Grouse News 41.pdf - Galliformes-sg.org

GROUSE NEWS 

Newsletter of the Grouse Group of the 

IUCN/SSC-WPA Galliformes Specialist Group 

Galliformes Specialist Group 

Issue 41 May 2011

Grouse News 41 Newsletter of the Grouse Group 

Contents 

From the Editor 3 

From the Chair 3 

Conservation News 

Urgent measures for conservation of the Cantabrian capercaillie Tetrao urogallus cantabricus in its 

habitat in the Cantabrian mountain range 

Research Reports 

Effects of large-scale human land use on capercaillie Tetrao urogallus (L.) populations in Finland - 

A short summary of the PhD Thesis 

5 

Why the capercaillie population Tetrao urogallus (L.) in the West Carpathians decline? 6 

Hazel grouse in open landscapes 13 

Investigating the importance of thermal refugia for white-tailed ptarmigan at the southern extent of 22 

their range 

Is GPS and satellite telemetry an option for grouse research? 24 

GPS satellite telemetry used on capercaillie Tetrao urogallus 25 

Experiences of satellite tags on rock ptarmigans 26 

Book Reviews 

Tetraonidae and Phasianidae of the USSR – Ecology and Morphology 28 

Conferences 

The 12 th International Grouse Symposium postponed 1 year 30 

Reminder – 29 th Prairie Grouse Technical Council (PGTC) Meetings October 4 - 6, 2011 31 

6 th European Conference Black Grouse Endangered Species. First announcement 31 

Recent grouse literature 32 

Erratum 

Error in the article on black grouse in issue 40 37 

Snippets 

The effect of climate change on grouse populations in Norway 38 

Amazing species required 38 

Back Page Picture 

Japanese rock ptarmigan 

2 

4


From the Editor 

First, the disasters of the earthquake, tsunami, and the subsequent nuclear crisis have been troubling news 

to the entire world, and we send our condolences to all the citizens of Japan. We wish the best for a rapid 

and complete recovery for this fine country and its people. As for the necessary postponement of the 11 th 

International Grouse Symposium, it is ONLY a conference, and most of us have plenty of other 

opportunities to exchange information and ideas. We will most certainly have a dynamite conference in 

2012. Thank you, Hiroshi, for the effort put into planning the symposium, and for having to foresight to 

know that waiting another year would be best for everyone. 

This issue of Grouse News contains several updates and summaries of recent and ongoing Capercaillie 

research, plus additional articles on Hazel Grouse and White-tailed Ptarmigan. Additionally, this issue 

contains some discussion and early results of using GPS technology for studying grouse. As this 

technology continues to improve, we will likely see more wide-scale usage by researchers everywhere, 

and regular updates on its applicability from those all that are currently using GPS units would be 

beneficial to many other researchers. The proceedings of the 2008 IGS are in the final stages of 

completion, and for sure many of our members are currently reviewing the page proofs, so the final 

project should be available before our next Grouse News. And finally, don‟t forget these upcoming 

grouse conferences: The 29 th Biennial Prairie Grouse Technical Council meeting will be held at Hays 

Kansas, USA, 3 October to 6 October 2011 and the XXX th IUGB and Perdix XIII Congress will be held 5 

September to 9 September 2011 in Barcelona, Spain. In September 2012 the 6 th European conference on 

black grouse will be held in Sweden. 

Tor Kristian Spidsö, Editor Grouse News, TKS.Grouse@gmail.com 

Don Wolfe, Co-editor North America 

G. M. Sutton Avian Research Center, University of Oklahoma, P.O. Box 2007, Bartlesville, OK 74005, 

dwolfe@ou.edu 

From the Chair 

Grouse researcher should be familiar with stochastic events and their potentially far-reaching effects on 

system dynamics. Yet, we were not well prepared to anticipate circumstances that forced us to derive 

from our triennial cycle of gatherings held dear for more than 30 years since the first symposium back in 

1978. As you will know by now, due to the earthquake, tsunami, and related nuclear disaster in Japan in 

March, there will be no International Grouse Symposium in 2011. The decision to postpone the 12 th IGS 

in Japan by one year was made in agreement between the local organisers in Japan and the Grouse 

Group‟s committee members. Even as Fukushima appears to be out of the international news by now, it 

will take a while until the situation can be considered under control, and an international event such as the 

IGS safe to plan and hold. 

I am most grateful to our colleague and IGS organizer Hiroshi Nakamura and his team for the work 

they have invested so far, and the difficulties they go through, in maintaining their commitment and 

dedication to invite the IGS to Japan. We now plan for the IGS to be held in July 2012. All Symposium 

plans will be kept, and a new registration process will be started in time. Please spread the information 

among your colleagues and students, and advertise for a successful IGS 2012. 

Updates of the situation and plans will be posted on the IGS website regularly. I am positive to see 

you all in Matsumoto in summer 2012. 

Ilse Storch, Chair, Grouse Group within the IUCN-SSC/WPA Galliformes SG (GSG), 

Co-Chair, IUCN-SSC/WPA Galliformes SG. 

Department of Wildlife Ecology and Management, Institute of Forest Zoology, University of Freiburg, D- 

79085 Freiburg, Germany, ilse.storch@wildlife.uni-freiburg.de 

3


CONSERVATION NEWS 

Urgent measures for conservation of the Cantabrian capercaillie 

Tetrao urogallus cantabricus in its habitat in the Cantabrian mountain 

range 

Ignacio Torres Ruiz-Huerta 

The EU LIFE+ project “Conservation of the Cantabrian capercaillie Tetrao urogallus cantabricus in its 

habitat in the Cantabrian Mountain range" began on October 1, 2010. The project main goal is to curb the 

drastic decline in this subspecies endemic to the Iberian Peninsula, whose numbers have greatly dwindled 

over the last few decades, and to offer a last hope for its recovery. The Cantabrian capercaillie has been 

classified by the IUCN as critically endangered, a category analogue to its status in the different Spanish 

Regions (Comunidades Autónomas) and in a national level (where it is considered in risk of extinction). 

The project is coordinated by the Fundación Biodiversidad, financed 50% by the European 

Commission through LIFE+ funds and the Regional governments of Asturias, Cantabria, Castilla y León 

(through Fundación Patrimonio Natural de Castilla y León) and Galicia, the National Parks Authority 

(depending of Spanish Ministry of Environment, Marine and Rural Affairs) and SEO/BirdLife are 

partners and the Fundación Iberdrola is also an important co financer. It will run until September 2014, 

with a budget of 7.3 million Euros. 

This project coordinates all the different regional governments‟ actions, where several preliminary 

actions are being undertaken. The aim is to surpass the pilot activity phase in order to implement a full, 

homogenous, cross-cutting project for the entire area where the species is found, combining in-situ 

conservation measures (i.e. habitat improvement, control of predators and competitors and non-natural 

mortality reduction) with ex-situ activities (breeding in captivity programs and restocking). Civil society 

is also involved and the inclusion of a bird conservation NGO, SEO/BirdLife, is a crucial part of this 

participation. Private landowners are also included through several tools including the land stewardship, 

contributing through innovation to the conservation of the species. Besides, a complete awareness raising 

programme to explain the species situation will be developed. 

The important degree of rural exodus in the project area (more than 600,000 hectares included in 16 

different SPA (Special Protection Areas) brings out the need to maintain the population by generating 

more employment in this area, undoubtedly one of the basic objectives of the project which will 

contribute to rural development and therefore to the conservation of the species. 

In addition, not only significant scientific research will be carried out to ensure the objective success, 

but also cooperation with both international and Spanish experts will be ensured in order to come up with 

the better proposals for the species survival. We consider the project implementation a priority in Spain 

and at a European level in order to prevent the extinction of this emblematic tetraonidae whose current 

situation is alarming. The official website of the project is www.lifeurogallo.es. 

For further information: Ignacio Torres Ruiz-Huerta. Spanish Ministry of Environment, Marine and 

Rural Affairs. 

Ignacio Torres Ruiz-Huerta, Director de Estudios y Proyectos. Fundación Biodiversidad, C/ Fortuny 7; 

28010 Madrid (Spain). itorres@fundacion-biodiversidad.es. 

4


RESEARCH REPORTS 

Effects of large-scale human land use on capercaillie Tetrao urogallus 

(L.) populations in Finland - A short summary of the PhD Thesis 

Saija Sirkiä 

Spatial requirements of capercaillie Tetrao urogallus are known to be impressive: lekking areas require a 

minimum of 300 hectares of forest (Wegge & Larsen 1987, Storch 1997, Wegge et al. 2003) and because 

of the seasonal changes in habitat use, individual home ranges may cover up to several tens of square 

kilometers (Wegge & Rolstad 2002). Thus, the species is often used as a focal species in landscape 

ecological studies. In Finland, capercaillie populations have decreased by approximately 40–85%, with 

the declines likely to have started in the 1940s (Lindén & Rajala 1981, Lindén 2002). Although the 

declines have stabilized from the 1990s onwards, it is obvious that the negative population trend was at 

least partly caused by changes in human land use. 

The aim of my thesis was to study the connections between human land use and capercaillie 

populations in Finland, using several spatial scales. First, I studied the effect of forest age structure on 

capercaillie population trends in 18 forestry board districts in Finland during 1965–1988. Second, I 

compared the abundances of capercaillie and moose Alces alces (L.) in terms of several land-use variables 

on a scale of 50 × 50 km grids and in five regions in Finland. Third, I investigated the effects of forest 

cover and fine-grain forest fragmentation on capercaillie lekking area persistence in three study locations 

in Finland, on 1000 and 3000 m spatial scales surrounding the leks. The analyses considering lekking 

areas were performed with two definitions for forest: > 60 m 3 ha –1 and > 152 m 3 ha –1 of timber volume. 

The results showed that patterns and processes at large spatial scales strongly influence capercaillie in 

Finland. Positive associations between capercaillie abundance and high forest cover and/or low human 

impact were most pronounced in southwestern and eastern Finland, where high proportion of fields and 

lakes, respectively, limit the amount of suitable habitats (Sirkiä et al. 2010a). Forest cover (> 60 m 3 ha –1 of 

timber) positively affected lekking area persistence only at the larger landscape scale (3000 m radius 

surrounding the lekking sites, covering approximately 28 km 2 , Sirkiä et al. in press). According to Wegge 

& Rolstad (2002), the annual home range size of an individual capercaillie may cover 30–80 km 2 . If the 

males stay in close vicinity of the lekking sites almost throughout the year (Wegge & Larsen 1987), it is 

reasonable that our result fits this range. 

The effects of older forest classes were hard to assess due to scarcity of older forests in several study 

areas. Young and middle-aged forest classes were common in the vicinity of areas with high capercaillie 

abundances especially in northern Finland (Sirkiä et al. 2010a). The increase in the amount of younger 

forest classes did not provide a good explanation for capercaillie population decline in 1965–1988 (Sirkiä 

et al. 2010b). In addition, there was no significant connection between mature forests (> 152 m 3 ha –1 of 

timber) and lekking area persistence in Finland (submitted manuscript). It seems that in present-day 

Finnish landscapes, area covered with old forest is either too scarce to efficiently explain the abundance 

of capercaillie and the persistence of the lekking areas, or the effect of forest age is only important when 

considering smaller spatial scales than the ones studied in this thesis. Our findings are supported by other 

recent studies considering forest age and capercaillie abundance (e.g. Miettinen et al. 2008) or lekking 

sites (e.g. Rolstad et al. 2007). It has also been suggested that habitat quality has been deteriorated in the 

older, managed forest stands, especially in northern Finland (Miettinen et al. 2009, 2010). 

In conclusion, larger spatial scales should be considered for assessing the future capercaillie 

management. According to the proposed multi-level planning the first priority should be to secure the 

large, regional-scale forest cover, and the second priority should be to maintain fine-grained, 

heterogeneous structure within the separate forest patches. A management unit covering hundreds of 

hectares, or even tens or hundreds of square kilometers, should be covered, which requires regional-level 

land-use planning and co-operation between forest owners. 

References 

Lindén, H. (Ed.), 2002. Metsäkanalintututkimuksia: Metsäkanalintukannat. Gummerus Kirjapaino Oy, 

Saarijärvi (in Finnish). 

Lindén, H., Rajala, P., 1981. Fluctuations and long-term trends in the relative densities of tetraonid 

populations in Finland, 1964–77. Finn. Game Res. 39, 13–34. 

Miettinen, J., Helle, P., Nikula, A., Niemelä, P., 2008. Large-scale landscape composition and capercaillie 

(Tetrao urogallus) density in Finland. Ann. Zool. Fenn. 45, 161–173. 

5


Miettinen, J., Helle, P., Nikula, A., Niemelä, P., 2009. Changes in landscape-scale habitat selection of 

capercaillie (Tetrao urogallus) in managed north-boreal forest. Silva Fenn. 43, 595–608. 

Miettinen, J., Helle, P., Nikula, A., Niemelä, P., 2010. Capercaillie (Tetrao urogallus) habitat 

characteristics in north-boreal Finland. Silva Fenn. 44, 235–254. 

Rolstad, J., Rolstad, E., Wegge, P., 2007. Capercaillie Tetrao urogallus lek formation in young forest. 

Wildl. Biol. 13 (suppl. 1), 59–67. 

Sirkiä, S., Helle, P., Lindén, H., Nikula, A., Norrdahl, K., Suorsa, P. & Valkeajärvi, P. Persistence of 

Capercaillie (Tetrao urogallus) lekking areas depends on forest cover and fine-grain fragmentation of 

boreal forest landscapes. – Ornis Fenn. in press. 

Sirkiä, S., Lindén, A., Helle, P., Nikula, A., Knape, J. & Lindén, H. 2010b. Are the declining trends in 

forest grouse populations due to changes in the forest age structure? A case study of Capercaillie in 

Finland. – Biol. Conserv. 143, 1540–1548. 

Sirkiä, S., Pellikka, J. & Lindén, H. 2010a. Balancing the needs of capercaillie (Tetrao urogallus) and 

moose (Alces alces) in large-scale human land use. – Eur. J. Wildl. Res. 56, 249–260. 

Storch, I., 1997. Male territoriality, female range use, and spatial organization of capercaillie Tetrao 

urogallus leks. Wildl. Biol. 3, 149–161. 

Wegge, P., Kvålsgard, T., Hjelhord, O., Sivkov, A.V., 2003. Spring spacing behaviour of capercaillie 

Tetrao urogallus males does not limit numbers at leks. Wildl. Biol. 9, 283–289. 

Wegge, P., Larsen, B.B., 1987. Spacing of adult and subadult male common capercaillie during the 

breeding season. Auk 104, 481–490. 

