changes in forest landscapes, including the resultant increase in predation. In
a tentative broader analysis, we found that the decline of the Capercaillie strongly correlated with
GNP, which doupled during the same period. That is understandable because especially in the
1960's and 1970's GNP was tightly dependent on forest sector, and the finding fits well with
"reversed" U-curve commonly used to describe the relationship between GNP and environmental
quality. Lowered quality of grouse habitats is probably associated to many other factors than clearcuttings
and resultant higher predator densities only. Among the other known but poorly quantified
mortality factors, the most influential may be 10-fold increase in traffic density, constructing dense
networks of forest roads, telephone and power lines, peatland diching (1.5 million kms since 1963)
and two new predator species etc. Many of these are known to reduce bird fauna locally rising the
question how grouse can survive, when "locally" is everywhere?

DISPERSAL FROM LEKS AND SPATIAL DISTRIBUTION OF MALE CAPERCAILLIE
DURING SUMMER IN RELATION TO DISPERSION AND FRAGMENTATION OF HABITAT
Olav Hjeljord, Per Wegge, Marina Ivanova, Alexander B. Beshkarev & Jørund Rolstad
In summer, capercaillie males are known to occupy richer habitats than during late winter and
spring. We compared movement of radio-equipped male capercaillie from leks to summer range in
three different regions in the boreal zone, viz. in south-east Norway (Varaldskogen), in north-west
Russia (upper Pechora basin) and in south-west Russia (upper Volga basin). The three areas
differed in dispersion of summer habitats and extent of fragmentation from logging. The hypothesis
tested was that dispersal from leks to summer range is related to these landscape variables. At
Varaldskogen (fine-grained mosaic and highly fragmented) mean distance to summer range was 1.5
km, but highly variable (0.8 to 10.0 km, N=44), in Pechora (coarse-grained and unfragmented oldgrowth
taiga) mean distance was 1.2 km (range 0.2-2.4 km, N=17), and in Volga (unfragmented old
spruce surrounded by large clearcuts) the distance averaged ca. 3.0 km (range ca 2-5 km, N=7).
The results were not consistent with the prediction. The spatial distribution of logged fragments and
non-territorial summer behaviour when males used small and overlapping ranges may have masked
a clearer relationship between dispersion of summer habitats and dispersal distance from leks.

ADAPTIVE RADIATION IN NECK ANATOMY FOR MATING CALLS IN MALES OF
GROUSE SPECIES
Ingemar Hjorth
Mating calls in birds are often highly adapted to overcoming attenuation due to the physical
environment and ambiguity due to the voices of other species. In grouse species (family
Tetraonidae), the females are very undifferentiated in the syrinx, trachea, oesophagus and feather
arrangement of the neck as well as in the sound production. The males, on the other hand, are
extremely species-specific in all these aspects. Anatomy analyses of deep frozen specimens of
North American and Scandinavian species have been compared with sound recordings from mating
calls and with films from displaying cocks. Sound production anatomy in the grouse family divers
in four groups. In the first group we recognize species with a ”normal” syrinx and without apparent
enlargement of the oesophagus (Bonasa, Canachites, Tetrastes and Lagopus). Tetrao urogallus
(and T. urogalloides?) have evolved a remarcable loop of the trachea. In the third group, we find
Dendragapus, Tympanuchus, Pedioecetes and Lyrurus tetrix with an inflatable oesophagus. In
Centrocercus, extreme oesohageal inflation is combined with heavy contractions of the trachea,
producing remarkable plops and visual signals. These results will be discussed as adaptations for
the acoustical efficiency of the mating sounds in species-specific habitats.

OCCURRENCE OF HAEMATOZOA IN GROUSE IN NORTHERN FINLAND
Tuula Hollmén, J. Christian Franson, Osmo Rätti, Pekka Helle, Einari Väyrynen & Ellis Greiner
During 1996 and 1997, blood samples were collected between May and October from four species
of grouse (Tetraonidae) at five locations in northern Finland to determine the host and geographic
distribution of blood parasites. Blood samples were collected from hunter-killed or live-captured
capercaillie Tetrao urogallus, black grouse Tetrao tetrix, hazel grouse Bonasa bonasia, and willow
grouse Lagopus lagopus, and blood parasites were identified from Giemsa-stained smears.
Leucocytozoon lovati, Trypanosoma avium, and a filariid nematode (microfilariae) were found in all
four grouse species studied. Haemoproteus mansoni was found in blood samples collected from
black grouse, capercaillie, and hazel grouse, but not in samples collected from willow grouse.
Prevalences varied from 64 to 100% for L. lovati, 0-17% for T. avium, 9-56% for microfilariae, and
0-20% for H. mansoni. Seasonal changes in prevalence of blood parasites were studied in captive
capercaillie and black grouse at Meltaus Game Research Station. Prevalences were lower during the
winter months (November-April): prevalences of L. lovati and microfilariae dropped from 100% to
80% and 50 to 40%, respectively. Furthermore, circadian patterns in blood parasitemia were
observed in capercaillie in Meltaus during July, 1997.

THE RADIATION OF GROUSE
Jacob Höglund
Sequence data from two protein coding (cytochrome b and NADH dehydrogenase subunit 2) and
two rRNA (12S and 16S) mitochondrial genes were analysed to reveal phylogenetic relationships
among grouse (Tetraoninae). Parsimony analyses showed that the grouse comprise a monophyletic
group with a closest relative within the Phasianidae. The hazel grouses Bonasa appear basal within
the Tetraoninae. Most previously recognized genera with the exception of Dendragapus group
together but some of the deeper branches within the clade are still uncertain (bootstrap support

WILLOW GROUSE POPULATION TRENDS FROM SWEDISH LINE TRANSECT COUNTS
Maria Hörnell & Tomas Willebrand
his study compares two related methods of distance sampling to determine willow grouse Lagopus
lagopus densities in the Swedish mountain range. One method uses the exact distances to each
observation and the other method only records if the observations are within or beyond a specified
distance (binomial method). The censuses have been made in the counties of Norrbotten (since
1994), Jämtland (since 1992) and Västerbotten (since 1998), and between 1,200-1,400 km are
censured each year. In each county 2-3 municipalities, and in each municipality 2-6 study areas are
censused, and the basic data collected are observed distances to willow grouse. Distances were
compared between years, counties, municipalities and areas. There were no differences between
municipalities in the same counties during the same year, except for one in the county of
Norrbotten. It was not possible to calculate densities at the area level because of too low sample
size. When comparing the three counties there were always larger observation distances in Jämtland
compared to Norrbotten. Estimates were fairly precise and gave coefficients of variation (CV)
between 20–40%. CVs varied more between lines in the same area than among counties,
municipalities and years. Unlike the method when exact distances are measured, the binomial
method requires fewer observers and larger areas can be sampled in a shorter time. However, there
appears to be a systematic underestimation of the density due to methodological difficulties.

DAMPED DYNAMICS, HARVESTING AND GROUSE POPULATIONS
Niclas Jonzén, Esa Ranta, Per Lundberg & Veijo Kaitala
A major challenge to ecology is the search for an understanding of temporal and spatial population
fluctuations in terms of the processes that produce them. This understanding should also constitute
the biological underpinning for a sound management of renewable resources such as game birds.
Even though all populations live in spatially structured and noisy environments, most harvesting
theory is built on assumptions of a spatially uniform and deterministic environment. The proposed
management strategies then has to cope with uncertainty caused not only by observation uncertainty
and the probabilistic nature of demographic rates, but also a considerable uncertainty about the
underlying processes.
A number of processes and mechanisms have previously been suggested to explain the cyclic
appearance of many grouse populations. The knowledge on eventual effects of hunting on
population stability is, however, still poor. We will demonstrate that harvesting per se may cause a
population, showing damped internal dynamics in the absence of harvesting, to become cyclic if
harvest rate is not temporally constant. Implications for population management will be discussed.