Wegge, P., Rolstad, J., 2002. Storfuglen og skogbruket: et sammandrag fra 20 års undersøkelser i Varald 

statskog, Hedmark, in: Seminarrapport IBN-SEVU. Norges Landbrukshøyskole, Ås, pp. 15–23 (in 

Norwegian). 

Sirkiä, S. 2010. Effects of large-scale human land use on Capercaillie (Tetrao urogallus L.) populations in 

Finland. - Ph.D. Dissertation, University of Helsinki. 

Opponent: Professor Tomas Willebrand 

Saija Sirkiä, Saija Sirkiä, Department of Biosciences, Viikinkaari 1, P.O. Box 65, FI-00014 University of 

Helsinki, Finland, saija.sirkia@gmail.com. 

Why the capercaillie population Tetrao urogallus (L.) in the West 

Carpathians decline? 

Miroslav Saniga 

From 1981-2010, population dynamics of capercaillie Tetrao urogallus L. was studied on 43 leks in the 

West Carpathians (Slovakia). Nest and chick losses were also studied. Altogether 94 nests, 124 hens with 

chicks in June and 132 in the period between 1 st August and 15 th September were checked. Results 

demonstrate a marked decrease (>50%) in numbers of cocks and hens on 12 monitored leks (28%) and a 

slight decrease (


LINDÉN, 1989, STORAAS et al. 1999). In terms of landscape ecology this large-scale change in forest 

mosaic is expected to have profound effects on spacing pattern and range use of wildlife species, 

especially those having home ranges and cruising radii within the critical area interval (ROLSTAD & 

WEGGE 1989a). Capercaillie belongs to this area-sensitive category, inhabiting old forest most of the year, 

and having seasonal ranges between 10 and 1000 hectares in size (WEGGE and LARSEN, 1987). 

In the recent few decades, populations throughout most of Western Europe have declined markedly 

(e.g. NOVÁKOVÁ & SŤASTNÝ 1982, KLAUS et al., 1986, KLAUS & BERGMANN 1994, SANIGA 1999). A 

decline in capercaillie populations has also been observed during the last 20-30 years in Fennoscandia and 

Russia (e.g. RAJALA & LINDÉN 1984, ROLSTAD & WEGGE 1989a). 

This paper reports on the findings of a 21-year capercaillie population study in the mountains of 

central Slovakia (West Carpathians). This study is aimed at: (a) monitoring population dynamics of the 

capercaillie on leks and the surroundings; (b) chick losses during the summer; (c) evaluation of the sex 

ratio in the chicks; (d) the relationship between nest and chick losses and predation factors; and (e) 

explore reasons for the persistent downward trend in numbers that was documented over the study period. 

Material and methods 

The field work was conducted in the mountains of central Slovakia 600 m a.s.l. to 1,530 m a.s.l (Veľká 

Fatra Mts., Malá Fatra Mts., Kremnické vrchy Mts., Starohorské vrchy Mts., and Nízke Tatry Mts., 

18°50‟ - 19°10‟E; 48°47‟ - 49°19‟N) from 1981-2010. The climate is moderately continental with a mean 

temperature of the warmest month (July) of 14.5°C and minus 5.5°C for the coldest (January). Yearly 

mean precipation is 1,000 - 1,400 mm, and the ground is usually covered with snow from mid-November 

to late March or April (depending on the altitude and exposure). 

In the study area mixed forest biocoenoses consisting of the spruce-beech-fir vegetation tier dominate 

(90%) (Picea abies (L.), Abies alba Mill., Fagus sylvatica L., Acer pseudoplatanus L.). Coniferous 

forests of the spruce vegetation tier constitute around 10% of the study area (Picea abies (L.) dominated, 

sprinkled with Acer pseudoplatanus L., Fagus sylvatica L., and Sorbus aucuparia L.). 

Capercaillie is difficult to count at throughout the year, but it is practicable to count the number of 

cocks displaying on leks in spring (KLAUS et al., 1986). Accuracy of the quantitative investigations 

depends on the exact timing of the census. In the initial phase of display activity (late March), cocks do 

not visit leks regularly. The period between 20 th April and 10 th May is most suitable for surveys of the 

capercaillie in Central Europe (SANIGA, 1998a). In this period, hens also visit the leks regularly. 

In 1981-2010, a total of 43 leks were monitored during the spring display season. The study was 

largely carried out by observing birds from the vicinity of the leks not to disturb the lek. Observation sites 

were usually occupied in the evening before the arrival of the males and were usually left when the 

morning display ended. Capercaillie was counted at least twice during the spring display season on the 

lek. Leks were censused especially during the period 20 th April to 10 th May (peak of lekking activity). A 

possible bias in the material is that data from some leks were not obtained during this peak period. The 

number of hens present on the leks is considered underestimated in comparison to cocks, as hens are 

much less conspicuous on the leks. Altogether 652 evening and 1088 morning observations were carried 

out on the forty-three leks during the spring display season. 

Results and discussion 

Spring density in natural forests 

During 1989-1991, spring density of capercaillie in natural forests varied between 0.5 ind/100 ha in 

dwarfed pine stands, 1.4-1.9 ind/100 ha in forests of the spruce vegetation tier, and 2.0-2.7 ind/100 ha in 

forests of the spruce-beech-fir vegetation tier. In the period 1999-2001, a dramatic decline in the spring 

density was found in the natural forests compared to 1989-1991 with 0.1 ind/100 ha in dwarfed pine 

stands, 0.7-1.1 ind/100 ha in forests of the spruce vegetation tier and 0.9-1.3 ind/100 ha in forests of the 

spruce-beech-fir vegetation tier. The density estimate for dwarfed pine stands was affected by a 

methodological bias due to the proportionately shorter length of transects. In contrast to the situation in 

Fennoscandia and Russia, long-term population studies of capercaillie based on censuses of leks during 

the display season are scarce in Central Europe (MüLLER, 1974). In Norway, WEGGE (1983) found a 

density close to 2.5 ind/100 ha. RAJALA (1974) found the capercaillie density to be 5.98 ind/100 ha in 

central Finland. According to Klaus et al. (1986), this density is the upper limit for optimal habitat. 

Population dynamics on leks 

Results demonstrate a marked decrease (>50%) in numbers of cocks and hens on 12 monitored leks 

(28%) and a slight decrease (


birds per lek was 6.3 cocks and 6.0 hens when the monitoring started in 1981. It declined to 1.5 cocks (r = 

0.83, p = 0.001, y = -0.189x + 20.588) and 1.5 hens (r = 0.67, p = 0.001, y = -0.142x + 15.823) per lek in 

2010 (Table 1). Findings in this study concerning the numbers of cocks and hens visiting the leks during 

the display season agree roughly with KOIVISTO & PIRKOLA (1961), who monitored 185 leks in Finland. 

They found 2.3-5.2 cocks and 2.8-3.9 hens per lek. 

Table 1. Presence of capercaillie on examined 43 leks during spring display season West Carpathians, 

Slovakia, 1981-2010). 

Year Checked Individuals Average per lek 

leks Males Females Males Females 

1981 4 25 24 6.3 6 

1982 4 24 27 6 4.3 

1983 7 31 27 4.4 3.9 

1984 7 31 28 4.4 4 

1985 7 23 29 3.3 2.7 

1986 6 24 17 4 2.8 

1987 8 18 19 2.3 2.4 

1988 8 27 17 3.4 2.1 

1989 13 40 28 3.1 2.2 

1990 23 74 43 3.2 1.9 

1991 23 78 52 3.4 2.3 

1992 21 68 52 3.2 2.5 

1993 23 67 62 2.9 2.7 

1994 17 53 55 3.1 3.2 

1995 25 71 61 2.8 2.4 

1996 22 50 46 2.3 2.1 

1997 22 41 46 1.9 2.1 

1998 10 21 27 2.1 2.7 

1999 10 19 25 1.9 2.5 

2000 9 16 22 1.8 2.4 

2001 10 17 26 1.7 2.6 

2002 62 118 104 1.9 1.7 

2003 72 116 128 1.6 1.8 

2004 69 124 119 1.8 1.7 

2005 79 137 110 1.8 1.4 

2006 73 124 108 1.7 1.5 

2007 66 112 93 1.7 1.4 

2008 73 110 116 1.5 1.6 

2009 74 111 108 1.5 1.4 

2010 56 84 82 1.5 1.5 

Altogether 903 1854 1701 2.1 1.9 

Surrounding habitats are expected to influence lek population size in capercaillie (LARSEN & WEGGE 

1985). Forest stands 80 years and older with suitable spatial structure covered 20-90% of the area within 

1 km of the checked lek centres. On four leks surrounded by forest with only 20-30% old growth, 1-2 

cocks displayed. On the contrary, on six leks with 80-90% old growth 5-12 cocks displayed (Table 2). 

Comparing the number of cocks on 43 leks with the proportion of old-growth forest (over 80 years old 

with suitable spatial structure) within 1 km radius of a lek, a statistically highly significant correlation 

between the amount of old-growth forest and the number of cocks attending a lek was found (Pearson 

correlation coefficient r = 0.725, p < 0.01). This supports WEGGE & ROLSTAD‟s (1986) findings that leks 

surrounded by a high proportion of old-growth forest supported more males than leks in fragmented 

areas. On 9 of 11 leks where capercaillie disappeared completely during the study period the surrounding 

8


habitat changed drastically. Presence of old-growth forest with suitable spatial structure within 1 km 

radius of a lek declined to less than 20%. 

Table 2. Relationship between the amount of old forests (over 80 years old) within a 1 km radius of the 

lek centres and the maximum number of cocks attending a lek (West Carpathians, Slovakia, 1981-2010, r 

= 0.025, p < 0.01). 

Old forest Number of cocks 

% 1 2 3 4 5 6 8 12 Sum leks 

21-30 3 1 

4 

31-40 2 2 1 

5 

41-50 2 2 1 

5 

51-60 

3 1 

4 

61-70 

3 2 

5 

71-80 

4 8 1 1 

14 

81-90 1 3 1 1 6 

Sum leks 7 5 12 11 2 4 1 1 43 

When old natural forests are fragmented by clearcuts, the landscape loses qualities which are very 

important to this tetraonid. Change of the forest landscape from old-growth forests to clearcuts and 

younger stands may increase the number of small rodents. This presumably favours higher densities of 

generalist predators (especially marten and fox), which prey on capercaillie eggs and chicks (ROLSTAD & 

WEGGE 1989b). Furthermore, the fragmentation of continuous forest habitat and its replacement with 

young stands unsuitable for capercaillie disrupts the social organization of capercaillie populations, 

particularly the formation of lek communities (KLAUS & BERGMANN 1994). 

Chick losses during summer 

Between 1 st August and 15 th September, 81 of 131 capercaillie hens had chicks (Table 3). The mean 

number of juveniles per hen was 1.9 over the whole study. The average number of chicks accompanying a 

hen significantly decreased during the study period (r = 0.77, p = 0.0003, y = -0.409x + 41.155). 

Predation pressure on chicks was high in spite of the fact that as breeding season progressed food 

offer for predators was continually increasing. Mean clutch size in capercaillie was 6.8 eggs (n = 94). In 

nests which were not destroyed or abandoned (n = 37), on average 5.7 chicks hatched. Hens had only 2.4 

chicks (n = 124) in June on the average and only 1.9 chicks (n = 131) in the period between 1st August 

and 15th September. 

Nest losses in capercaillie depend on many factors as habitat type, vegetation cover, timing of the egglaying, 

egg colour, nest localization and weather conditions (MüLLER 1984). Predation pressure on 

capercaillie chick show regional differences, but also depends on season and other factors (KLAUS et al., 

1986). Uncamouflaged nest is with high probability detected and robbed by corvid birds (raven, jay). On 

the contrary, mammalian predators use much scent, thus nest camouflage does not play a significant role. 

Sex ratio in the chicks 

Of all broods observed in the mountains of central Slovakia in the period between 1 st August and 15 th 

September, 96 were counted reliably. Chick number varied between 2 and 6. Broods of 3 were most 

frequently found, making up more than 1/3 of all broods seen. The mean brood size was 2.2. Broods were 

significantly larger in the beginning of the decline than later (Table 3). 

Only 53 broods totalling 171 chicks were reliably sexed. They consisted of 69 males and 102 females. 

Female chicks were consistently outnumbering male chicks. The deviation from 1:1 ratio was in each 

case significant at ten percent probability or smaller when tested by 2 using YATE‟s correction ( 2 = 

12.41, p< 0.001). The sex ratio was clearly related to the size of the brood. In 13 2-chick broods, 7 

consisted only of females, in 5 there was one of each sex and in one only males. In both cases the 

deviation from a 1:1 ratio was statistically significant. The difference seemed to even out as broods 

become larger, but the pattern was inconsistent. Table 4 illustrates the predominance of female chicks and 

how the sex ratio is related to the size of brood. There was a remarkable increase in the proportion of 

female chicks when brood size decreased from three to two. When comparing the sex composition 

between successively larger broods, there were no statistical difference between sizes three, four, five, 

9


and six, whereas the difference between two and three was significant at ten percent probability (t = 2,25, 

p < 0.10, 31 df, one-tailed t-test). Brood sizes one and two had significantly fewer males than did any 

other brood size. 

Table 3. Observations of capercaillie hens with chicks between 1 st August and 15 th September (West 

Carpathians, Slovakia, 1983-2010, n =132). 

Number of chicks 

Year 0 2 3 4 5 Total Average per hen 

1983 

1 1 1 3 4 

1984 

1 

1 

2 4 

1986 

1 2 1 4 4 

1987 

1 1 

1 3 3.3 

1989 

2 3 

5 3.6 

1990 

2 2 2 

6 2.7 

1991 2 3 3 

8 1.9 

1992 3 

2 2 

7 2 

1993 2 1 2 

5 1.6 

1994 

1 1 

2 3.5 

1995 3 

2 2 

7 2 

1996 3 

2 2 

7 2 

1997 4 2 2 1 

9 1.6 

1998 3 

1 2 

6 1.8 

1999 4 

1 2 

7 1.6 

2000 2 3 2 

7 1.7 

2001 3 

2 2 

7 2 

2002 1 2 

3 1.9 

2003 3 

1 

4 1.6 

2004 2 2 

4 1.6 

2005 2 2 

4 1.9 

2006 3 

1 

4 2 

2007 3 

1 1 

5 2 

2008 2 1 1 

4 1.9 

2009 3 

1 

4 1.2 

2010 3 1 4 0.8 

Total 51 20 32 25 3 131 1.9 

Table 4. Composition of capercaillie broods sampled between 1 st August and 15 th September (West 

Carpathians, Slovakia, 1983-2010, n = 53). 

Proportion 

Brood size n males females M/F % males % females 

2 13 0.54 1.46 0.37 26.9 73.1 

3 20 1.2 1.8 0.67 40 60 

4 16 1.75 2.25 0.78 43.7 56.3 

5 3 2.33 2.67 0.88 46.7 53.3 

6 1 3 3 1 50 50 

Altogether 53 1.3 1.92 0.68 40.4 59.6 

10


Predators influencing the capercaillie populations 

Predation appears to be of major importance in limiting numbers of birds, including capercaillie. Out of 

75 capercaillie clutches 49 (65%) were destroyed. Main mammalian egg predators of the capercaillie 

were found pine marten Martes martes (L.), stone marten Martes foina (Erxl.), mustelids Mustela sp. and 

red fox Vulpes vulpes (L.) (altogether 18%), wild boar Sus scrofa (L.) (6%) and brown bear Ursus arctos 

(L.) (4%). According to KLAUS (1984), predation of wild boar on capercaillie nests can locally reach 

30%. Main avian egg predators were corvid birds, particularly jay Garrulus glandarius (L.) and raven 

Corvus corax (L.) (altogether 18%). 

In the years with very cold weather during May (heavy snowfall), nests were destroyed by snow cover 

and abandoned (21%). Four clutches (8%) were found abandoned, the hens were probably predated by 

goshawk Accipiter gentilis (L.), golden eagle Aquila chrysaetos (L.) and ural owl (Strix uralensis Pall.), 

or by some of the mammalian predators - lynx Lynx lynx (L.) in 12 destroyed and abandoned nests (25%) 

the reason was unknown. 

Predation pressure on capercaillie nests decreased significantly during the incubation period (74% nest 

losses during the first half of May, n = 49; 54% in the second half of May, n = 35). A decrease in nest 

losses during the incubation period was expected because at the start of the capercaillie egg-laying no 

other bird species except birds of prey and owls are nesting. Capercaillie nests quite early in the spring 

prior to the onset of breeding of most sedentary and migratory birds. Thus the predation pressure on this 

forest-dwelling grouse is much higher in the first half of May than later when 53-59 bird species breed 

(SANIGA 1994, 1995a, 1995b). A second factor is that nests placed on the ground at the beginning of May 

may be too exposed to predators until the vegetation has adequately developed (FULLER 1995). Thus the 

predations pressure on the early breeding birds is expected to be much higher than later as the breeding 

season progresses. 

Fences, wires and disturbances 

Of 23 perished adult capercaillie, hitting fences was a common cause of death to 11 (48%). Fences used 

in mature forests where browsing by deer is preventing the growth and development of natural 

regeneration does present a considerable hazard to capercaillie, especially when a fence runs through a 

forest. Two capercaillie were found to be killed on cables of ski-lifts. 

Enthusiastic gamekeepers and birdwatchers anxious to view a lek were harmful to capercaillie display. 

Similarly forest management during the display season caused continued disturbance and had a 

deleterious effect on lek capercaillie populations. Tourism is a significant limiting factor in most 

capercaillie habitats in Central Europe (KLAUS & BERGMANN, 1994). 

Conclusion 

Several factors have contributed to the dramatic recent decline in capercaillie population in the mountains 

of the West Carpathians. Habitat deterioration has probably played a main role. The correlation between 

the amount of old forest and the number of cocks attending a lek has been significant. The presence of 

older trees has appeared to be important for capercaillie in West Carpathians, as elsewhere. Other factors 

have also contributed to the rapid decline. The recent decline in numbers has also been associated with an 

increase in rainfall and snowfall in early June. The number of rain- and snow-days in this crucial period, 

when most of the chicks hatch, has been inversely associated with capercaillie breeding success. 

Deteriorating climatic conditions for capercaillie could override any improvements in habitat quality 

(MOSS & PICOZZI 1994). Predation has appeared to be of major importance in limiting numbers of 

capercaillie populations. Fences have also been an important cause of capercaillie mortality. Continued 

disturbance caused by gamekeepers, enthusiastic birdwatchers and forest managers also has had a 

deleterious effect on capercaillie lek populations. The future of capercaillie populations in the West 

Carpathians will depend on the way in which the forest resources will be used and also on the effects of 

air pollution on forest health, ground vegetation and the abundance of insects available to chicks during 

the first weeks of their life (PORKERT 1991). Habitat improvement via forest management practices 

should be the most successful way to save the species. 

Acknowledgement 

This contribution/publication is the result of the project implementation: Centre of Excellence „Adaptive 

Forest Ecosystems“, ITMS: 26220120006, supported by the Research & Development Operational 

Programme funded by the ERDF. 

References 

FULLER, R.J. 1995. Bird life of woodland and forest. - Cambridge: Cambridge University Press. 244 p. 

11


HELLE, P. 1985. Effects of forest fragmentation on bird densities in northern boreal forests. - Ornis 

Fennica 62: 35-41. 

KLAUS, S. 1984. Predation among capercaillie in a reserve in Thuringia. - In LOVEL, T. AND HUDSON, P. 

(eds). - International Grouse Symposium 3: 334-346. 

KLAUS, S. & BERGMANN, H.H. 1994. Distribution, status and limiting factors of capercaillie (Tetrao 

urogallus) in central Europe, particularly in Germany, including an evaluation of reintroductions.- 

Gibier Faune Sauvage, Game Wildl., 11: 57-80. 

KLAUS, S., BERGMANN, H.H., ANDREEV, A.V., MÜLLER, F., PORKERT, J. & WIESNER, J. (eds) 1986. - Die 

Auerhühner. Wittenberg-Lutherstadt: Ziemsen Verlag. 276 p. 

KOIVISTO, I. & PIRKOLA, M. 1961. Behaviour and number of the capercaillies and black grouses on leks. - 

Suomen Riista 14: 53-64. 

LARSEN, B.B. & WEGGE, P. 1985. Habitat characteristics of territorial capercaillie cocks during the 

breeding season. In LOVEL, T. AND HUDSON, P. (eds). - International Grouse Symposium 3: 236-246. 

LINDÉN, H. 1984. Changes in Finnish tetraonid populations and some factors influencing mortality. - 

Finnish Game Research 39: 3-11. 

LINDÉN, H. 1989. Characteristics of tetraonid cycles in Finland.- Finnish Game Research 46: 34-42. 

MOSS, R. & PICOZZI, N. 1994. Management of Forests for Capercaillie in Scotland. - Bulletin HMSO 113: 

1-32. 

MÜLLER, F. 1974. Hahn in Ruh. - Pirsch 26: 50-52. 

MÜLLER, F. 1984. The loss of capercaillie clutches - an evaluation of a ten year study on simulated nests 

in the western Rhön mountains. In LOVEL, T. AND HUDSON, P. (eds). - International Grouse 

Symposium 3: 347-353. 

NOVÁKOVÁ, E. & ŠŤASTNÝ, K. 1982. Bestand und Bestandsentwicklung des Auerhuhns in Böhmen und 

Mähren. In KEMPF, C. (ed.). Actes Coll. Int. Grand Tetras: 35-42. 

PORKERT, J. 1991. Nebelfrost als das Aussterben von Tetraoniden förderner Faktor in den Ostsudeten. 

Acta ornithoecol. 2: 195-209. 

RAJALA, P. 1974. The structure and reproduction of Finnish populations of Capercaillie, Tetrao urogallus, 

and Black Grouse, Lyrurus tetrix, on the basis of later summer census data from 1963-66. - Finnish 

Game Research 35: 1-51. 

RAJALA, P. & LINDÉN, H. 1984. Finnish tetraonid populations in 1982-83 according to the August routecensuses. 

- Suomen Riista, 31: 92-99. 

ROLSTAD, J. 1986. Effects of logging on capercaillie Tetrao urogallus leks. II. Cutting experiments in 

south-eastern Norway. - Scandinavian Journal Forest Research 4: 99-109. 

ROLSTAD, J. & WEGGE, P. 1987. Habitat characteristics of Capercaillie Tetrao urogallus display grounds 

in southeastern Norway. - Holarctic Ecology 10: 219-229. 

ROLSTAD, J. & WEGGE, P. 1989a. Capercaillie Tetrao urogallus populations and modern forestry - a case 

for landscape ecological studies. - Finnish Game Research 46: 43-52. 

ROLSTAD, J. & WEGGE, P. 1989b. Effects of logging on capercaillie Tetrao urogallus leks. III. Extinction 

and Recolonization of Lek Populations in Relation to Clearfelling and Fragmentation of Old Forest. - 

Scandinavian Journal Forest Research 4:129-135. 

SANIGA, M. 1994. Bird community of the forests of the spruce-beech-fir vegetation tier in the Veľká and 

Malá Fatra mountains. - Biológia, Bratislava, 49: 787-794. 

SANIGA, M. 1995a. Breeding bird communities of the fir-beech to the dwarfed-pine vegetation tiers in the 

Veľká and Malá Fatra mountains. - Biológia, Bratislava, 50: 185-193. 

SANIGA, M. 1995b. Seasonal dynamics of the bird assemblages in the natural forests of the spruce 

vegetation tier. - Folia Zool., 44: 103-110. 

SANIGA, M. 1996a. Habitat characteristics of Capercaillie (Tetrao urogallus) leks in central Slovakia. - 

Biológia, Bratislava, 51: 191-199. 

SANIGA, M. 1996b. Distribution, habitat preferences and breeding biology of the Capercaillie (Tetrao 

urogallus) population in the Veľká Fatra mountains (West Carpathians). - Biológia, Bratislava, 51: 

201-211. 

SANIGA, M. 1996c. Population study of Capercaillie (Tetrao urogallus) in the Ľubochňa valley (Veľká 

Fatra mts., Slovakia). - Folia Zool., 45: 17-29. 

SANIGA, M. 1998a. Daily activity rhythm of capercaillie (Tetrao urogallus). - Folia Zool., 47: 161-172. 

SANIGA, M. 1998b. Quantitative-qualitative damage of the woody plants by the mammals and birds in the 

plantings and natural seedings. - Forest. Jour., 44: 101-109 (in Slovak, with a summary in English). 

SANIGA, M. 1999. Population dynamics of Capercaillie Tetrao urogallus on leks in Central Slovakia in the 

perid 1981-1997. - Vogelwelt 120, Suppl.: 235-240. 

STORAAS, T., KASTDALEN, L. & WEGGE, P. 1999. Detection of forest grouse by mammalian predators: A 

possible explanation for high brood losses in fragmented landscapes. - Wildlife Biology 5: 187-192. 

12


VÄISANEN, R.A., JARVINEN, O. & RAUHALA, P. 1986. How are extensive, human-caused alterations 

expressed on the scale of local bird populations in boreal forests? - Ornis Scandinavica 17: 282-292. 

WEGGE, P. & ROLSTAD, J. 1986. Size and spacing of capercaillie leks in relation to social behaviour and 

habitat. - Behavioural Ecology Sociobiology 19: 401-408. 

WEGGE, P. & STORAAS, T. 1990. Nest loss in capercaillie and black grouse in relation to the small rodent 

cycle in southeast Norway. - Oecologia 82: 527-530. 

Miroslav Saniga, Institute of Forest Ecology, Slovak Academy of Sciences, Research station, SK-976 02 

Staré Hory, Slovakia, uelsav@bb.sanet.sk. 

Hazel grouse in open landscapes 

Marc Montadert & Siegfried Klaus 

Introduction 

The hazel grouse is a forest-dwelling species rarely seen outside of forests and is reported to be a poor 

disperser (Swenson 1991). Therefore, forest fragmentation has a potentially strong negative impact on 

individual behaviour and population dynamics. At the population level, studies of hazel grouse 

occurrence in forest fragments scattered in agricultural landscapes have shown that birds do not inhabit 

forest patches isolated by an open gap of 100 m in south-central Sweden (Aberg et al. 1995) and 200-250 

m wide along the northern border of the Bohemian Forest (Sewitz & Klaus 1997; Klaus & Sewit 2000). 

Landscape structure could also impede range expansion in mountainous regions. In the south-eastern 

French Alps, the recolonisation front is blocked by open alpine meadows and rocky ridges acting as 

barriers to hazel grouse dispersal (Montadert & Léonard 2006). Telemetry studies suggested that the 

impact of forest fragmentation on hazel grouse distribution could be mainly explained by an unusual 

juvenile dispersal pattern where males moved further than females (Montadert & Leonard in press) 

In numerous regions, viable populations of hazel grouse are living in heterogeneous landscapes where 

forests were fragmented by human activity centuries ago (Fang & Sun 1997; Klaus & Sewit 2000; Sun et 

al. 2003). In such areas the local decline of traditional agriculture and forest exploitation can lead to the 

development of habitat favourable to hazel grouse in the form of young forest stages and habitat edges 

with deciduous trees and shrubs, if natural forest succession is allowed on former fields and logged sites. 

An example is the habitat that develops on meadows abandoned in the Bohemian Forest (Klaus 1995). 

In this paper, we want to use observations of hazel grouse in non-forest or edge habitats to shed light 

on three questions: 

- Is the male-biased juvenile dispersal in hazel grouse that is suggested by telemetry confirmed by 

anecdotal observations of hazel grouse outside of forests? 

- To what extent do open alpine ridges constitute definite barriers to hazel grouse dispersal? 

- To what extent are edge and corridor structures in agricultural landscapes used by hazel grouse 

and are they incorporated by settled individuals into their regular home ranges? 

Study area and Methods 

Hazel grouse outside of forests. 

Using an informal network of ornithologists and "grousers" in France, Switzerland and Germany, we 

collected observations of hazel grouse in unusual habitats by requesting the following information: sex, 

age, date, habitat type and distance from nearest forest. 

Hazel grouse using edge and corridors. 

We used personal data collected in two study sites: Auzet in the south-eastern French Alps and Sumava 

mountains in Czechia. 

Auzet. 

The Auzet study site is located in a mountainous landscape at 1,200-1,700 m a.s.l. and 76% of the area is 

forested. About half of the forest consists of old stands of fir Abies alba or beech Fagus sylvatica, the 

remainder being young coniferous stands originating from natural reforestation/succession on either clearcuts 

made 40-50 years ago or on abandoned pastures. Deciduous trees providing food were unequally 

distributed over the study area, from very abundant and diversified in mixed-stands to very few in young 

pine-spruce forests on poor soils (see Montadert & Léonard (2003) for more detailed description of study 

area). Winter feeding behaviour of hazel grouse appeared highly opportunistic in this area, depending on 

local resource availability. In rich stands, birds selected catkins/buds of hazel Corylus avellana, birch 

Betula pendula, beech, and rowan Sorbus aucuparia, whereas in poor stands, hazel grouse fed on all 

13


available deciduous trees, but leaves of bearberry Arctostaphyllos uva-ursi represented the staple food 

(unpublished data). 