HYPOTHESES ABOUT THE SEX-RATIO OF CAPERCAILLIE -THE EFFECT OF
DIFFERENT ASSUMPTIONS ABOUT MORTALITY RATES
Annika Kangas & Sami Kurki
In southern Finland the relative amount of female capercaillies is much lower than in northern
Finland. The change is assumed to have happened within the last 20 - 30 years. Since the sex-ratio
of hatched juveniles has been similar across the country, the distortion of sex-ratio is due to
differences in mortality rates between the sexes. In this paper we present calculations about the
development of the capercaillie population with different assumptions about mortality rates. The
calculations are based on Leslie matrix approach, assuming different mortality rates for female and
male capercaillie. We examine how large differences are required to produce the observed sexratios,
what is the time required for the observed change and what is the effect of assumed
differences on the age structure of capercaillie population. We also examine the long-term
persistence of the population under such circumstances.

TESTING OF CRITICAL ASSUMPTIONS FOR THE USE OF DOGS IN LINE TRANSECTS OF
WILLOW PTARMIGAN
Leif Kastdalen, Hans Chr. Pedersen, Tomas Willebrand & Harald Steen
In order to census the density of game birds, line transects with dogs has often been used in recent
years. We tested some assumptions for using dogs (spring and autumn), where the measured
distance from the transect line to the observation point of the bird is the basis for calculating the
area of coverage (DISTANCE model).
The assumptions tested were: 1) The birds are detected at their initial location, not moving as a
response to the approaching census team; 2) All birds on the line are detected; 3) The detectability
of the birds is the same on both sides of the transect line and equal for the entire length of the line.
In addition, we tested if the detection function was depending on flock size. These assumptions
were tested on radio-collared willow ptarmigan, as well as reanalysis of over 4000 flush
observations reported from ordinary line transects. The testing showed that the method used fulfils
assumptions 1 and 2. Testing also showed that single birds and pairs have a different detection
function than groups of three or more birds. Assumption 3 was investigated using GPS receivers
mounted on the dogs during ordinary line transect sampling. Buffer analysis in a GIS was used to
calculate the area covered by the dogs.

BLACK GROUSE ARENAS IN THE BELGIAN HAUTES-FAGNES: LANDSCAPE
FEATURES AND MANAGEMENT PROPOSALS
Christine Keulen, Servais Houbart & Jean-Claude Ruwet
This study describes the black grouse Tetrao tetrix habitats in Europe and in the Belgian Hautes-
Fagnes, where the precise locations of the arenas have been mapped. Vegetation analysis were
carried out in the arenas within the radius of 150 m from the lekking centre, and vegetation maps
were designed. The vegetation assemblages differed considerably between different districts, but
consistent relationships were found between habitat structure and black grouse. Factor analysis was
used to investigate the relationships between the quality of black grouse arenas and their
characteristics: site topography, soil nature, landscape opening, vegetation structure, presence of red
deer and wild boars, distances to woods or bushes. This analysis provided evidence that tree
overgrowth, soil humidity, and presence of deer or wild boars are detrimental to black grouse
arenas. On the other hand, proximity of Ericaceae bushes and landscape opening are enhancing the
quality of the arenas. Based on the ecological requirements of black grouse, suggestions for a better
management of habitat resources are put forward.

TERRITORIAL AND MATING BEHAVIOUR OF CHINESE GROUSE AT LIANHUASHAN,
GANSU PROVINCE, CHINA
Siegfried Klaus, Yue-Hua Sun, Yun Fang & Wolfgang Scherzinger
The poorly investigated endemic Chinese grouse Bonasa sewerzowi inhabits the mountain
coniferous forests of Gansu, Qinghai, Sichuan, Yunnan and Tibet. In April 1995 and May 1997, we
obtained photographs, video and sound recordings in the Lianhuashan Natural Reserve (Gansu
Province), at elevations between 2,700 and 3,200 m a. s. l., which allowed first insights into
territorial (including fighting), advertising, feeding, and comfort behaviour. Chinese grouse did not
react to imitations and playback of hazel grouse B. bonasia or ruffed grouse B. umbellus songs and,
in contrast to the hazel grouse, does not sing. The main elements of display are flutter jumps
producing a deep instrumental sound and rarely territorial flights. The biphasic flutter jump is
composed of 12 ± 0.9 wing beats during the rising phase (duration 0.8 ± 0.1 s), followed by a short
gliding (0.14 ± 0.03 s) and landing with 3.4 ± 0.8 wing beats (duration 0.2 ± 0.06 s), often with a
turn (mean of 15 flutter jumps). The main difference from the flutter jump of the hazel grouse was
found to be the landing (6.0 wing beats, duration 0.3 ± 0.02 s). Males uttered a special offensive
vocal song during combat. Fighting and mating behaviour (not different from that of hazel grouse)
is described, based on video analysis. Fighting by territorial cocks involves standing in front of
each other in a highly erected pose, moving the head up and down, jumping in the air with wingbeat
attacks and pecking, followed by one male quickly chasing the other. Precopulatory behaviour
involves crouching and head-shaking by both sexes, and the male circling around the female with
tail and wing spread. During copulation the male bits the neck feathers of the female and spreads his
wings to the ground. Thereafter, the male runs after the female with an extremely erected tail and
wings dropping to the ground.
Our observations on morphology and behaviour of B. sewerzowi support Short's (1967) hypothesis
that the Chinese grouse more closely resembles the ancestral form than either of the other Bonasa
species. Activity was diurnal, territorial activity (flutter jumping) dominated in the morning and
foraging prevailed in the afternoon.

DEMOGRAPHY OF CAPERCAILLIE IN NORTHERN FINNISH BOREAL FORESTS
Paavo Kumpu, Pekka Helle & Veli-Matti Vallinkoski
According to wildlife censuses capercaillie Tetrao urogallus population has declined dramatically
during the last decades in Finland. The species is, however, still reasonably abundant and hunted in
most parts of the country. We studied demography of capercaillie in northern Finland by
radiotagging 22 adult male capercaillie in 1988-1992 and 34 females 1997-1999. The annual
mortality of both male and female capercaillie was 20%. Main cause of death was predation, mainly
by the red fox and goshawk. Breeding success was studied in 1997-1999 and it was especially low
during the last two years. Nest losses varied between 27 and 55%. Chick mortality was highest
during the first week after hatching and settled down after four weeks. The factors behind the chick
mortality are not known. The reproduction of our study birds was very low during two out of three
study years. Our results agree fairly well with the wildlife triangle censuses from other parts of
northern Finland.
It is not an exceptional result to have two or three consecutive seasons of low productivity during
the decreasing phase of the population cycle. In our case the result was unexpected, since the
previous peak year of capercaillie in Lapland was 1994, and we actually should be in the increasing
phase of the cycle. There are signs, however, that the fairly regular population cycles of grouse
seem to be disappering.

THE COST OF INCUBATION IN PTARMIGAN
Torgeir Lande, Hans Chr. Pedersen & Peter J. Pekins
The number of chicks produced by ptarmigan species can vary a lot among years. Changes in
predation rates is often the main source to this variation, but also weather conditions and the food
situation during the breeding period may cause some of this variation. In years with bad weather
conditions it might be too costly to raise big broods. Both the ambient temperature, amount of rain
and wind can increase the energetic cost of incubation for the hen. In addition to climatic factors,
the body condition of the hen can reflect how much energy she can spend on incubation. We wanted
to test if the energetic cost of incubation was dependent on climatic factors, the hen's body
condition, age, total clutch volume and timing of incubation. We also tested if metabolic cost varied
among birds with different nest cover, and finally if energetic cost during incubation was related to
number of fledged chicks per brood. We measured the metabolic rate of rock ptarmigan Lagopus
mutus and willow ptarmigan Lagopus lagopus hens in mid-Norway with the use of the doubly
labelled water method. Ptarmigan hens in this study had high energy expenditure during incubation
compared to other studies of tetraonids. These findings will be discussed.