At Auzet, use of edge and corridors by hazel grouse was studied by radio-tracking of one male from 

24 April to 24 December 2000, (N=64 fixes) and one female from 8 October 2004 to 22 June 2006 

(N=198 fixes). The male was captured in spring in his first-year and he settled in a forest fragmented by 

pastures, close to human settlements (Figure 3). The female was 2 months-old when captured. Only fixes 

obtained after her natal dispersal ended (9 October) were used for the analysis of edge selection. Her adult 

home range was implanted around a clearing grazed by cattle in a largely forested area (Figures 4 & 5). 

Annual and seasonal home ranges were defined by the Minimum Convex Polygon method. To check 

for habitat selection in relation to distance to edge, we divided the annual home range into three areas 

representing different distances to pasture edges: 20 m (edge habitat), 20 to 100 m and 100 m (interior 

habitat) and calculated the relative proportion of each area. These relative proportions represented the 

availability of each habitat. Radio-fixes were classified according to their location in one of the three 

areas. 

The selection of edge was studied using the selection ratio (Wi) and the difference with random use of 

habitat. We tested this difference with a Log-likelihood ratio test (LR) (Manly et al. 2002). Analyses were 

implemented using "AdeHabitat" library in R (Calenge 2006). 

Sumava Mountains (Bohemian Forest) 

Sewitz & Klaus (1997) recorded within a 10-m circle around observations of grouse the altitude (600- 

1,300 m a.s.l.), exposition, tree species composition, number of stems and cover (ground vegetation, 

shrub layer, tree layer). Four main classes of habitats were defined in the study area depending on the 

altitude: 1) valleys with alder as the dominant deciduous food tree (about 500-700 m a.s.l.), 2) lower 

slopes with birch and hazel as the dominant potential winter food (about 700-900 m a.s.l.), 3) mountain 

mixed forests with spruce, beech and fir where beech buds are the main winter food as indicated by 

qualitative fecal analysis (900-1,100 m a.s.l.), and 4) the natural mountain spruce forests where rowan is 

the principal food in winter when the snow is deep and bilberry shoots are no longer available (1,100 – 1, 

300 m a.s.l.). The main winter foods of hazel grouse in this area were primarily buds and catkins of these 

five species (Kucera, Kämpfer-Lauenstein, Lieser pers. comm. and own unpublished observations). 

Results 

Hazel grouse in open landscapes outside forests 

Between 1951 and 2009, we obtained 32 observations of hazel grouse leaving closed forests in France, 

Switzerland and Czech Republic: 21 males, 7 females and 4 unsexed specimens (Table 1). Age was 

known for 6 of the 32 birds among which 5 were classified as juveniles. Age was determined from 

prepared museum specimens or dead individuals using criteria of first and ninth primaries (Bonczar & 

Swenson 1992, Montadert & Leonard 2009). 

Of the 32 observations, 56% were individuals found dead, often killed by collision with human 

infrastructures: window (n=9) or electric wire (n=1). One was found dead in a hen house next to a farm, 

one killed by a predator. The other 44% were direct sightings or birds captured alive and released (one 

was radio-equipped and monitored during 7 months, 4 were captured in bird nets at Cou-Bretolet ringing 

station). 

Twenty-one of the hazel grouse were observed in hamlets, villages or even in cities. Six birds were 

noticed in open sub-alpine or alpine habitat. The highest recorded elevations were 2,200 and 2,460 m. 

Other observations involved birds in agricultural areas or marsh not far from forest. 

Observations were concentrated in autumn (18 out of 28), spring (n=9) and winter (n=2). Females 

were observed only in autumn (median date: 21 September, min-max: 16/08 – 14/10, n=6). Males were 

observed equally in autumn (median date: 25 September, min-max: 12/09 – 27/10, n=8) and spring 

(median date: 4 April, min-max: 20/03 – 23/05, n=9). Two were contacted in February (1 and 22). 

14


Table 1. Observations of hazel grouse outside of closed forest. 

Date Sex Age Habitat 

Distance to 

forest 

Location Commune Country Observer Comments 

sixties ? ? city ? 

1.8 km from next 

University of 

science 

Besançon 

(Doubs) 

France Cretin JY killed by collision with window 

23/11/2008 ? ? city 

forest but 20 km 

from closest 

known inhabited 

territories 

in a small 

garden 

Payerne Switzerland Henrioux P 

direct sighting in a small garden in 

residential area 

in residential Sala 

Swiss killed by collision with a window at 

20/10/2008 ? ? city 100-500 m area with Capriasca Switzerland Ornithological 3 km from closest known 

thickets (Tessin) 

Station territories 

open agriculture 

hedgerow 

20/09/1995 ? ? landscape with 100 m connected to Ancelle (05) France Borg flushed out in a hedgerow 

scattered woods 

little wood 

17/09/1990 female ? city ? martigny school Martigny Switzerland Keim C. killed by collision with window 

1921 female ? city 1 km Zool. Garten Basel Switzerland 

in Blattner 

(1995) 

found dead conserved at National 

Museum 

16/08/1994 female ? subalpine moor 50-100 m in the pass 

Bretolet 

pass(valais) Switzerland 

Swiss 

Ornithological 

Station 

ringing data. not far from open 

forest and alder moor 

10/10/2008 female ? subalpine moor 50-100 m in the pass 

Bretolet pass 

Switzerland 

(valais) 

Swiss 


Station 


forest 

14/09/2006 female juvenile village 1-1.5 km in the village Bramois Switzerland Arlettaz R 

killed by collision with window. 

preserved in St Maurice college 

14/10/2007 female ? village 55 m in the village Ardon (Jura) France 

hunting 

association of 

Jura 

Found dead without apparent 

cause in fenced garden 

25/09/2002 female ? 

village in plain 

area 

50-100 m in the village 

Bad Ragaz 

(St Gal) 

Switzerland 

Swiss 


Station 

killed by a collision with a window 

of a city building. known territories 

higher up 500-1000m hight. forest 

close to the village 

10/10/2009 male ? 

alpine rocky 

meadow 

1 km 

top of Piolit peak 

Ancelle (05) 

(2460 m alt) 

France Itier. G 

seen alive at the top of the 

mountain. See picture Fig. 1 

23/03/1984 male ? city 1.25 km in the city 

Solothurn 

(SO) 

Switzerland 

in Blattner 

(1995) 

found dead conserved at National 

Museum 

20/03/2003 male ? 

forest edge. 

agriculture area 

and village at 

615 m 

50 m? 

Steffisburg 

(Berne) 

Switzerland 

Swiss 


Station 

killed found in forest edge at 4- 

5km from known territories. not far 

from Aar valley 

08-09 1990-91 male ? 

isolated farm in 

agriculture area 

> 500 m 

Faucogney 

(70) 

France Mattieu B. 

found dead in a hen house of a 

farm surrounded by open 

agricultural landscape 

02/05/2004 male ? marsh 

1 km but small 

thicket at 200m 

belts of 

Neuchatel lake 

Coffrane Switzerland Sinz C 

direct sighting in reeds of a marsh 

with scattered willows 

04/05/2008 male 

snow in alpine 

juvenile 

meadow 

2-3 km 

Grimsel 

mountain. (2200 

m alt) 

Guttannen Switzerland Stigler M. 

found dead lying on the snow. 

See pictures Fig. 2 

27/10/2001 male 

open clearing in 

juvenile 

a forest 

10-50 m Ardon Switzerland Monney JC 

killed by collision with window of a 

isolated building in the opening 

16/09/1971 male adult subalpine moor 50-100 m in the pass 

Bretolet 

pass(valais) Switzerland 

Swiss 


Station 



12/09/2004 male ? subalpine moor 50-100 m in the pass 

Bretolet pass 

Switzerland 

(valais) 

Swiss 


Station 



captured alive in a cellar injured 

03/04/1951 male ? village ? in the village Autumn France Combe J (de la) by a phone wire during an 

overnight storm 

freshly dead after a collision with 

23/05/1973 male ? city 1.5 km In the city Kraliky Chequia Novotny. D. window, conserved in Kraliky 

Museum 

15


22/02/2006 male ? village ? Aiglun (04) France Normant F killed by collision with window 

05/10/1968 male ? village 0.2 km in the village Zofingen (AG) Switzerland 

in Blattner 

(1995) 

1987 male ? village 0.6 km in the village 

Oensingen 

(SO) 

Switzerland 

in Blattner 

(1995) 1 

16/09/1966 male ? village 1.6 km in the village 

Le Russey 

(Doubs) 

France Michelat JM 

walk into a room by open door of 

the post office in the village 

14/09/2006 male ? village 100 m in the village 

Bois d'Amont 

(39) 

France Lamy G. 

captured alive in an open hangar 

and release 

captured alive after collision with 

04/04/2009 male juvenile village 100-200 m 

in the center of 

the village 

Contamine- 

Montjoie (74) 

France Garcel. G 

a window. radio-equipped and 

tracked up to November (when 

killed by a predator) 

23/03/2009 male juvenile village 4 km in the village Dizy Switzerland Favre P. killed by collision with window 

23/03/2000 male ? village 50 m les billets Samoëns (74) France Desmet. JF killed by collision with electric wire 

17/10/2009 Male ? castle estate 50 m 

01/02/1993 male ? 

warehouse in a 

village 

100- 500 m 

in the kitchen 

garden 

close to or in the 

village 

16 

Chenecey- 

Buillon (25) 

Les 

Geneveyssur-Coffrane 

France Cannelle J. Observed alive in a walled garden 

Suisse 

Von Allmen C.- 

A. 

1 Blattner, M. 1998. Der arealschwund des Haselhuhns Bonasa bonasia in der Nordwestschweiz. - Ornis 

Beobatcher 95:11-38. 

2 Mulhauser, B. 2003. Vie de la Gélinotte des bois Bonasa bonasia dans les forêts du Haut-Jura Franco- 

Suisse. - Bulletin de la Société Neuchâteloise des Sciences Naturelles 126:15-53. 

Distance to the nearest forest averaged 0.4 km for females (n=6) and 0.8 km for males (n=18). 

Maximum distances were 1.2 km for females and 4 km for males. The most surprising observations 

involved two cases of males in alpine meadows, one found dead lying on the snow at 2,200 m, and the 

other photographed alive at 2,460 m at the rocky top of a mountain (see Figures 1 & 2). 

Figure 1. Hazel grouse cock at the top of the Piolit 

Mountain, south-eastern French Alps, 10/10/2009. 

Photo Gilles Itier. 

Figure 2. Hazel grouse cock dead on a snow pack, 

Swiss Alps, Grimsel Mountain, 4/05/2008. Photo 

Mathieu Stigler. 

Use of hedges and corridors. 

Case studies at Auzet. Space behaviour of radio-equipped hazel grouse in forest fragmented by pastures 

- Male "Diapason" 

Annual home range covered 27.5 ha, encompassing 3.8 ha of pastures with some hedgerows (Figure 3). 

Size of seasonal home ranges was 2.8 ha in spring, 9.2 in summer and 18.3 in autumn. Mating status of 

this male was not clearly established in spring but a pair was flushed at the capture site two days before 

capture. In autumn, he was observed two times with a female. 

observed alive and photographed. 

in Mulhauser 2003 2

Annual 

Autumn 

Winter 


Figure 3. Localisations and seasonal home ranges (MCP) of a radio-equipped hazel grouse male at Auzet 

(south-eastern French Alps). 

Fixes close to forest edges (< 20 m) represented 38 % of total fixes and selection analysis showed a 

positive selection for edges and rejection of forest interior in the male's home range (Table 2). The 

tendency to use forest edges changed during seasons with main use in autumn (Table 2). 

Table 2. Analysis of edge selection of two radio-equipped hazel grouse in south-eastern French Alps. LR: 

value of Log-likelihood ratio test, Wi: selection ratio standard error, P: probability value, NS : Nonsignificant. 

* Only two locations per nests were kept for analysis to prevent over weighting of nest habitat 

characteristics. Significant threshold was fixed to 0.016 using Bonferroni correction. 

Male "Diapason" Female "Jeunette" year 1 * Female "Jeunette" year 2 * 

LR P Wi P LR P Wi P LR P Wi P 

edge (20m) 

3.1


- Female "Jeunette". 

The spatial behaviour of this female was studied during 21 months. Annual home range (October to 

August) of the first year covered 94 ha, encompassing 10.1 ha of pastures (Figure 4). Autumn, winter, 

spring and summer home 

ranges covered 43.5, 19.8, 

11.9 and 17.4 ha 

respectively. This female did 

not raise a brood because of 

predation on both a first and 

second clutch. The second 

year (September to June 

when battery failed) covered 

47.7 ha encompassing 4.8 ha 

of pastures. Autumn, winter 

and spring home ranges 

covered 13.8, 29.8 and 12.1 

ha respectively (Figure 5). 

Figure 5. Seasonal home 

range (MCP) and 

localisations of a radioequipped 

hazel grouse 

female "Jeunette" during 

the second year of 

monitoring. White arrows 

and stars show location of 

nests. A: location of the 

ruin of an old farm (see 

text and Figure 6). 

18 

Figure 4. Seasonal home 

range (MCP) and 

localisations of a radioequipped 

hazel grouse 

female "Jeunette" during the 

first year of monitoring. 

White arrows and stars 

show location of nests. A: 

location of the ruin of an old 

farm (see text and Figure 6).


During the 21 months 23% of fixes were in edge (< 20 m) and selection analysis showed a clear trend 

to over use of edge and rejection of forest interior (Table 2). This female, usually accompanied by a nonequipped 

male, showed particular attraction for a small grove (0.1 ha) near a farm ruin surrounded by 

pasture (Figure 6). From autumn to spring 17 % (first year) and 18% (second year) of localisations were 

recorded in this spot (not counting fixes during incubation). This attraction could be related to abundance 

of feeding trees (notably domestic trees such as plum, cherry and pear trees mixed with hazel and 

hawthorn). 

Figure 6. View of the ruin of an old farm house in a small grove surrounded by pasture where female 

"Jeunette" was often localised, Auzet, south-eastern French Alps. 

We located four nests and only the last one of the second year succeeded, the others being destroyed 

by carnivores. Only the replacement clutch of the first year was located in forest edge (Figure 4). It was 

destroyed one or two days before hatching and the female barely escaped, leaving numerous feathers 

(stress moult) in the nest. This event likely explained the sudden shift of the summer range (Figure 4). 