CHARACTERISTICS OF DISTRIBUTION AND HABITAT STRUCTURE OF HAZEL
GROUSE IN KOREA
Woo-Shin Lee & Shin-Jae Rhim
This study was conducted to investigate the characteristics of distribution and habitat structure of
hazel grouse Bonasa bonasia from October 1998 to May 1999 in Korea. Hazel grouse were
observed from the responses to calls imitated by whistles in winter and spring seasons. Line transect
method was used in this study. Hazel grouse were distributed in most of high mountain forest areas
in Korea. Habitat utilization varies with altitude in the range of 300 - 1,200m a.s.l. At 600 - 900m,
the highest density of hazel grouse was observed, at low altitude in winter and high altitude in
spring. They inhabited mixed, deciduous, and coniferous forest areas. Especially, a lot of
individuals were observed in planted Japanese larch Larix leptolepis forests. The coverage and
density of understory vegetation are well developed in habitat of hazel grouse. The inhabitation of
hazel grouse seems to be related more to the development of understory vegetation than to the forest
type.

DISTRIBUTION, MOVEMENTS AND HABITATS OF SAGE GROUSE ON THE UPPER
SNAKE RIVER PLAIN OF IDAHO: CHANGES FROM THE 1950'S TO THE 1990'S
Kara M. Leonard, Kerry P. Reese, & John W. Connelly
Sage grouse Centrocercus urophasianus population levels on the Upper Snake River Plain of
southeastern Idaho declined significantly over the last 40 years. We investigated migration patterns
and seasonal ranges of these birds, and compared these data to data previously collected on birds in
the study area from the 1950's and 1960's. We also examined land use changes in the area that
occurred between 1975 and 1992. Migration patterns have not changed substantially since the
1950's. These grouse currently make annual round-trip movements of up to 125 km, and use an
annual population range of at least 276,000 ha. The only major change in land use on the area since
1975 has taken place in sage grouse winter range. Between 1975 and 1992, 29,762 ha of sagebrush
Artemisia spp. rangeland were converted to cropland, which represented a 74% increase. A total of
70,126 ha of sagebrush have been converted to cropland in the winter range of these grouse.
Regression analysis suggested a close relationship between habitat loss and grouse population
decline. Approximately 124,425 ha of privately-owned sagebrush on the study area could
potentially be converted to cropland, which likely would have serious negative implications for the
sage grouse population.

THE RESEARCH PROGRAMME FOR FINNISH GROUSE
Harto Lindén
Grouse research was the flagship of the organized Finnish game research for decades. This “state
monopoly” collapsed some 20 years ago, and an era of increasing cooperation started. Now grouse
scientists can be found in most universities and research organizations all over the country.
Astonishingly and luckily enough, these about 30 scientists are able and willing to cooperate across
the existing administrative borders within the very loose framework called “The research
programme for Finnish grouse”.
Monitoring game populations is the national duty of the Finnish Game and Fisheries Research
Institute, and the annual grouse censuses were started as early as 1963. The field work is made by
voluntary hunters, so we at the research institute strongly feel that we cannot “own” the data or its
applications, but our duty is to enhance its wise utilization. The data of the modern version of the
grouse censuses - wildlife triangle scheme - is accessible for everybody presenting a sensible
cooperative research plan. Wildlife triangles build the base for most of the grouse research in
Finland.
The high quality zoological research in Finland has “found” the grouse research resulting in
outstanding achievements especially in the fields of population dynamics and landscape ecology .
Professor Rauno Alatalo and the black grouse studies made “under his wings” represent the best
behavioral ecology of its kind. Professor Raimo Hissa with his groups has studied grouse species
from a physiological point of view with an understanding of the ecological background. The main
task of the programme - in addition to making good science - is to collect all the relevant
information and distribute it to those responsible for management.

NON-STABLE AGE DISTRIBUTIONS OF CYCLIC CAPERCAILLIE POPULATIONS IN
FINLAND
Harto Lindén & Esa Pienmunne
We studied the variation in age structure in Capercaillie Tetrao urogallus during population cycles
in Finland. More than 2000 skulls were collected from hunters for age determination, which based
on several ossification criteria. There are evidence that one-year-old females are poor breeders and
that the oldest females reproduce most successfully. We observed marked changes in the age
structure of the adult population in different phases of population cycles. The patterns were
different for the sexes owing probably to the difference in longevity between them. Age structure,
expressed as the average age of the female population, accounted for a half of the yearly variation in
the number of juveniles per adult female in August. We argue that the population age structure is an
important, but rarely documented, factor associated with Capercaillie population cycles.

EFFECTS OF HAND-REARING ON THE MORPHOLOGY AND PHYSIOLOGY OF THE
CAPERCAILLIE
Tuija Liukkonen-Anttila, Risto Saartoala, Tiina Mäkinen & Raimo Hissa
We studied morphological and physiological differences between hand-reared and wild capercaillie
Tetrao urogallus. Hand-reared birds differed from their wild counterparts in the mass of pectoral
muscles (P = 0.033). Captive males had heavier pectoral muscles than wild males, on the other
hand, these muscles were heavier in wild than in captive females. Wild birds had heavier heart (P <
0.001), liver (P < 0.001) and gizzard (P = 0.051) and longer small intestine (P < 0.001) and caeca (P
= 0.024). In wild capercaillie cytochrome c-oxidase activity was higher both in pectoral muscle (P <
0.001) and heart (P = 0.014) than in captive birds. In wild birds the protein concentration of pectoral
muscle (P = 0.039) and heart (P = 0.009) was higher than in captive birds. The chemical
composition of liver differed between wild and hand-reared capercaillie (P < 0.001). The uric acid
(P < 0.001) and T4 (P = 0.032) concentrations were higher in hand-reared than in wild birds. The
differences in digestive system and liver can result in lowered ability of captive birds to utilise
natural food nutrients. Decreased cytochrome c-oxidase activity of hand-reared birds can affect
their flying capacity and increase their vulnerability to predation. These facts can partly reduce the
hand-reared birds' survival after release. Our results on capercaillie support previous studies on grey
partridge, which show that hand-reared individuals may not be fit for living in the wild.

INFLUENCE OF THE CLIMATE ON THE DYNAMICS OF THE BLACK GROUSE
POPULATION IN THE BELGIAN HAUTES-FAGNES SINCE 1967
Michele Loneux, J.K. Lindsey & Jean-Claude Ruwet
Yearly spring demographic data on the varying population size of black grouse Tetrao tetrix living
in the Hautes-Fagnes are related to data on local meteorological conditions having a known
influence on the grouse survival during their life cycle. The data were modelled by Poisson stepwise
multiple regression. The dynamics studied cover thirty years of censuses. The predictions from the
best statistical model follow the observed values well: the weather explains a lot. The model
diverges more or less from the observed value in five of the 30 years, in one direction or the other.
When the predictions are far from the observed values, other variables than the climatic ones used
in the model must dominate. It is rather reassuring that this climatic model does not agree perfectly
with the observations because that would mean that all other factors are negligible, contradicting
many publications on the subject. It seems reasonable to ignore the effects of illness in favour of
factors involving ‘quality of habitat’ or ‘disturbances’, and also of ‘predation’, thought to be
negligible in a viable population, but which may have had more impact the last four years because
of the higher density of foxes and the lower density of grouse.

PATHOLOGY ENCOUNTERED IN CAPTIVE BLACK GROUSE BRED IN THE
EXPERIMENTAL GAME BIRD FARM OF THE NATIONAL CORP OF FORESTERS AT
ORECCHIELLA NATURAL PARC (TUSCANY, APENNINE)
Paolo Mani, Giacomo Rossi & Marco Bagliacca
The authors describe the pathology encountered in black grouse Tetrao tetrix bred in captivity
during the period 1988-1998 in the experimental game bird farm of the National Corps of Foresters
in the nature Park of Orechiella in the upper Tuscanian Apennines. The research consisted of
clinical and epidemiological examinations and anatomyhistopathological tests on birds found dead.
For laboratory diagnostic procedures, virological, bacteriological and parasitological tests were
performed on pathological material taken during necropsy and on fecal samples or other material
taken from the game farm. A total of 51 black grouse of various age were necropsied. The various
laboratory diagnostic procedures revealed the following pathologies:
- bacterial: Coligranulomatosis, aerosacculitis and E. coli-associated enteritis, spirochete-associated
typhlitis;
- protozoan: Trichomonas spp. and coccidia-associated enteritis, and/or typhlitis;
- parasitological: capillariasis of the oesophagus and crop;
- stressors: heat stroke, injuries and fractures, cannibalism, perforation of the gizzard, etc.
Clinical control in the game bird farm and samples taken from live animals often revealed
epidemiological risks and permitted suitable treatments and preventive measures to be taken.

EFFECT OF LANDSCAPE STRUCTURE ON HABITAT SELECTION AND SURVIVAL OF
BLACK GROUSE BROODS
Arto Marjakangas, Ari Nikula & Anu Koski
We studied the effect of landscape structure on habitat selection and survival of black grouse Tetrao
tetrix broods during 1990-93 in eastern central Finland at various spatial scales, using data on radiomarked
hens with brood. As landscape data we used classified satellite images. Landscape
characteristics around mean brood locations and random points were analysed in Geographic
Information System (GIS), using four different radii (200 m-2.5 km). Broods avoided landscapes
with mature forest (dominant habitat type) and agricultural land, and they preferred landscapes with
open bog, other open habitat including clearcut, and pine bog, especially at larger spatial scales.
Futhermore, the amount of habitat patches and edges were higher in brood landscapes than in
random landscapes. These results suggest that broods preferred fragmented landscapes, and that
edge effect may play an important role in their habitat selection. Considering all years, 68 hens
fledged one or more chicks, while 32 hens lost their broods totally by early August. However, we
found no important differences in landscape structure between successful and unsuccessful broods.

WHITE-TAILED PTARMIGAN REPRODUCTION IN YEARS OF EXTREME WEATHER
CONDITIONS IN COLORADO: WHEN TO OPT OUT, AND HOW TO COMPROMISE?
Kathy Martin & Karen L. Wiebe
Although ptarmigan have exceptional abilities to reproduce successfully in harsh weather
conditions, occasionally there are years that are extreme even for tundra habitats. We evaluate the
reproductive tactics and possible fitness costs imposed on white-tailed ptarmigan in such extreme
conditions. In 1995, there was heavy and persistent snow cover throughout May and June, and
breeding was delayed about 20 days later than the study mean. We observed no overlap in clutch
initation dates for first nests with the other 8 years of the study. Although early reproduction has
the highest success, all females attempted nesting. We looked for reproductive, somatic or survival
compromises females made to initiate reproduction under these extreme conditions. In 1995, hens
initiated clutches with lighter body mass, had smaller clutches, took more incubation recesses of
longer duration, and renested less. However, egg size, shell thickness and the usual female agedependent
relationships for initiation dates and clutch size remained. Given their short life span and
normally high reproductive failure, ptarmigan initiated breeding as soon as ground nesting was
possible. In extreme years, ptarmigan lay smaller clutches, and adjust incubation schedules to
compensate for reduced breeding condition. Opting out is not part of the ptarmigan reproductive
repertoire!

BLACK GROUSE BREEDING AND BIOMETRIC PARAMETERS IN SAMPLE AREAS OF
EASTERN ALPS (ITALY)
Silvano Mattedi, Fabio Perco & Andrea Cadamuro
Laying and hatching data of about 30 black grouse Tetrao tetrix nests, found in Friuli - Venezia
Giulia (Eastern Alps, Italy) are reported. The nests described have been localized by random
searches or radio-equipped individuals over a period of 12 years (1987-1998). 16 nests are
described according to the method suggested by P. De Franceschi (1993). This allows us to describe
the relevant vegetation characteristics, predation impact, hatching dates and success. 12 nests were
found within the sample area of Lanza - Val Dolce in the Province of Udine. Egg weights and other
measurements are reported. Biometric data of adult or young black grouse caught during the
research work are also summarized.

DISTRIBUTION OF CAPERCAILLIE IN RELATION TO AGE OF FOREST STANDS
Emmanuel Ménoni, Laurent Tautou, Yann Magnani, Jean Poirot & Laurent Larrieu
Many small-scale studies show that capercaillie Tetrao urogallus is associated with older forest
stands. In the Vosges (NE France), forests are intensively managed, mostly as even-aged stands.
The capercaillie population in the Vosges is declining, but still occupies a range covering about
25,000 ha distributed through 400,000 ha of forest. The objective of our study was to determine if
the present distribution of capercaillie at this scale could be explained by age and species
composition of forest stands. We found statistically that capercaillie frequently occupies stands
more than 120 years old, and less often younger stands.

WINTER MORTALITY OF PTARMIGAN IN SOUTH-WEST ICELAND
Olafur K. Nielsen
The Icelandic ptarmigan Lagopus mutus population has a c. 10-year cycle of numbers. Winter
mortality determines population change. Radio telemetry was used to follow ptarmigan in southwest
Iceland, from fall through spring for two winters, 1995-96 and 1996-97. Approximately 100
birds have been radio tagged. Mortality was high in fall and early winter but low after December.
The main mortality factors were hunting and predation. The hunting rates are such that this
population is probably over-harvested.

FORAGING BEHAVIOUR OF CAPERCAILLIE IN SPRING
Morten Odden, Per Wegge, Mats H. Finne & Sigrunn Eliassen
During March - end of May of 1997, 199 fecal samples of capercaillie Tterao urogallus (140 from
males and 59 from hens) were collected at Varaldskogen, SE Norway. Besides Scots pine Pinus
sylvestris, the birds exploited successively a larger proportion of ground foods, predominately
berries of cowberry Vaccinium vitis-idaea, berries of cranberry Oxycoccus quadripetalus, flower
buds of bog cottongrass Eriophorum vaginatum and bilberry stems Vaccinium myrtillus. Diet
composition differed significantly between the sexes during the whole period, with the largest
difference in May. The proportion of Scots pine was consistently higher among males, and
graminoids were almost solely detected among hens. However, the short-term temporal variations
were strikingly similar due to the influence of temperature, food plant phenology and food plant
abundance, which all affected the diet composition of both sexes. An exception occured in late
April, coinciding with the peak of mating. At this time male diet seemed to be less affected by
environmental factors, probably due to different territorial behaviour among the sexes. Studies of
radiomarked males revealed temporal changes in habitat use and daily movement as the diet
changed towards the end of spring. The relationship between size of male day ranges (N=20) and
their nutritional quality was also tested, but no correlation was found.