In the second winter of monitoring, this female again showed a strange behaviour when she suddenly 

left her regular home range to move to the home range of a radio-equipped female neighbour (Figure 5). 

From mid-December to the end of January, these two females, accompanied by a third radio-equipped 

female coming from another neighbouring territory, seemed to form a stable group. This temporary shift 

of home range could not be associated with the search for highly attractive resources, as this place was 

not especially rich in deciduous feeding trees. This exceptional association with two females in winter, 

never noticed in other radio-equipped hazel grouse and not cited in the literature (apart from special 

behaviour of east-Siberian populations (Swenson et al. 1995)) could be linked to loss of mate but we 

could not get any proof of that. 

- Observations of hazel grouse in corridors in Sumava. 

Between 2005 and 2010 we recorded 11 cases without telemetry when hazel grouse had left closed forests 

and were found in linear vegetation structures (width 4-50 m, mean 18.5 m), surrounded by open 

grassland (Table 3). Eight birds were seen in 6 occasions, 5 data resulted from indirect evidence like a 

dust bath and/or droppings. Distances of birds from closed forests were between 20 and 250 m (mean 82 

m). The vegetation of linking structures included bands of forest and shrubs, as well as stone walls 

bordered by birch, pine Pinus sylvestris, hazel, spruce Picea abies, hawthorn Crataegus sp., juniper 

Juniperus communis and mountain ash Fraxinus excelsior. 

19


Table 3. Corridors used by hazel grouse in Sumava. 

N° Date Kind of observation 

Distance from 

forest (m) 

Type of linking structure 

Width of 

corridor (m) 

Site No. Sex 

1 14.10.2005 pair, feeding 90 old pine, birch 10-20 77 M,F 

2 29.03.2006 droppings 90 old pine, birch 10-20 77 ? 

3 07.10.2005 dust bath 70 stone wall, single spruce 2-5 96 ? 

4 02.04.2006 old burrow 20 old birch-bordered road 4-6 3 ? 

5 07.08.2005 dust bath 80 old spruce, birch, hazel 20-40 7 ? 

6 10.10.2009 flight from ground 60 old spruce spot 30 121 M 

7 10.10.2009 

flight from 

mountain ash 

70 maple, mountain ash, cherry 10 120 ? 

8 22.10.2010 flight 30 ash, birch-corridor 10 territory N M 

9 22.10.2010 flight from ground 50 juniper, stone-walls, pine 20 137 M,F 

10 21.10.2010 male feeding 90 pine, birch, hawthorn 10-20 77 M 

11 21.10.2010 dust bath 250 pine, spruce, hazel, birch 50 75 ? 

Discussion 

Dispersal and landscape fragmentation 

Our previous telemetry studies in the French Alps led us to formulate several conclusions and hypotheses 

concerning dispersal patterns in hazel grouse and the impact of landscape structure on individual 

movements (Montadert & Léonard 2006; Montadert & Leonard in press). Our main conclusions were: 

1. Natal dispersal distances in hazel grouse are usually short (< 5 km, n = 35) and mostly undertaken 

under the secure cover of continuous forest. The dispersal distance of males is longer than that of females 

beca 

dispersal (> 10 km, n = 4) which sometimes lead them to cross open habitat with temporary use of small 

wooded patches or hedgerows in an agricultural landscape. Dispersal period is exclusively autumn for 

juvenile females but both autumn and spring for juvenile males, in particular for long dispersers. 

2. Dispersal distance per se does not lead directly to a higher mortality risk, but longer use of 

unfamiliar habitat seems to increase mortality. 

3. Hazel grouse range expansion in the south-eastern French Alps is characterized by a relatively low 

expansion rate (estimated 1.5 km/year ) and the avoidance of open alpine ridges over 2,000 m high. 

New data gained in this study partially confirm the above conclusions: 

- A hazel grouse out of forest is a rare event. For instance, among 851 data of geo-localised sightings 

of hazel grouse in the French Jura between 1996 and 2010 (ONCFS/GTJ Data base), only three involved 

a bird out of forest. 

- Clearly, frequentation of unfamiliar sites is risky, in particular human settlements where numerous 

birds were killed by colliding with windows or wires. 

- Females were rarely observed in open habitats (only in autumn), whereas males were recorded both 

in fall and spring in unsuitable open habitats. The fact that juvenile male hazel grouse in the Alps move 

further during dispersal than juvenile females is also supported by our new data as the sex ratio of 

occasional sightings of birds outside forests is clearly male biased. These data suggest that males are more 

willing to cross open landscapes. The male-biased dispersal is contrary to what is currently known in 

other grouse species (see Montadert & Leonard (in press) for synthesis of natal dispersal patterns in 

grouse). 

- Furthermore, we collected two observations of males in alpine meadows, suggesting that high rocky 

ridges could be crossed during dispersal even though none of our radio-equipped birds did so. Therefore 

open alpine meadows may not be absolute barriers to hazel grouse and this could explain several 

occasional sightings reported in remote upper valleys of the Maritime Alps (still free of permanent hazel 

grouse population). These valleys can only be reached by crossing high open alpine habitat. Females 

seem to prefer denser and safer forest or shrub habitats, with open ground being more of a barrier to their 

movements. These observations strengthen the hypothesis that colonization of habitats in a fragmented 

landscape is limited more by female dispersal than by male dispersal. 

The use of corridors and edge by hazel grouse 

Tracking data obtained at Auzet showed that, in particular conditions, some adult hazel grouse can use 

edge and corridor structures as regular resources in their home ranges. Without telemetry at Sumava, we 

20


could not assess age and territorial status of hazel grouse observed in corridors, but it is likely that in 

several cases these linking structures were part of permanent adult territories. 

As to natal dispersal, even if some juvenile males can occasionally cross open gaps, the few longdispersing 

males tracked in the south-eastern French Alps passed through cover of wooded corridors and 

isolated groves. These linking structures could play a major role for the even rarer long-disperser females 

by permitting them to cross open lands that otherwise would be barriers to reaching the next habitat patch. 

It is possible that the importance of edge structures as regular resources differed between the two 

regions. At Auzet, such use appeared to be rare as only 2 birds among 89 radio-equipped cocks and hens 

settled in fragmented forest and showed particular attraction to forest edges (M. Montadert, unpublished 

data). Several other radio-equipped birds had part of their regular home range bordered by pastures, but 

we found no permanent attraction for immediate forest edge among these birds, apart from occasional 

sightings in attractive feeding trees (mainly willow Salix caprea in spring and hawthorn in winter). It is 

also noteworthy that local radio-equipped broods did not use forest-pasture edges, but rather small 

clearings imbedded within forest (M. Montadert, unpublished data). At Auzet, feeding trees was 

randomly scattered at variable density in "closed" forests with a semi-open and coarse-grained structure, 

which permitted growth of light-demanding trees and a well-developed herbaceous field layer. A forest 

with gap structure like that found at Auzet is really different from the closed forests of Sumava. In this 

latter study area where intensive forestry has resulted in spruce monocultures, a more intensive use of 

forest edges by hazel grouse is suspected because feeding trees and a well-developed field layer are often 

concentrated at stand borders. The comparison of the two different study areas shows that the hazel 

grouse is above all a forest bird preferring young succession and/ or multi-layered old stands with a gap 

structure and that in sub-optimal habitat conditions it can compensate for food resources that are rare and 

clumped by integrating richer edges into its home range. Yet, such behavioural plasticity could affect 

individual fitness if vulnerability to predators is higher at forest edge. 

Acknowledgements 

Many thanks to all contributors of hazel grouse data out of forest: Bernard Frochot, Dominique Michelat, 

Bruno Matthieu, Jean-François Desmet, Jean-François Normand, Geoffrey Garcel and special mention for 

Swiss team and Ornithological Swiss Station: Martin Blattner, Bertrand Posse, Pierre Henrioux, Raphael 

Arlettaz, Alain Barbalat, Anatole Gerber. We also need to thanks Patrick Léonard, Patricia Michel, Roger 

Izoard and A. Sewit for indispensable field assistance at Auzet and Sumava. Finally, we are grateful to 

Larry Ellison who kindly reviewed the manuscript. 

References 

Aberg, J., Jansson, G., Swenson, J.E. & Angelstam, P. 1995. The effect of matrix on the occurrence of 

hazel grouse (Bonasa bonasia) in isolated habitat fragments. - Oecologia 103: 265-269. 

Bonczar, Z. & Swenson, J.E. 1992. Geographical variation in spotting patterns on Hazel Grouse Bonasa 

bonasia primary feathers: consequences for age determination. - Ornis Fennica 69: 193-197. 

Calenge, C. 2006. The package "AdeHabitat" for the R software: A tool for the analysis of space and 

habitat use by animals. - Ecological Modelling 197: 516-519. 

Fang, Y. & Sun, Y.-H. 1997. Brood movement and natal dispersal of hazel grouse Bonasa bonasia at 

Changbai Mountain, Jilin Province, China - Wildlife Biology 3: 261-264. 

Klaus, S. 1995. Hazel Grouse in the Bohemian Forest - results of a 20-year study. Pp. 27-33 in D. Jenkins 

(editor). Proceedings of the International Grouse Symposium 6, World Pheasant Association, 

Reading, Great Britain. 

Klaus, S. & Sewit, A. 2000 Ecology and conservation of hazel grouse Bonasa bonasia in the Bohemian 

forest (Sumava, Czech Republic). Pp. 138-146 in P. Malkova (editor). Proceedings of the 

International Conference in Ceské Budejovice. Korsach, Vyskocil, V., Ceské Budejovice, Czech 

Republic. 

Manly, B.F.J., McDonald, L.L., Thomas, D.L., McDonald, T.L. & Erickson, W.P. 2002. Resource 

Selection by Animals: Statistical Design and Analysis for Field Studies. - Kluwer Academic 

Publishers, Dordrecht, The Netherlands. 

Montadert, M. & Leonard, P. 2009. Age determination of hazel grouse in the south-western limit of its 

European range. - Grouse News 37: 7-14. 

Montadert, M. & Leonard, P. in press. Natal dispersal patterns in Hazel grouse: consequences for 

population dynamics in heterogeneous landscapes. - Studies in Avian Biology. 

Montadert, M. & Léonard, P. 2003. Survival in an expanding hazel grouse Bonasa bonasia population in 

the southeastern French Alps. - Wildlife Biology 9: 357-364. 

Montadert, M. & Léonard, P. 2006. Post-juvenile dispersal of Hazel grouse Bonasa bonasia in an 

expanding population of the southeasten French Alps. - Ibis 148: 1-13. 

21


Sewitz, A. & Klaus, S. 1997. Besiedlung isolierter waldinseln im vorland des böhmerwaldes durch das 

haselhuhn (Bonasa bonasia). - Beiträge zur Jagd- und Wildforschung, Bd. 22: 263-276. 

Sun, Y.-H., Piao, Z.-J. & Swenson, J.E. 2003. Occurrence of hazel grouse Bonasa bonasia in a heavily 

human-impacted landscape near the Changbai Mountains, northeastern China. - Wildlife Biology 9: 

371-375. 

Swenson, J.E. 1991. Is the Hazel Grouse a poor disperser? Pp. 347-352 in S. Csanyi & J. Ernhaft 

(editors). Proceedings of the Congress of the International Union of Game Biologists. 20, Gödöllö, 

Hungary. 

Swenson, J.E., Andreev, A.V. & Drovetski, S.V. 1995. Factors shaping winter social organization in 

Hazel Grouse Bonasa bonasia: a comparative study in the eastern and western Palearctic. - Journal of 

Avian Biology 26: 4-12. 

Marc Montadert, ONCFS, les Granges Michel, 25300 Les Verrières de Joux, France. 

Marc.montadert@wanadoo.fr. 

Siegfried Klaus, Max Planck Institute for Biogeochemistry, Hans-Knöll-Str. 10, D-07745 Jena, 

siegi.klaus@gmx.de. 

Investigating the importance of thermal refugia for white-tailed 

ptarmigan at the southern extent of their range. 

Don Wolfe and Lena Larsson. 

Due to the aridity, extreme temperatures, and sparse vegetation, high alpine environments can be 

exceptionally harsh, and are inhabited by only a few vertebrate species that have various adaptations that 

allow them to persist. For some species, migration to lower elevations or latitude during winters permit 

them to utilize the high alpine in summers, while other species that remain alpine hibernate for upwards 

of 5-6 months, or cache food in burrows. Most avifauna employ the first of those options mentioned, 

however a few species, such as rock ptarmigan Lagopus muta and white-tailed ptarmigan Lagopus 

leucura can survive the harsh winters by consuming little more than twigs or buds, with occasional downslope 

forays for other vegetation, and by making use of snow burrows for shelter from the cold (in 

addition to their various physiological and morphological adaptations). While we all recognize and 

perhaps marvel at how well ptarmigan are adapted for extreme cold, another question is how they adapt to 

the high solar radiation in the summer? (e.g. occasionally in excess of 50° C, Potapov 2004, Acta 

Zoologica Sinica 50:970-977). 

Like some populations of rock ptarmigan in Europe and Asia, the Rocky Mountain populations of 

white-tailed ptarmigan have adapted to live in high alpine habitats at a much lower latitude than what one 

might expect. The southernmost extent of the white-tailed ptarmigan distribution is in north-central New 

Mexico, at a latitude of just below 36 degrees North. However, the sustainability of this population may 

be precarious, due to upslope advancement of trees, vegetational shifts due to climate change, and to 

general warming that adversely affect the species‟ ability to thermoregulate. Over the past 5 years, we 

have surveyed peaks and high alpine ridges of the Sangre de Cristo Mountains, New Mexico, to 

determine the present occupied range of ptarmigan within the state. Preliminary results indicate that 

ptarmigan in New Mexico are found predominately above 3750 meters (over 99% of sightings or sign), 

and are always found within or in close proximity to large boulder fields. We theorize that the crevices 

within these boulder fields not only provide protection from the ever present aerial predators, but also 

provide thermal refugia for ptarmigan and other alpine obligates. In 2010, we attempted to determine the 

value of such thermal refugia by the use of temperature/relative humidity data loggers. We deployed 12 

pairs of data loggers, each consisting of a logger in a likely refugium location, and a second nearby on the 

alpine surface. Six sets of loggers were deployed at the Wheeler Peak Wilderness Area at elevations 

ranging from 3720 to 3906 meters a.s.l., and six sets were deployed at the Pecos Wilderness Area at 

elevations ranging from 3768 to 3923 meters a.s.l. Generally, the refugia loggers recorded cooler 

temperatures during afternoons and warmer temperatures during nights and early mornings. (Figures 1, 

2). For all sets, the mean afternoon high temperatures for the refugia loggers were lower than for the 

surface loggers; although some sets showed a much greater difference than others (e.g., see Wheeler Set 1 

and Wheeler Set 3, Figure 3). 