INFLUENCE OF METEOROLOGICAL VARIABLES ON BLACK GROUSE AND ROCK
PTARMIGAN POPULATIONS IN THE CARNIAN AND JULIAN ALPS
Fabio Perco, Paolo F. de Franceschi & Stefano Filacorda
The populations of black grouse Tetrao tetrix tetrix and rock ptarmigan Lagopus mutus helveticus
in the Alps are showing a dramatic decrease and the climate is changing. The 'Osservatorio
Faunistico del Friuli-Venezia Giulia- Udine' has studied the influence of monthly and seasonal
meteorological variables on the hunting bags (HB) obtained in the Carnian and Julian Alps, and on
the censuses of populations (CE) and the reproductive index (RI), for 12-26 years. The
meteorological variables were introduced in density-dependent models, with a stepwise procedure
(to enter: P

HABITAT SUITABILITY CHARACTERISTICS FOR CAPERCAILLIE IN CENTRAL
SWEDEN AND NORTH ITALY - A COMPARISON
Rainer Ploner & Hartmut Gossow
Habitat evaluation procedures (HEP) play an important role in wildlife management, especially
where overabundance, or declines, or (re)introductions are a major problem. For the capercaillie
Tetrao urogallus, Fennoscandian and Alpine habitats appear to be still the most suited ones in
Western Europe. But what are the most relevant habitatcharacteristics in these two different
landscape types and their respective forests?
This question was comparatively studied in a mixed mountain forest in the Alps of North Italy, and
in a boreal forest near Grimsö in Central Sweden: As most relevant appear (medium) canopy
closures of mainly coniferous forests (tree species less critical), Vaccinium cover and fructification
(although not generally essential), ground vegetation by height and coverage, and the successional
stage; exposition and incline could not be compared in this investigation because of the flat boreal
forest topography.

WHY IS ARTIFICIAL BREEDING OF TETRAO TETRIX MALE X TETRAO UROGALLUS
FEMALE HYBRIDS NECESSARY?
Jan Porkert & Karl-Heinz Schindlatz
Among the outcome of tetraonid crosses, hybrids between male black grouse Tetrao tetrix and
Capercaillie T. urogallus female are most common. In contrast to males of which some may be
fertile, the female hybrids are generally nonfertile. Thus, males of F1 generation can cross with
females of parent species resulting in the F2 generation. The real origin of different specimens of
assumed F2, F3 and F4 hybrids collected in museums need further verification. It is assumed, that
fertility increases in direction from F2 to F4 hybrids. When successive backcrossing from F2, F3
and F4 and further on were performed with capercaillie females, the offspring resembles more and
more the capercaillie.
Because of their slower growth during the first days of life, reduced body mass, intermediate diet
between parent species and lower food consumption, we assume a better survival of hybrid cocks in
fragmented and polluted habitats compared with capercaillie males. With help of backcrossing
experimens, the genetics determining morphophysiological characteristics and behavior as well as
the fertility of the female hybrids can be clarified. These experiments were planned by us and the
first experiment has been carried out.

SOCIAL RELATIONS BETWEEN CAPERCAILLIE AND HYBRID MALE TETRAO TETRIX
MALE X T. UROGALLUS FEMALE ON ONE CAPERCAILLIE DISPLAY GROUND IN S.
NORWAY
Jan Porkert, Roar Solheim & Arne Flor
Since spring 1996 we have observed 1-3 male hybrids and 1-2 male capercaillies on one
capercaillie display ground in Ovre Landvik in S. Norway. Both hybrids and male capercaillies
defended their respective territories when challenged. Female capercaillies chose where to mate
with the dominating male capercaillie. Although hybrids were fighting other hybrids, the
dominating hybrid concentrated his attention towards the display territory of the dominating male
capercaillie. Hence, the dominating male capercaillie showed great aggressiveness towards the
dominating hybrid. Feathers and skin found on the display ground in 1998 indicate that the
dominant hybrid was seriously injured, probably causing his death.

UNDERSTANDING DYNAMICS OF POPULATIONS: FINNISH GROUSE AS AN EXAMPLE
Esa Ranta
An overview is made of long-term fluctuations of grouse populations (capercaillie Tetrao urogallus,
black grouse Tetrao terix, hazel grouse Bonasa bonasia) in Finland. It will be shown that: (1) The
dynamics is cyclic for all species with 6-7 year period. (2) The populations of the three species tend
to fluctuate in synchrony over large areas both within species and across species. (3) In all species
the level of synchrony tends to go down against distance between the populations compared. (4) For
all species the skeleton of the population fluctuations is damped dynamics. (5) Recent data indicate
that the cyclic component of the Finnish grouse dynamics has become less prominent.
When damped dynamics is disturbed by random density-independent external forcing, the Moran
effect, cyclic dynamics can be sustained over long periods. The external forcing also synchronises
dynamics of populations with shared density-dependent structure. If the Moran effect is spatially
autocorrelated, or if there is small scale dispersal, the level of synchrony goes down against distance
among the populations compared. Distance-dependent dispersal topped off with increasing habitat
fragmentation (e.g. due to forestry practice) is capable of reducing the prominence of the cyclic
fluctuations.
Summarising, with the long-term quantitative population data courtesy of Finnish Game and
Fisheries Research Institute — combined with theoretical research — all the relevant features in
population dynamics of Finnish grouse can be understood.

INFLUENCES OF HUMAN ACTIVITY ON THE UTILIZATION OF SANDSAGE HABITAT
BY LESSER PRAIRIE-CHICKENS IN KANSAS
Robert J. Robel, Brent E. Jamison, Gregory C. Salter & Thomas L. Walker, Jr.
Lesser prairie-chickens Tympanuchus pallidicinctus in North America originally inhabited vast
shortgrass prairies of Texas, New Mexico, Oklahoma, Colorado, and Kansas. Populations of these
prairie grouse have declined range-wide because of habitat alterations, mainly conversion of
suitable habitats to rowcrop agriculture and grazing by livestock. Large remnants of sand sagebrush
Artemisia filifolia prairie in southwestern Kansas support viable populations of lesser prairiechickens.
However, human activity is gradually altering the pristine nature of these habitats. We
initiated this study in 1996 to determine habitat preferences and movements of lesser prairiechickens
in fragmented sandsage prairie. Over 250 lesser prairie-chickens have been trapped, 115 of
which have been fitted with radio transmitters and monitored daily. A data set of approximately
18,000 radio-determined locations of lesser prairie-chickens is being used to assess the impact of
roads, oil/gas wells, gas compressor stations, an electric generating plant, and inhabited buildings
on the utilization of sandsage habitat by lesser prairie-chickens. Initial analyses suggest some
negative impacts of human disturbance extend up to 1 km from the site of the activity. We believe
our data will be applicable to other grouse species sensitive to human disturbances, e.g., sharp-tailed
grouse Tympanuchus phasianellus, capercaillie Tetrao urogallus, and black grouse Tetrao tetrix.

EVOLUTION OF BLACK GROUSE HABITATS AND FLUCTUATIONS OF COCKS
NUMBERS FROM 1966 TO 1999 IN THE BELGIAN HAUTES-FAGNES
Jean-Claude Ruwet, Serge Fontaine & Servais Houbart
Black grouse Tetrao tetrix is an endangered species in Central and Western Europe, and the Hautes-
Fagnes plateau (summit of Belgium, altitude 500- 694 m), a mixture of peat bogs and moorlands
(4,000 ha out of 5,000 ha under protection), is one of the last and probably its most western
sanctuary on the continent. The number of cocks censused in spring since 1966 has fluctuated
between an exceptional peak of 200 in 1971-72 and a low of about 25 individuals in the mid sixties
as well as at present, with less pronounced increases and declines meanwhile. Thanks to important
recruitment years in the late 1960s, the very low population grew up to 198 males in 1971; then it
collapsed to 42 cocks in 1976. During the following 20 years, it fluctuated between about 80 and
less than 40 males. The latest surveys revealed less than 30 cocks. The present situation is thus
more critical than in the mid 1970s, and it was even worsened by the spontaneous change of some
parts of the Nature Reserve from attractive moorlands to less attractive woodlands. Considering that
the populations are vulnerable to unpredictable weather dependent variation in recruitment, hard
work and strict policies are urgently needed to restore habitats and to increase populations, bearing
in mind that a stock of 50 cocks is the lower limit to face hazards of any kind during low density
periods.

SELECTIVE HUNTING OF PARASITIZED GROUSE
Osmo Rätti, Pekka Helle & Tuula Hollmén
Caecal parasites have been observed to make red grouse vulnerable to mammalian predation during
breeding. This has been confirmed by a study where a trained pointing dog was less likely to find a
nest of a female with experimentally reduced parasite burden. Here we study whether intestinal
parasites make grouse (capercaillie Tetrao urogallus, black grouse Tetrao tetrix, and hazel grouse
Bonasa bonasia) vulnerable to hunting aided by a dog in Finland. We collected intestinal samples
from hunters during hunting season and asked whether a dog was used. Parasites were examined at
laboratory and the data were analysed by hierarchical log linear model selection. Nematodes were
predominantly parasitizing adult grouse whereas cestodes were parasitizing juveniles. Therefore,
the tests were performed separately for adults and juveniles. In adults, the use of dog selected
species, which were disproportionally infected by nematodes. In juveniles the use of dog selects
species and individuals infected by cestodes. In both age classes, the bag is most often capercaillie
and less often hazel grouse when the dog was used. Among juveniles, the use of dog yielded most
often a bag that was infected by cestodes. Thus, either the dog found infected grouse easier or the
dog was better able to hold the infected grouse at halt. Most of the dogs used in hunting were
barking bird dogs (Finnish spitz).

POPULATION DYNAMICS OF CAPERCAILLIE IN THE JURA MOUNTAINS
Sebastien Sachot, Nicolas Perrin & Cornelis Neet
Capercaillie populations are declining in the French and Swiss Jura Mountains. To understand the
factors influencing their dynamics, we are building a spatially structured population model in order
to perform a population viability analysis.
The model considers the dynamics of the female sex, using estimates of vital rates collected from
these populations over 20 years of field studies. It integrates spatial references through dispersal
between populations, modelled as a negative exponential function that also takes into account the
size of the target populations. Demographic stochasticity influences the proportion of reproductive
females, fecundity, number of dispersers, as well as the survival probability of both juveniles and
adults. Environmental variance is taken into account for each population, which underlines the
asynchronisation of local dynamics.
Preliminary results indicate that the 40 populations forming the metapopulation face a serious threat
of extinction in the next 100 years. A sensitivity analysis performed on the vital rates shows that
female survival is the most important parameter influencing population dynamics.

COMPARING GEOGRAPHIC DISTRIBUTION AND GENETIC DISTANCE IN
CAPERCAILLIE AND BLACK GROUSE: GENETIC MARKERS AS A TOOL FOR LARGE
SCALE CONSERVATION
Segelbacher Gernot & Ilse Storch
Capercaillie Tetrao urogallus and black grouse Tetrao 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 (meta)population dynamics and persistence. To identify
conservation priorities, it is important to know if and to what extent exchange between local
populations can be anticipated. To provide this information, we investigate the dispersal patterns of
grouse using non-invasive genetic methods. We assess the genetic differentation of spatially distinct
populations using microsatellite loci analyses based on DNA extracted from feathers. We expect a
correlation between geographic and genetic distance. Our approach allows to identify critical
geographic distances beyond which demographic connectivity between populations is not assured.

TELEMETRY STUDY OF RELEASED BLACK GROUSE IN SCHLESWIG-HOLSTEIN,
GERMANY
Christiane Seiler
Due to the rapid decline of the black grouse populations in northern Germany during the late 70’s,
hunters in Schleswig-Holstein founded a “Black Grouse Association” that aimed to prevent the
regional grouse population from extinction. Among other measures the association tried to restock
the grouse population by annual releases of captive reared birds. In 1996/1997 a scientific project
was conducted to assess the effects of the measures taken so far. It included a telemetry study of the
black grouse released in 1996, which is presented in the poster.
The region monitored by the association contains about 3.500 ha of fragmented moorland, suitable
as black grouse habitat. In autumn 1996, three moorland patches were chosen as release areas for 58
birds, out of which 29 were radio tagged. Locally, the radio tagged grouse stayed within the
moorlands and were seldom tracked on surrounding pastures, but individual birds dispersed up to
17 km. Only one cock was known to survive until spring display 1997; the radio signal of 4 birds
was lost. As expected, predation was the main cause of death. In total, the proportions of avian
(goshawk) and mammal (red fox) predation were almost equal, but differed between the release
areas.

THE DISPLAY OF BLACK GROUSE COCKS IN A MIRE LANDSCAPE IN EASTERN
SWITZERLAND: INFLUENCE OF LANDSCAPE STRUCTURE AND HUMAN ACTIVITIES
Ursula Sieber
In a mire landscape of national importance, distribution, choice of habitat and behaviour of the
black grouse Tterao tetrix cocks on the display were studied in spring. I wanted to elucidate
whether or not human utilization of the area had any effect on the display of black grouse cocks. In
the whole area, habitat quality is not very suitable for the black grouse. Accordingly, the area is
sparsely populated. Compared to other sites, the black grouse cocks display in this area in very
steep locations. The reason for this may be the great steepness at the tree line or the fact that all
flatter regions are used by humans (tourism, agriculture, military). At the beginning of the mating
season the duration of the display was about the same at a site in the military zone and in an area
used by tourists. At the end of May, however, when tourism increased, the cocks in the military
zone displayed for longer periods than those in the tourist region. In the present situation of this
area, tourism is affecting the black grouse more than military activities, and in the future tourism
will probably be the more important source of danger.

INCREASE IN GROUSE HUNTING YIELD IN EARLY 20TH CENTURY NORWAY: WHAT
HAPPENED?
Chr. Anton Smedshaug
In Norway there was a period of high willow grouse hunting yield and non-cyclicity in the early
century (1906-1914). At the same time a Norwegian Program for Extermination of Predators
(NPEP) was going on, which means that additional predator species were bountied (from 1900) and
the intensity was increased. The increase in willow grouse hunting yield has been attributed to this
predator program. However, others have argued that an unusual case of several peak rodent years
coinciding with the NPEP caused the high grouse hunting yield, as predicted by the alternative prey
hypothesis. These peak rodent years are said to be caused by predator extermination.
In order to address these hypotheses, time series of vole peak years (1885-1914), grouse hunting
reports (1885-1914 and 1900/1-1914), grouse export statistics (1885-1914) and predators bountied
(1885-1914) from the same period, were analyzed for South, East and Central Norway.
The results make it likely that the registered effort in predator reduction on regional scale was not
sufficient to cause the increase in small game. In two regions with both predator reduction and
unusually high rodents number there was a significant increase in grouse hunting yield. But in the
third region where it was just predator reduction and not high rodent numbers in successive years,
there was no increase in grouse hunting yield.