22


Figure 1. Average daily temperatures for the entire month of August 2010 at Wheeler Peak Wilderness 

Area. 

We plan to collect additional data logger climate data in the summer of 2011 (perhaps even longer), 

and will attempt to quantify the amount of suitable habitat at various elevations and other factors (such as 

size and location of willow thickets). We also plan to quantify ptarmigan abundance as related to the 

available thermal refugia. Thus far, the temperature differences seen between refugia and alpine surface 

seem to support our hypothesis on the association of boulder fields and ptarmigan occupancy. 

Figure 2. Average daily temperatures for the entire month of August 2010 at Pecos Wilderness Area. 

23


Figure 3. Mean high temperatures for all data loggers from 15 July through 31 August 2010. 

Don Wolfe and Lena Larsson, G. M. Sutton Avian Research Center, University of Oklahoma, Bartlesville, 

OK, USA. dwolfe@ou.edu. 

Is GPS and satellite telemetry an option for grouse research? 

Since the 1960s, radio telemetry has become the standard method to study activity patterns, movements 

and habitat use of many wildlife, including grouse. Very High Frequency (VHF) radio transmitters 

located with hand-held receivers have been widely used in grouse studies; yet, it remained labour 

intensive. The Argos satellite system became available for wildlife use in the 1980s, and allowed 

convenient automatic tracking over long distances. However although technical improvements have 

reduced the size and weight of Argos Platform Transmitter Terminals (PTTs) significantly, major 

shortcomings remained in large location errors (150-1,500m) and varying degrees of successful 

transmitter uplinks depending on topography, latitude and power level (Kaczensky et al. 2010). 

In the mid-1990s, Global Positioning System (GPS) collars became available for wildlife tracking. By 

now, GPS telemetry systems promise precise locations on a pre-defined schedule, remote data access, and 

easily calculable and predictable costs. Various reports have addressed whether the new equipment can 

live up to these high expectations (Kaczensky et al. 2010 and references therein). So far, published 

applications of Argos and GPS telemetry in grouse research are rare. We are aware of a single study on 

capercaillie in Norway (Wegge et al. 2007). The most critical questions when considering using satellitebased 

telemetry systems for grouse are these: 

24


How to fix tags on grouse, and is the backpack-method acceptable? How frequently do I have to 

expect technical problems, and particularly transmitter failures? How about precision and reliability of 

locations, particularly in forested and mountainous areas? Are there any unexpected and unforeseeable 

costs involved? 

Here, we report experience from published work on capercaillie in the boreal forest of Norway 

(Wegge et al. 2007) and from an ongoing study on rock ptarmigan in the tundra of Svalbard. Please do 

report further experience to share with colleagues – we will follow up on this topic in later Issues of GN. 

References 

Kaczensky, P., Ito, T.Y. & Walzer, C.2010. Satellite telemetry of large mammals in Mongolia: what 

expectations should we have for collar function? - Wildlife Biology in Practice 2010; 2(6); 108-126. 

Wegge, P., Finne, M.H. & Rolstad, J. 2007. GPS satellite telemetry provides new insight into capercaillie 

Tetrao urogallus brood movements. - Wildlife Biology 13 (Suppl. 1): 87-94. 

Ilse Storch, Department of Wildlife Ecology and Management, Institute of Forest Zoology, University of 

Freiburg, D-79085 Freiburg, Germany, ilse.storch@wildlife.uni-freiburg.de 

GPS satellite telemetry used on capercaillie Tetrao urogallus 

The movement patterns of young capercaillie broods was studied using GPS satellite telemetry (Wegge et 

al. 2007). The aim was to test if this new technology could be applied to gain more detailed insight into 

behaviour and habitat selection at a small spatial scale, and if so to compare the broods‟ relative use of 

planted and older, naturally regenerated forests. Hens of four broods with chicks 2-7 days old were 

captured and fitted with backpacks of 90g containing GPS units and VHF transmitters. The GPS units 

were programmed to record positions every 15 minutes, the shortest interval possible. The transmitter had 

a storage capacity of 450 positions and movements could then be monitored for ca 4.5 days (Wegge et al. 

2007). 

In the study area at Varaldskogen with moderate topography, the GPS technology performed quite 

well (Wegge et al. 2007). A total of 1,277 positions were obtained (84% of potential maximum), of which 

77% were within 20 m of the true position of the brood. It was surprising that 84% of the potential 

number of recordings was recorded since brood hens move on the ground in dense vegetation and under a 

canopy of trees. The transmitter of 5-6% of the birds‟ weight did not seem to affect the birds negatively 

(Wegge et al. 2007). The GPS locations with


Experiences of satellite tags on rock ptarmigans 

Eva Fuglei & Åshild Ønvik Pedersen 

Here we report our experiences by use of satellite telemetry on a subspecies of the rock ptarmigan 

Lagopus muta, the Svalbard rock ptarmigan Lagopus muta hyperborea from an ongoing pilot study 

(http://svalbardrype.npolar.no/en/). 

Among many unknown aspects of the Svalbard rock ptarmigan‟s biology one is whether the birds 

migrate seasonally within the Svalbard archipelago (Pedersen et al. 2005). No knowledge exists about 

their wintering areas and possible migration routes during the Arctic winter (October through March). 

Claims have been made that this species could perform long-range migration, a behavior that can allow 

them to track seasonal shift in suitable feeding areas (Pedersen et al. 2005; Gudmundsson 1972). 

The Svalbard rock ptarmigan is the 

only resident terrestrial bird in the high 

Arctic Svalbard archipelago, Norway 

(74-81°N, 10-30°E). Svalbard, approx. 

62,700 km 2 , is a remote area, with 

almost no people living and no road 

connections between the few 

communities, so the ability for 

retaining tagged ptarmigan is limited. 

Therefore, the most effective method 

for gathering reliable data about bird 

migrations and overwintering areas is 

satellite telemetry. Since we were able 

to capture birds in May only, the tags 

needed to last for more than 12 

months. We selected Argoscompatible 

bird tracking PTTs 

Figure 1. Svalbard rock ptarmigan mounted with a 20 g 

satellite tag. Photo E. Fuglei. 

26 

(Platform terminal transmitters) 

designed and commercialized by 

NorthStar Sciences and Technology 

(http://www.northstarst.com/). We 

used 20 gram battery powered PTTs with a battery life time expectancy of up to 500 hours. For the tags to 

work for more than 12 months they were programmed with a duty cycle (i.e. pre-defined transmitting 

period) of 5 h every 5 day. It is important to note that the Svalbard rock ptarmigan are heavier than rock- 

and willow ptarmigan Lagopus lagopus living for instance in mainland Norway. Rock ptarmigan in 

Svalbard show large seasonal variations in 

body weight due to heavy fat deposition in 

autumn, and their bodyweight can vary 

from 500-550 g in summer to 900-1200 g 

in winter (Steen and Unander 1985). The 

weight of the tag should not exceed more 

than about 4 % of the birds body weight. 

We also based our selection of satellite tags 

on the experiences conducted on ivory gulls 

(Gilg et al. 2010). The transmitters were 

attached to the birds like a backpack that 

consisted of a plastic covered neck ring (4- 

5 cm in diameter) with a flat “tail” to fasten 

the PTT on (Figure 1, Figure 2). 

We captured and mounted 8 ptarmigans 

with satellite tags in May 2009 (5 hens and 

3 males) and 10 in May 2010 (4 hens and 6 

males). Since the Svalbard rock ptarmigan 

is extremely tame and show very little 

marked fear behavior we were able to get 

Figure 2. Svalbard rock ptarmigan flying with a satellite 

tag mounted on its back. Photo Å. Ø. Pedersen. 

close and capture them by using to different methods. (1) With a hand held Supertalon net gun 

(http://lawenforcementmall.com/ supertalon.html) where a net is shot from a distance of 7-10 m over the


Figure 3. One of the methods used for capturing the 

Svalbard rock ptarmigan. Guiding of ptarmigans against 

the mist net mounted on two bamboo poles. Photo K. 

Lone. 

27 

birds or (2) by using a mist net mounted 

on two long bamboo poles held by two 

people, while two other persons guided 

the ptarmigans against the net that was 

put over them (Figure 3). 

All of the 8 satellite tags deployed in 

May 2009 functioned satisfactorily (of a 

total of 4586 positions 84 % where of 

good quality, bearings that was good 

enough for us to estimate a location). In 

detail, after losing one bird during the 

hunt in September 2009, we lost contact 

with the first tag in March 2010, the 

second in late April, the third and fourth 

in mid-May, and the last three still 

worked by July 2010. Our experience 

with the 10 tags deployed in May 2010 is 

somewhat different. Here we also lost 

one bird during the hunt in September 

2010, but we lost contact with the first 6 

tags already during summer 2010, the 

next in October 2010, while the last two tags are still working by February 2011. We are currently 

discussing with NorthStar Sciences and Technology the reason why we lost contact with almost all the 

tags we mounted in 2010. So far we have not been able to solve the problem. In order to get more 

experience with the satellite tags we are planning to put out up to 5 more tags in May 2011. 

Thanks to support from the Governor of Svalbard, Svalbard Environmental Protection Fund, 

Nansenfondet, Sparebanken Nord-Norges gavefond and Norwegian Polar Institute 

References. 

Gilg, O., Strøm, H., Aebischer, A., Gavrilo, M. V., Volkov, A. E., Miljeteig, C. & Sabard, B. 2010. Postbreeding 

movements of notheast Atlantic ivory gull Pagophila eburnea populations. J. Avian Biol, 

41: 532-542. 

Gudmundsson, F. 1972. Grit as an indicator of the overseas origin of certain birds occurring in Iceland. 

Ibis 114: 582. 

Pedersen, Å. Ø., Overrein, Ø., Unander, S. & Fuglei, E. (2005) Svalbard rock ptarmigan (Lagopus mutus 

hyperboreus): a status report. Rapportserie no. 125, Norwegian Polar Institute. pp. 23. 

Steen, J. B. & Unander, S. (1985) Breeding biology of the Svalbard rock ptarmigan (Lagopus mutus 

hyperboreus). Ornis Scand. 16: 191-197. 

Eva Fuglei and Åshild Ønvik Pedersen, Norwegian Polar Institute, FRAM Centre, NO-9296 Tromsø, 

Norway, Email: eva.fuglei@npolar.no


BOOKS REVIEWS 

Tetraonidae and Phasianidae of the USSR – Ecology and Morphology 

[Maria A. Kuz’mina. Teterevinye I Fazanovye SSSR: Ekologo-Morfologicheskaya Kharakteristika] 

Don Wolfe 

Most book reviews are of books that have recently been published or updated. So, why am I writing a 

review of a book that was originally published over 30 years ago, and then translated into English nearly 

20 years ago? Mainly because, I, as I assume with many westerners, have only vaguely been aware of the 

tremendous amount of grouse research that has been conducted and is still being conducted in the former 

Soviet Union. I feel that language barriers and politics have effectively, even if unintentionally, 

suppressed this vast accumulation of knowledge. Additionally, this compilation is rarely cited, and the 

actual book can be difficult to find. I first became aware of it through a citation in an article on ptarmigan 

morphology, and was able to view a copy of the book through interlibrary loan. My initial reading left me 

fascinated and greatly desiring my own copy, so after nearly a year of searching, I finally found and was 

able to purchase a copy. I would like to make other grouse researchers aware of the book, and the 

valuable information contained within. 

Maria Kuz‟mina apparently was a very detail oriented person, and this book not only includes a 

compilation of her decades of research on galliforms, but also many summaries of many other researchers 

within the former Soviet Union, other areas of Europe and Asia, and even North America. She goes into 

meticulous detail on differences in anatomy and morphology between families, between species, and even 

within species inhabiting different habitats or regions. Then, she draws conclusions on how these 

differences are of importance to differing migration patterns, mobility patterns, foraging behaviors, and 

adaptations to climates. She also is equally meticulous in summarizing diets, often comparing diets for a 

single species from several locations, upwards of thousands of kilometers apart. 

The organization of this book may be somewhat different than several other sources. Whereas in most 

books, a chapter or section is devoted to a single species or family, usually following an introductory 

chapter, Kuz‟mina devotes entire chapters to one particular aspect of galliform ecology, and discusses the 

similarities and differences of that aspect between a large number of species of grouse, partridges, quail, 

and pheasants. While this format is valuable and may be of great interest, it can be a bit difficult and 

tedious if the reader is attempting to glean information on only a single species. The various chapters are 

as follows: 

1. SOME ASPECTS OF THE ORIGIN AND TAXONOMY OF GALLIFORM BIRDS 

2. REVIEW OF GALLIFORM BIRDS OF THE SOVIET UNION 

3. FEEDING 

4. ECOMORPHOLOGICAL ADAPTATIONS TO CLIMATIC CONDITIONS 

5. ECOMORPHOLOGICAL CHARACTERISTICS OF LOCOMOTOR ORGANS 

6. ECOLOGICAL GROUPS OF GALLIFORM BIRDS 

7. CONCLUSION 

This book is amply illustrated with line drawings and black and white photographs that correspond to 

the differences in anatomy and morphology between species. There are 16 tables, distributed throughout 

the text, that list measurements and ranges of anatomical and morphological features. It also includes 

range maps of most species, but some of the maps, especially those of species with very limited 

distribution, are difficult to interrupt unless the reader is already intimately familiar with that particular 

region. Approximately 650 citations are included in the bibliography, some in Russian, some in German, 

and some in English. Most, but not all, of the Russian and German citations also include an English 

translation of the title. However, one fairly important part of the bibliography is lacking, that being the 

page numbers of the journal articles. Appendences are included that list all the animal and plant species 

mentioned in the text, with the English, Russian, and scientific names. It also appears that Mr. Siegel- 

Causey did a superb job in translating the text. Throughout the book, there are numbers in the left-hand 

column that indicate the page number of the original text in Russian. 

Overall, I believe that this book should be of great interest and value to all grouse researchers, not 

only those working in the various states that once were a part of the U.S.S.R. An updated, revised edition, 

including research conducted since the 1970s would especially valuable. Even if that were not to happen 

in the near future, making this book more readily available, either through reprinting or in electronic 

28


format would still allow greater access to this amazing resource. Perhaps this endeavor can be attempted 

through the IUCN Galliformes Specialist Group. 

Kuz‟mina, M.A. 1977. Tetraonidae and Phasianidae of the USSR – Ecology and Morphology [Teterevinye 

I Fazanovye SSSR: Ekologo-Morfologicheskaya Kharakteristika] Nauka Publishers, Alma-Ata. 