GROUSE POPULATIONS, MANAGEMENT AND MOORLAND VEGETATION
Adam Smith
Management for the production of red grouse Lagopus lagopus scoticus comprises three activities:
habitat manipulation, predator and disease control. It has been proposed that improving moorland
habitat quality may increase the density and productivity of red grouse, thus offsetting predation by
raptors. In order to determine how grouse abundance changes with habitat we studied the
relationships between moorland habitat and red grouse density and productivity on eighty-three
sites on 61moors managed for red grouse in upland Britain.
The location of sites, their altitude and whether or not they were managed explained 57% of the
variation in red grouse density in spring. Once these factors were controlled for, heather phosphorus
content was the only habitat variable that explained a significant amount (5%) of variation in spring
density. Grouse had greater breeding success on managed sites. In addition, breeding success was
higher on sites with higher July temperatures, higher heather phosphorus content, more cotton grass
and reduced amounts of young heather shoots. These five variables accounted for 50% of the
variation in grouse breeding success. Heather cover had no effect on either grouse density or
breeding success, though all but one site had more than 40% heather cover.
Similar data were collected from 14 sites in the north Pennines that had low heather cover. On these
sites, a strong negative relationship indicated that grouse densities were lower where heather cover
was less. This study suggests that above 30% heather cover, the physical characteristics of moors,
predator and disease control to have a greater effect on red grouse abundance than habitat.

RENESTING IN CAPERCAILLIE
Torstein Storaas, Leif Kastdalen & Per Wegge
During 1979-88, hatching dates of 234 capercaillie nests and broods were observed or estimated in
southeastern Norway. Between 8.8 and 87.3% (mean 35.7%) of the females that had their first nest
depredated, renested. The probability of renesting was highest the year before the small-rodent
peak. During the year of highest renesting, the predation rate decreased in the course of the summer,
but in the small-rodent crash year the predation rate increased. The weight of adult females on leks
was higher in the two years of highest renesting rates.
Renests contained fewer and smaller eggs than first-nests, and during incubation renesting females
left the nest more often and for longer periods than firstnesters. Weight of yearling hens at leks was
lower, and they took more recesses during incubation than older females, and no yearling was found
to renest. Over a six-year period, 21 percent of all successfully hatched broods came from renests.

STATUS, THREATS, AND CONSERVATION OF GROUSE WORLDWIDE: AN OVERVIEW
Ilse Storch
A major task of the Grouse Specialist Group is to compile an Action Plan to communicate the status
of the grouse, major threats to their long-term survival, and recommendations for their conservation.
In preparation of the Action Plan, a questionnaire regarding the status, population trend, threats, and
conservation needs was sent to researchers, state agencies, and NGOs in most countries within the
range of the 17 grouse species worldwide. About 100 completed questionnaires from >30 countries
were returned. Additionally, the recent literature was searched and data on conservation and legal
status were compiled. The data were analysed in relation to geographic distribution and frequency.
Three species, the Caucasian black, Chinese, and Siberian spruce grouse, and several subspecies are
listed as globally threatened. These taxa should be considered global priorities for grouse
conservation. At a national level, populations at the southern edge of a species’ range and in densely
populated countries are most often red-listed. Habitat degradation due to human land use activities
was named as the major threat to grouse viability. Accordingly, most correspondents recommended
habitat preservation as the most urgent conservation need. Additional threats such as exploitation
and human disturbance were regionally felt to be critical.

DENSITY VARIATION AMONG ALPINE CAPERCAILLIE POPULATIONS AT MULTIPLE
SPATIAL SCALES: HOW MUCH IS EXPLAINED BY THE HABITAT?
Ilse Storch
he paper presents a Habitat Suitability Index (HSI) model for the capercaillie Tetrao urogallus in
the Bavarian Alps, Germany, and explores effects of spatial resolution on model performance and
applicability in conservation. The model was developed based on local-scale habitat preferences
revealed by telemetry. To validate the model, habitat suitability and observed frequency of
capercaillie use from six different areas were compared at different spatial scales. Among 0.04 ha
sample plots, the model achieved a close correlation (Taub = 1.0) between HSI scores and the
probability of capercaillie use. At larger scales, i.e., 100 ha and 400 ha grid cells and 2000 ha study
areas, respectively, HSI scores explained about 40% of the variation in capercaillie abundance.
These results illustrate that there may be a discrepancy between a HSI model’s power to predict the
distribution of a species over habitat types, and its limited potential to predict its abundance at
larger scales. Two points need to be considered in model applications: 1) wildlife-habitat
relationships rarely are consistent throughout all levels of spatial scale; and 2) HSI models typically
are restricted to habitat-related components of carrying capacity, although populations may be
influenced by a variety of other factors as well.

THE DECLINE OF A BLACK GROUSE POPULATION IN A FRAGMENTED MOOR
LANDSCAPE
Egbert Strauss
The objective of the reintroduction project "Wurzacher Ried" (1978 - 1993) was to establish a
stable black grouse Tetrao tetrix population after the extinction of the autochthonous population in
1978 in the scattered moorlands of "Oberschwaben" (South-Germany). Losses and changes in
habitats were quantified in ten peat bogs on the basis of historical maps and aerial photos as
potential causes of extinction. The diet (162 fecal samples from July 1988 - December 1991) and
the habitat utilization of released and radio tracked black grouse were analyzed to estimate the
carrying capacity on the basis of food and habitat resources. There were no ethological or parasitic
particularities to be noticed in the released black grouse.
The early peat exploitation in the 19th century was intensified in the beginning of the 20th century.
Litter meadows, surrounding the rain bogs, were meliorated into intensively used pasture land. The
drained and peat cutted areas developed into heath and dwarf shrub heath. In the middle of the 20th
century draining and peat cutting were discontinued, so that pristine rain bog areas could be
preserved in a large extent. Approximately 1,200 ha of the investigated area (3,600 ha) are optimal
habitats for black grouse. The patchy distribution of the moorlands in the landscape and the habitat
resources in the peat bogs are comparable to the time period of the highest density of black grouse.
The decline in the larger bogs is not due to changes and losses of habitats in contrast to the small
bogs. The main cause of the extinction and the failure of reintroduction is probably the impact of
predators from the surrounding areas into the habitat islands, as a consequence of edge effects and
increasing predator density.

DISTRIBUTION AND STATUS OF CHINESE GROUSE
Yue-Hua Sun
After the first discovery of the endemic Chinese grouse Bonasa sewerzovi in 1873 in Gansu, the
distribution area in China has enlarged due to new recordings, and in 1997 the species was found in
Jiali County in Tibet. Its present distribution should be defined to include Gansu, Qinghai, Sichuan,
Yunnan and Tibet, corresponding with the distribution of mountainous conifer forest in central
China, with the altitude between 2400 to 4300 m. The habitat requirements of Chinese grouse
should include two main factors: the one is coniferous trees serving cover, the species being spruce,
fir or cypress in different areas; the other is deciduous trees and shrubs for food, mainly willow
Salix spp. according to latest studies. Checking the forest distribution map in China, we found that
historically most Chinese grouse habitats were connected with each other in the past. However, due
to intensive cutting of virgin forest in past decades, the present distribution is much more isolated
and fragmented. The bird is listed as an endangered species by China Red Data Book. According to
our ecological study in Lianhuashan, second growth forest could serve as winter habitat for Chinese
grouse, and the population density in Lianhuashan, where forest was slightly disturbed in the past,
is higher than that in old conifer forest. We suggest that although the Chinese grouse could stand for
a moderate scale of selective cutting, large area of clear cutting is disastrous. Conservation
suggestions on forestry management and designation of protected areas are given.