Translated into English by Douglas Siegel-Causey, 1992; Gulab Primiani, Oxonian Pvt. Ltd, New 

Delhi. ISBN 81-7087-064-X. 

Don Wolfe, G. M. Sutton Avian Research Center, University of Oklahoma, P.O. Box 2007, Bartlesville, 

OK 74005,USA dwolfe@ou.edu 

29


CONFERENCES 

The 12 th International Grouse Symposium postponed 1 year 

Hiroshi Nakamura 

IGS News No. 5 (4th April) 

Dear Grouse Scientists. Due to the unprecedented disasters and continued uncertainty about the nuclear 

situation in Japan, The Local Organizing Committee of IGS2011 regretfully announces that the IGS2011 

must be canceled this year. However, The Local Committee proposes the following: Postpone the IGS 

for a year. 

As we announced in IGS News No.4, we judged that it was possible to open the symposium in this 

July in Matsumoto as it was planned. We announced our decision because neither Tokyo nor Matsumoto 

had damage from the major earthquake and tsunami. We had hoped for the nuclear situation to be 

resolved in a short time. Unfortunately, the status of the damaged nuclear plant remains unforeseeable. 

The reports of the IAEA http://www.iaea.org/newscenter/news/tsunamiupdate01.html and other sources 

indicate that this is a situation of great uncertainty and a significant risk of a sudden deterioration leading 

to serious large-scale radiation. This uncertainty will continue for weeks and, perhaps, months. 

Thus, our Local Committee and the IGS Program Committee feel that we cannot carry forward the 

responsibility for the symposium and its participants because of the current situation. In addition, Japan is 

facing another new problem this summer, which is the shortage of electric power. The loss of the nuclear 

power plant has resulted in a regulated periodic reduction in power in Tokyo and its surrounding areas. 

These reductions have succeeded and the electrical problem may disappear in a short time. However, late 

June is the peak time of electric demand. We are afraid that we may need again the periodic reduction in 

power, which causes traffic problems. 

For these reasons, our Local Committee decided to postpone the IGS for a year. We are sorry if this 

has caused anyone any inconvenience, but we feel under the circumstances that we have no choice in our 

decision. The Committee of the Grouse Group within the IUCN Galliformes Specialist Group is 

supportive of our decision. We all hope that the nuclear situation will be under control by the end of this 

summer. 

All our symposium plans may be kept and adjustments can be made as needed. Current registrations 

will be cancelled and a new registration process will be started in the fall of 2011 as we did last year. 

Those who paid registration fees for 2011 will have a refund returned as soon as possible. We will also 

try to find a way to help those who should already have booked their flights. If you need our assistance 

with cancelling your flights, please contact us. My secretary, Yukiiri, and I will contact the people 

needing cancelation help. Our hope is that everyone who has already purchased a ticket can have a 

refund. We know some airlines are allowing refunds voluntarily, but if you encounter problems, we will 

try to help. 

Our Local Committee will renew our symposium plans more attractively and, hopefully, allow more 

participants to attend more easily in 2012. The website will be continued and information updated 

regularly. We would like to continue announcements using the IGS News forum until the IGS2012. I 

have received a lot of get-well messages by e-mail. We really appreciate your kindness. We believe that 

we Japanese can rise again from this unprecedented disaster. 

The reason that we accepted IGS in Japan is because many grouse scientists were interested in our 

presentation about the Japanese Rock Ptarmigan at the IGS2005 held in Luchon, France. The ptarmigan is 

extremely unique as I pointed out in my research reports in the Grouse News 40 (Nakamura 2010). We 

believe firmly that the IGS2012 held in Japan will contribute not only to the conservation of the 

ptarmigan but also to mark a new phase in IGS. Because we have a long history of its conservation and 

researches on the ptarmigan and unique ecological, cultural, and conservation status, we expect that many 

grouse scientists will want to participate in the IGS2012 to observe our work, the grouse, and the habitat 

of these isolated populations. 

We are looking forward to see you next year in Matsumoto, Japan. 

Hiroshi Nakamura, Organizer of the Local Committee of IGS2011, Faculty of Education, Shinshu 

University, Nagano, 380-8544, Japan, E-mail (DESK-PC): hnakamu@shinshu-u.ac.jp, E-mail 

(Secretary-PC): seitajm@shinshu-u.ac.jp, URL: http://cert.shinshu-u.ac.jp/eco_lab/. 

30


Reminder – 29 th Prairie Grouse Technical Council (PGTC) Meetings 

October 4 - 6, 2011 

The 29 th Prairie Grouse Technical Council (PGTC) Meetings October 4 - 6, 2011 hosted by Kansas Dept. 

of Wildlife and Parks, Fort Hays State University, Hays, Kansas. 

A conference announcement, call for abstracts, call for Hamerstrom Award nominations and other 

info will be sent out through the PGTC listserv very soon. If you are not on that listserv and desire 

information please contact David Dahlgren at dave.dahlgren@ksoutdoors.com. Any other questions can 

be directed to Dave as well. We look forward to an excellent conference. There will be a field trip to 

Lesser Prairie Chicken range which is sympatric with Greater Prairie Chicken range in western Kansas. 

Dave Dahlgren, Kansas Department of Wildlife and Parks, 1426 Hwy 183 Alt., PO Box 338, Hays, KS 

67601-0338, USA, dave.dahlgren@ksoutdoors.com. 

6 th European Conference Black Grouse Endangered Species 

First announcement 

Invitation 

The first International Black Grouse Conference was organized in 

Belgium, in 2000. Since then the conferences have been organized 

in different countries to gather black grouse specialists from across 

Europe. The 6 th meeting will be held in Sweden. 

We are pleased to invite you to the 6 th European Conference 

Black Grouse Endangered Species. 

The conference will be held in Gysinge, Sweden, September 2012. 

More information and possibilities to register on-line will added 

later. We hope that the conference will be a great opportunity to 

discuss the present situation of the black grouse in Europe and 

further initiatives concerning the protection of this species and its 

habitats. 

Language of the conference 

The official language will be English. 

Contact person: Jacob Höglund (jacob.hoglund@ebc.uu.se) 

31 

Photo: Hugh Jansman


RECENT GROUSE LITERATURE 

For a complete bibliography on grouse, go to: http://www.suttoncenter.org/pages/publications (please 

note that the link in previous editions may not be current). 

Adam, A., L. M. I. Webster, W. Mullen, L. F. Keller, P. C. D. Johnson. 2011. Quantifying fenbendazole 

and its metabolites in self-medicating wild Red Grouse Lagopus lagopus scoticus using an HPLC-MS- 

MS approach. Veterinary Parasitology XXX:XXX-XXX \(online early). 

Alda, F., P. Sastre, P. J. De La Cruz-Cardiel, and I. Doadrio. 2011. Population genetics of the 

endangered Cantabrian Capercaillie in northern Spain. Animal Conservation XXX: XXX - 

XXX(onlineearly). 

Augustine, J. K., and B. K. Sandercock. 2011. Demography of female Greater Prairie-Chickens in 

unfragmented grasslands in Kansas. Avian Conservation and Ecology 6(1):2. [online] URL: 

http://www.ace-eco.org/vol6/iss1/art2/ 

Barnagaud, J.-Y., P. A. Crochet, Y. Magnani, A. B. Laurent, E. Menoni, C. Novoa, and O. Giminez. 

2011. Short-term response to the North Atlantic Oscillation but no long-term effects of climate 

change on the reproductive success of an alpine bird. Journal of Ornithology XXX: XXX-XXX 

(online early). (Black Grouse). 

Barry, P. D., and D. A. Tallmon. 2010. Genetic differentiation of a subspecies of Spruce Grouse 

(Falcipennis canadensis) in an endemism hotspot. Auk 127: 617-625. 

Behney, A. C., C. W. Boal, H. A. Whitlaw, and D. R. Lucia. 2010. Prey use by Swainson‟s Hawks in the 

Lesser Prairie-Chicken range of the Southern High Plains of Texas. Journal of Raptor Research 44: 

317-322. 

Bendell, L. I. 2011. Trace metal depositional patterns from an open pit mining activity as revealed by 

archived avian gizzard contents. Science of the Total Environment 409:1193-1197. (Blue Grouse). 

Bijlsma, R. G., and E. Jansen. 2010. Het Korhoen, de Havik en Staatsbosbeheer. [The Black Grouse, the 

Goshawk and the state forestry service.] Takkeling 18:108-131. (in Dutch with English abstract). 

Blanco-Fontao, B., M. Quevedo, and J. R. Obeso. 2011. Abandonment of traditional uses in mountain 

areas: typological thinking versus hard data in the Cantabrian Mountains (NW Spain). Biodiversity 

and Conservation XXX:XXX-XXX (online early). (Capercaillie). 

Bochkov, A. V., and K. Skirnisson. 2011. Description of the life stages of quill mite Mironovialagopus 

sp. nov. (Acari: Syringophilidae) parasitizing the Rock Ptarmigan Lagopus muta (Phasianidae) from 

Iceland. Parasitology Research 108:715-722. 

Bollmann, K., R. F. Graf, and W. Suter. 2011. Quantitative predictions for patch occupancy of 

Capercaillie in fragmented habitats. Ecography 34:276-286. 

Broseth, H., and H. Chr. Pedersen. 2010. Disturbance effects of hunting activity in a Willow Ptarmigan 

Lagopus lagopus population. Wildlife Biology 16: 241-248. 

Bush, K. L., C. K. Dyte, B. J. Moynahan, C. L. Aldridge, H. S. Sauls, A. M. Battazzo, B. L. Walker, K. E. 

Doherty, J. Tack, J. Carlson, D. Eslinger, J. Nicholson, M. S. Boyce, D. E. Naugle, C. A. 

Paszkowski,and D. W. Coltman. 2011. Population structure and genetic diversity of Greater Sage- 

Grouse (Centrocercus urophasianus) in fragmented landscapes at the northern edge of their range. 

Conservation Genetics 12:527-542. 

Cardinal, C. J. 2011. Greater Sage-Grouse ecology, movements, and habitat use related to land use 

patterns in the vicinity of Bear Lake, 2010 Progress Report. Submitted to Idaho Department of Fish 

and Game, Utah Division of Wildlife Resources, Bureau of Land Management, U.S. Fish and Wildlife 

Service, U.S. Forest Service, Idaho Department of Lands, Idaho Governor‟s Office of Species 

Conservation, East Idaho Uplands Sage-Grouse Local Working Group, Rich County Coordinated 

Resources Management Working Group, Wyoming Department of Game and Fish, and Southwest 

Wyoming Sage-grouse Local Working Group. Utah State University. 11pp. 

Carpenter, J. E., A. Aldridge, and M. S. Boyce. 2010. Sage-grouse habitat selection in winter in Alberta. 

Journal of Wildlife Management 74:1806-1814. 

Cas, M. 2010. Disturbances and predation on Capercaillie at leks in Alps and Dinaric Mountains. 

Sumarski List 134:487-495. 

Cox, R., D. Newborn, D. Baines, C. J. Thomas, and T. N. Sherratt. 2010. No evidence for resistance to 

Fenbendazole in Trichostrongylus tenuis, a nematode parasite of the Red Grouse. Journal of Wildlife 

Management 74:1799-1805. 

Doherty, K. E., J. D. Tack, J. S. Evans, and D. E. Naugle. 2010. Mapping breeding densities of Greater 

Sage-Grouse: A tool for range-wide conservation planning. Bureau of Land Management Completion 

Report: Interagency Agreement # L10PG00911. 31pp. 

32


Drummer, T. D., R. G. Corace III, S. J. Sjogren. 2011. Sharp-tailed Grouse lek attendance and fidelity in 

upper Michigan. Journal of Wildlife Management 75:311-318 

Ehrbar, R., K. Bollmann, and P. Mollet. 2011. Ein Sonderwaldreservat für das Auerhuhn – das Beispiel 

Amden (Kanton St. Gallen). [A special forest reserve for the Capercaillie – the model of Amden 

(Canton St Gallen).] Schweizerische Zeitschrift fur Forstwesen 162:11-21. (In German with English 

abstract). 

Evers, L. 2010. Modeling Sage-Grouse habitat using a state-and-transition model. Ph. D. Dissertation, 

Oregon State University. 168pp. 

Fedy, B. C., and K. E. Doherty. 2011. Population cycles are highly correlated over long time series and 

large spatial scales in two unrelated species: Greater Sage-Grouse and cottontail rabbits. Oecologia 

165:915-924. 

Francis, I. 2010. A survey of Black Grouse in north-east Scotland in 2009. Scottish Birds 30:290-294. 

Gavashelishvili, A., and Z. Javakhishvili. 2010. Combining radio-telemetry and random observations to 

model the habitat of near threatened Caucasian Grouse Tetrao mlokosiewiczi. Oryx 44:491-500. 

Gillan, J. K., and E. K. Strand. 2010. Sage-grouse habitat in Idaho: a practical guide for land owners and 

managers. Contribution Number 1048 of the Idaho Forest, Wildlife and Range Experiment Station, 

University of Idaho, Moscow. 66pp. 

Guttery, M. R. 2011. Ecology and management of a high elevation southern range Greater Sage Grouse 

population: vegetation manipulation, early chick survival, and hunter motivations. Ph. D. Dissertation. 

Utah State University. 119 pp. 

Habibzadeh, N., M. Karami, and A. Tarinejad. 2010. Caucasian Black Grouse (Tetrao mlokosiewiczi) 

breeding display sites selection in Arasbaran Region, East Azerbaijan, Iran. Russian Journal of 

Ecology 41:450-457. 

Hansen, C. P., J. J. Millspaugh, and M. A. Rumble. 2011. Occupancy modeling of Ruffed Grouse in the 

Black Hills National Forest. Journal of Wildlife Management 75:71-77. 

Hansen, C. P., M. A. Rumble, and J. J. Millspaugh. 2011. Ruffed Grouse selection of drumming sites in 

the Black Hills National Forest. American Midland Naturalist 165:400-411 

Hansen, J. L. H. Hansen, and N. M. Schmidt. 2010. Bird monitoring at Zackenberg, northeast 

Greenland, 2007. Bird Populations 10:56-67. (Rock Ptramigan). 

Hayama, S., C. Hattori, Y. Ohata, and N. Mitano. 2010. Body mass changes of Japanese Rock 

Ptarmigan Lagopus muta japonicus in captivity. Ornithological Science 9:149-155. 

Hoglund, J. J. K. Larsson, C. Corrales, G. Santafe, D. Baines, and G. Segelbacher. 2011. Genetic 

structure among Black Grouse in Britain: implications for designing conservation units. Animal 

Conservation XXX:XXX-XXX. (online early). 