DETERMINATION OF CHINESE GROUSE HABITAT AT LANDSCAPE SCALE IN THE
LIANHUASHAN MOUNTAINS
Yue-Hua Sun, Siegfried Klaus, Yun Fang & Wolfgang Scherzinger
he endemic Chinese grouse Bonasa sewerzowi inhabits isolated remnants of the mountainous
coniferous forests of Gansu, Quinghai, Sichuan, Yunnan and Tibet in China. The decline and the
increasing isolation of this endangered Red data book species is caused naturally by the increasing
aridity of the area following the rise of the Himalaya and of the Tibetan Plateau, and artificially by
agriculture, logging and deforestation of mountain forests.
Male Chinese grouse hold territories in the mountainous spruce-fir forests mixed with deciduous
trees (mainly willow and birch), and all nests we found (n=34) were in this habitat type. This
suggests that spruce-fir forest is the key habitat for the survival and reproduction of this bird.
Satellite images were used to determine the distribution of spruce-fire forests in the Lianhuashan
Mountains, including one reserve and two forestry farms, totally an area of 400 km². The Chinese
grouse population in this area is isolated with the nearest neighboring population 30 km away. The
size of forest islands, the distance of open land barriers, as well as the presence of possible
movement corridors were evaluated visually using the images. Hopefully, our results will be used
by local forestry managers to link habitat islands with reforested corridors, essential also to other
wildlife.

STATUS AND MANAGEMENT OF THE GREATER PRAIRIE-CHICKEN IN NORTH
AMERICA
W.D. Svedarsky, R.L. Westemeier, R.J. Robel, S. Gough & J.E.Toepfer
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 those regions where climatic and/or soil factors
favored the retention of grassland. Historically they probably occurred in 20 states of the United
States and 4 Canadian provinces but presently occur in 11 states and no longer occur 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 those 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.

PREDATION IMPACT OF THE GOSHAWK ON GROUSE
Risto Tornberg
I studied predation of the goshawk Accipiter gentilis in 1988-1998 in Oulu district in northern
Finland focusing on the goshawk´s impact on four grouse species: willow grouse Lagopus lagopus,
black grouse Tetrao tetrix, capercaillie Tetrao urogallus and hazel grouse Bonasa bonasia. Grouse
made up 68% by biomass of the diet during nest-building and incubation period, 50% during
nestling period and 44% during fledgling period. Black grouse was the most important grouse as
prey (15-40%), others sharing 5-10% each. Predation impact of the resident goshawks during the
breeding season was in average 22% (5-54%) for willow grouse, 12% (8-21%) for black grouse, 2%
(0-5%) for capercaillie, 17% (7-25%) for hazel grouse and 10% (2-27%) for grouse chicks. Average
yearly predation impact was calculated only for years 1991-1995 when data of winter predation
were available. On yearly basis, willow grouse made up about 2%, black grouse 30%, capercaillie
4% and hazel grouse 6% in the diet. Calculated from August, when grouse censuses are made, to
the next August, breeding hawks and their offspring killed 24% of willow grouse, 15% of black
grouse, 2% of capercaillie and 20% of hazel grouse. Taken into account an estimated portion of
non-breeding birds (1/3 of the resident population) the respective percentages were 36%, 24%, 3%
and 30%. Goshawk´s share of the total yearly losses for black grouse were 30% (breeding birds)
and 46% (breeding + non-breeding birds), for hazel grouse 43% and 66% and for capercaillie 6%
and 10%, respectively.

THE DENSITY OF THE HAZEL GROUSE AND ITS HABITATS IN THE CENTRAL PART OF
ESTONIA
Ene Viht
The study area (26 km²) is situated in Kaarepere forest, in the central part of Estonia. The censuses
of the hazel grouse Bonasa bonasia were carried out with the whistle in spring and autumn during
six years within the period 1986-1998. All whistling birds were assumed to be territorial. The sites
of the birds, who react to the whistle were mapped. To characterize the habitats of the hazel grouse
territories the following parameters were used: forest site types, index of forest productivity, tree
species composition, age, stocking density, height of the main stand, second growth and
underwood.
The density of the hazel grouse varied between 4.6 and 15.6 ind./km². The hazel grouse has wide
range of habitat requirements; it uses all the different groups of forest site types but prefers some of
them: Hepatica, Oxalis, transitional bog, drained swamps and raised bog forest site types. The hazel
grouse prefers especially mixed forest, although the species occurs also in coniferous and deciduous
forests. The species lives in forests with different age, stocking density and height, but prefers older
forests and forests with stocking density of 0.7- 0.8.

CAPERCAILLIE IN FRAGMENTED FORESTS: DOES DISPERSAL STABILIZE BREEDING
POPULATIONS ON A LARGER LANDSCAPE SCALE?
Per Wegge & Jørund Rolstad
Recent focus on "source-sink" processes and metapopulation theory has rendered the traditional
approach of studying "local" populations less useful for understanding the population dynamics of
species. We present empirical data on demography and seasonal movement and dispersal of a local
population of capercaillie at Varaldskogen in south-east Norway during 18 years based mainly on
radio-equipped birds (N=200). The ca 80 km² study area is embedded in > 1000 km² contiguous
forested capercaillie habitat, but is highly fragmented due to industrial logging. Preliminary
analyses show that the study population has become a sink, maintained by a net import of females
from surrounding forests. Due to strong breeding site fidelity, dispersal among adults of both sexes
is minimal, although seasonal movements up to 10 km may occur. Data on natal dispersal indicate
that this is independent of density and longer among juvenile females, both during first autumn and
during spring before first breeding. The surrounding habitat of the sink within the periphery of
potential natal dispersal distance appears to be of no better quality, which raises the question of
location and demography of the donor "population(s)".
Following a review of published information on natal dispersal in this and other related polygynous
species, we examine probable explanations for the observed pattern.

FOREST DEVELOPMENT AND CAPERCAILLIE HABITATS WITHIN THE AUSTRIAN
ALPS
Hubert Zeiler, Monika Breuss & Hartmut Gossow
In opposition to declining populations of capercaillie Tetrao urogallus in Central Europe, a stable
population development is postulated for this species for Austria within its alpine area. To test the
relationship between habitat characteristics and capercaillie density, we compared the numbers of
displaying cocks in spring with the data of the Austrian Forest Inventory in 46 hunting districts. The
study area is the alpine (montane - subalpine) region within Austria. Significant positive
relationships can be shown between capercaillie density and the amount of protection forest within
the overall Austrian alpine area. Also higher elevations as well as old forest stand structures are in a
positive relationship to capercaillie occurrence on a landscape scale. On the other hand, no
relationship could be detected between capercaillie as a forest species and the overall amount of
forested area, which implies a clumped or fragmented distribution pattern of the species. This, and
the circumstance that a relationship between the occurrence of the species and forest property types
exists, leads to the conclusion that the still existing belief in a quite stable capercaillie population in
the Austrian Alps can not any longer be supported without strong reservations.

HABITAT SELECTION OF HAZEL GROUSE - THE INFLUENCE OF HETEROGENEITY
AND SPATIAL SCALE
Johan Åberg, Gunnar Jansson, Jon Swenson & Grzegorz Mikusinski
The habitat selection of hazel grouse Bonasa bonasia populations, based on censusing of males,
was studied in a forest reserve in Sweden during 11 years. The objective of the study was to analyze
the effect of stochasticity on hazel grouse habitat selection on several scales and to test the
applicability of the commonly used forest stand descriptions, for management of the species. The
censusing was performed in spring and in autumn each year at 197 censusing plots. The reserve,
dominated by coniferous forest, was divided in to a richer part and a poorer part based on
vegetation. The entire reserve was described by forestry stand descriptions, and a detailed
vegetation description was performed for the richer part. No effect of habitat type was found to
influence hazel grouse occurrence in the richer part using the stand descriptions, but a negative
effect of coniferous trees was evident for the occurrence of hazel grouse in the poorer part. The
occurrence of hazel grouse was significantly positively influenced by increasing amount of shrubs
in the western part at the largest scale, using the detailed vegetation description. A density
dependence pattern for the occurrence of hazel grouse was evident, but no habitat variable differed
significantly between type of grid-cells. Management of hazel grouse would benefit of improved
stand descriptions, especially in relatively suitable areas, and also by use of a landscape ecological
perspective.