Hunt, J. L., and T. L. Best. 2010. Vegetative characteristics of active and abandoned leks of Lesser 

Prairie-Chickens (Tympanuchus pallidicinctus) in southeastern New Mexico. Southwestern Naturalist 

55:477-487. 

Jarnevich, C. S., and M. K. Laubhan. 2011. Balancing energy development and conservation: a method 

utilizing species distribution models. Environmental Management XXX:XXX-XXX (online early). 

(Lesser Prairie-Chicken). 

Karlsson, K. 2010. Tjaderspel. [Capercaillie lek.] Anser 49:192-195. (in Swedish). 

Kielczynski, C. 2010. Wskaźniki biometryczne cietrzewi (Tetrao tetrix L.) w rozwoju osobniczym. 

[Biometrics rate of the Black Grouse (Tetrao tetrix L.) growth.] Aparatura Badawcza i Dydaktyczna 

15(2):125-136. (in Polish with English abstract). 

Klaus, S., and A. Christner. 2010. Ungewoehnliches Paarungsverhalten beim Birkhuhn Tetrao tetrix. 

[Atypical copulation behaviour in Black Grouse Tetrao tetrix.] Limicola 24: 140-145. (in German 

with English abstract). 

Lampila, P., E. Ranta, M. Monkkonen, H. Linden, and P. Helle. 2011. Grouse dynamics and harvesting 

in Kainuu, northeastern Finland. Oikos XXX:XXX-XXX (online early). (Capercaillie, Black 

Grouse, Hazel Grouse, Willow Grouse). 

Lande, U. S. 2011. Grouse – habitat relationships: monitoring, scale, and management. Ph. D. 

Dissertation. Swedish University of Agricultural Sciences. 46pp. (Capercaillie, Black Grouse, 

Willow Ptarmigan). 

Lees, J., R. Nudds, K.-A. Stokkan, L. Folkow, and J. Codd. 2010. Reduced metabolic cost of locomotion 

in Svalbard Rock Ptarmigan (Lagopus muta hyperborea) during winter. PLoS ONE 5(11):e15490. 

8pp. 

Lu, N., and Y.-H. Sun. 2011. Population viability analysis and conservation of Chinese Grouse Bonasa 

sewerzowi in Lianhuashan Nature Reserve, north-west China. Bird Conservation International 21: 49- 

58. 

33


Ludwig, G. X., R. V. Alatalo, P. Helle, and H. Siitari. 2010. Individual and environmental determinants 

of early brood survival in Black Grouse Tetrao tetrix. Wildlife Biology 16:367-378. 

Ludwig, G. X., R. V. Alatalo, P. Helle, and H. Siitari. 2010. Individual and environmental determinants 

of daily Black Grouse nest survival rates at variable predator densities. Annales Zoologici Fennici 

46:387-397. 

Ludwig, T., and I. Storch. 2011. Re-introduction and re-enforcement as a conservation measure for 

grouse. Gallinformed 4:18-21. 

Lukaszewicz, E., A. Kowalczyk, and Z. Rzonca. 2010. Characteristics of fresh semen of captive-bred 

Capercaillie Tetrao urogallus L. Zoo Biology XXX:XXX-XXX. (online early). 

Lycke, A., L. Imbeau, and P. Drapeau. 2011. Effects of commercial thinning on site occupancy and 

habitat use by Spruce Grouse in boreal Quebec. Canadian Journal of Forest Research 41:501-508. 

Martinez-Padilla, J., P. Vergara, L. Perez-Rodriguez, F. Mougeot, F. Casas, S. C. Ludwig, J. A. Haines, 

M. Zeineddine, and S. M. Redpath. 2011. Condition- and parasite dependent expression of a malelike 

trait in a female bird. Biology Letters XXX:XXX-XXX (online early). (Red Grouse). 

Mays, J. K., R. F. Silva, L. F. Lee, A. M. Fadly. 2010. Characterization of reticuloendotheliosis virus 

isolates obtained from broiler breeders, turkeys, and prairie chickens located in various geographical 

regions in the United States. Avian Pathology 39:383-389. 

McRoberts, J. T., M. J. Butler, W. B. Ballard, M. C. Wallace, H. A. Whitlaw, and D. A. Haukos. 2011. 

Response of Lesser Prairie-Chickens on leks to aerial surveys. Wildlife Society Bulletin 35:27-31. 

Miettunen, J. 2010. Winter food use and selection of Willow Grouse (Lagopus lagopus) in southern 

Finland. Suo 61(2):35-48. 

Moran, M. 2010. Estimas de abundancia y movimientos del urogallo cantábrico mediante análisis 

moleculares. [Estimate of abundance and movements of the Cantabrian Capercaillie by means of 

molecular analyses.] M. Sc. Thesis. Dept. Biología de Organismos y Sistemas, Universidad de 

Oviedo. (in Spanish). 

Naugle, D. E., K. E. Doherty, B. L. Walker, H. E. Copeland, and J. D. Tack. 2011. Sage grouse and 

cumulative impacts of energy development. Pp. 213-226 in P. R. Krausman and L. K. Harris (Eds.). 

Cumulative effects in wildlife management: impact mitigation. CRC Press, Boca Raton, Florida, 

USA. 

Norton, M. A., K. C. Jensen, A. P. Leif, T. R. Kirschenmann, and G. A. Wolbrink. 2010. Resource 

selection of Greater Prairie-Chicken and Sharp-Tailed Grouse broods in Central South Dakota. Prairie 

Naturalist 42:100-108. 

Nudds, R. L., L. P. Folkow, J. J. Lees, P. G. Tickle, K.-A. Stokkan, and J. R. Codd. 2011. Evidence for 

energy savings from aerial running in the Svalbard Rock Ptarmigan (Lagopus muta hyperborea). 

Proceedings of the Royal Society B XXX:XXX-XXX (online early). 

Ohman, K., L. Edenius, and G. Mikusinski. 2011. Optimizing spatial habitat suitability and timber 

revenue in long-term forest planning. Canadian Journal of Forest Research 41:543-551. (Hazel 

Grouse). 

Patricelli, G. L., and A. H. Krakauer. 2011. Assets and tactics in a mating market: economic models of 

negotiation offer insights into animal courtship dynamics on the lek. Current Zoology XXX:XXX- 

XXX (online early). (Greater Sage-Grouse). 

Patten, M. A., and J. F. Kelly. 2010. Habitat selection and the perceptual trap. Ecological Applications 

20:2148-2156. (Lesser Prairie-Chicken). 

Pearce-Higgins, J. W. 2010. Using diet to assess the sensitivity of northern and upland birds to climate 

change. Climate Research 45(special issue 25):119-U435. (Black Grouse, Rock Ptarmigan, Willow 

Ptarmigan). 

Peksa, L. 2010. Inwentaryzacja cietrzewia Tetrao tetrix i gluszca Tetrao urogallus w Tatrzanskim Parku 

Narodowym. [Census of Black Grouse Tetrao tetrix and Capercaillie Tetrao urogallus in the Tatra 

National Park.] Chronmy Przyrode Ojczysta 66:384-390. (in Polish with English abstract). 

Perez, T., J. F. Vazquez, F. Quiros, and A. Dominguez. 2011. Improving non-invasive genotyping in 

Capercaillie (Tetrao urogallus): redesigning sexing and microsatellite primers to increase efficiency 

on faeces samples. Conservation Genetics Resources XXX:XXX-XXX (online early). 

Pis, T. 2010. The link between metabolic rate and body temperature in galliform birds in thermoneutral 

and heat exposure conditions: The classical and phylogenetically corrected approach. Journal of 

Thermal Biology 35:309-316. 

Prather, P. R. 2010. Factors affecting Gunnison Sage-Grouse (Centrocercus minimus) conservation in 

San Juan County, Utah. Ph. D. Dissertation. Utah State University. 126pp. 

Rojas, M., I. Gonzales, M. A. Pavon, N. Pegels, P. A. Hernandez, T. Garcia, and R. Martin. 2011. 

Development of a real-time PCR assay to control the illegal trade of meat from protected Capercaillie 

species (Tetrao urogallus). Forensic Science International XXX: XXX-XXX (online early). 

34


Sandercock, B. K., E. B. Nilsen, H. Broseth, and H. C. Pedersen. 2011. Is hunting mortality additive or 

compensatory to natural mortality? Effects of experimental harvest on the survival and cause-specific 

mortality of Willow Ptarmigan. Journal of Animal Ecology 80:244-258. 

Sands, J. P, and M. D. Pope. 2010. A survey of galliform monitoring programs and methods in the 

United States and Canada. Wildlife Biology 16:342-356. 

Schaublin, S., and K. Bollmann. 2011. Winter habitat selection and conservation of Hazel Grouse 

(Bonasa bonasia) in mountain forests. Journal of Ornithology 152:179-192. 

Selas, V., G. A. Sonerud, E. Framstad, J. A. Kalas, S. Kobro, H. B. Pedersen, T. Kr. Spidso, and O. Wiig. 

2011. Climate change in Norway: warm summers limit grouse reproduction. Population Ecology 

53:361-371. (Black Grouse, Capercaillie, Willow Grouse). 

Signorell, N., S. Wirthner, P. Patthey, R. Schranz, L. Rotelli, and R. Arlettaz. 2010. Concealment from 

predators drives foraging habitat selection in brood-rearing Alpine Black Grouse Tetrao tetrix hens: 

habitat management implications. Wildlife Biology 16:249-257. 

Skrip, M. M. 2010. Fall-winter survival, habitat, and long-term population change of Ruffed Grouse in 

New York State. M. Sc. Thesis. State University of New York College of Environmental Science and 

Forestry, Syracuse. 120pp. 

Slipantschuk, J., E. Ullner, M. D. Baptista, M. Zeineddine, and M. Thiel. 2010. Abundance of stable 

periodic behavior in a Red Grouse population model with delay: a consequence of homoclinicity. 

Chaos 20(4):045117. 

Stevens, B. S., K. P. Reeese, and J. W. Connelly. 2011. Survival and detectability bias of avian fence 

collision surveys in sagebrush steppe. Journal of Wildlife Management 75:437-449. (Greater Sage- 

Grouse). 

Summers, R. W., D. Dugan, and R. Proctor. 2010. Numbers and breeding success of Capercaillies 

Tetrao urogallus and Black Grouse T. tetrix at Abernethy Forest, Scotland. Bird Study 57:437- 

446. 

Svobodova, J., G. Segelbacher, and J. Hoglund. 2011. Genetic variation in Black Grouse populations 

with different lekking systems in the Czech Republic. Journal of Ornithology 152:37-44. 

Thiel, D., S. Jenni-Eiermann, R. Palme, and L. Jenni. 2011. Winter tourism increases stress hormone 

levels in the Capercaillie Tetrao urogallus. Ibis 153:122-133. 

Tornberg, R., P. Helle, and E. Korpimaki. 2011. Vulnerability of Black Grouse hens to Goshawk 

predation: result of food supply or predation facilitation? Oecologia XXX:XXX-XXX (online early). 

Unger, C., and S. Klaus. 2010. Einfluss der topographischen Faktoren Hoehenlage, Hangneigung und 

Exposition auf die Habitatwahl umgesiedelter russischer Auerhuehner Tetrao urogallus in Thueringen. 

[Influence of the topographical factors altitude, slope inclination, and aspect on habitat choice of 

Capercaillies Tetrao urogallus translocated from Russia to Thueringen.] Anzeiger des Vereins 

Thueringer Ornithologen 7:49-56. (in German with English abstract). 

Villanueva, R. L. 2011. Implementación de una red telemática que facilite el anillamiento de la hembra 

de urogallo. [Deployment of an IP camera network which aims to determine the time when Western 

Capercaillie hatch out of eggs, in order to ring the female birds.] B. Sc. Thesis. Universitat Oberta de 

Catalunya. (in Spanish with English abstract). 

Vodehnal, B., A. Pierson, T. Doeden, M. Schultz, and M. D. Blount. 2010. Location of Sharptailed 

Grouse and Greater Prairie-Chicken display grounds in relation to NPPD Ainsworth Wind Energy 

Facility – 2006-2010. Annual Report, Nebraska Game and Parks Commission. 9pp. 

Warren, P. 2010. Black Grouse recovery in northern England. British Wildlife 21(6):383-391. 

Webster, L. M. I., L. V. Mello, F, Mougeot, J. Martinez-Padilla, S. Paterson, and S. B. Piertney. 2011. 

Identification of genes responding to nematode infection in Red Grouse. Molecular Ecology 

Resources 11:305-313. 

Webster, L. M. I., S. Paterson, F. Mougeot, J. Martinez-Padilla, and S. B. Piertney. 2011. 

Transcriptomic response of Red Grouse to gastro-intestinal nematode parasites and testosterone: 

implications for population dynamics. Molecular Ecology 20:921-931. 

Wegge, P., and J. Rolstad. 2011. Clearcutting forestry and Eurasian boreal forest grouse: Long term 

monitoring of sympatric Capercaillie Tetrao urogallus and Black Grouse T. tetrix reveals unexpected 

effects on their population performances. Forest Ecology and Management 261:1520-1529. 

Whiting, J. C., M. Morales, J. Handy, and K. Edwards. 2010. Ecological assessment of Sage Grouse in 

the area surrounding T-24 and T-25. Report prepared for CH2M-WG Idaho, LLC. Subcontract 

00732290. S. M. Stoller Corporation, Idaho Falls, Idaho. 9pp. 

Willebrand, T., M. Hornell-Willebrand, and L. Asmyhr. 2011. Willow Grouse bag size is more sensitive 

to variation in hunter effort than to variation in Willow Grouse density. Oikos XXX:XXX-XXX. 

(online early). 

35


Wilson, S., and K. Martin. 2011. Life-history and demographic variation in an alpine specialist at the 

latitudinal extremes of the range. Population Ecology XXX:XXX-XXX (online early). (White-tailed 

Ptarmigan). 

Wolfe, D. H., M. A. Patten, and L. C. Larsson. 2010. Status, distribution, and ecology of White-tailed 

Ptarmigan (Lagopus leucura) in the Sangre de Cristo Mountains, New Mexico. Final Report. 

Submitted to New Mexico Department of Game and Fish. Contract Number 10-516-0000-00018. 

15pp. 

Yang, C., Y. Fang, and Y.-H. Sun. 2011. Winter space use and social behaviors of Chinese Grouse 

(Bonasa sewerzowi) at Lianhuashan mountains, Gansu, China. Journal of Ornithology 152:297-305. 

36


ERRATUM 

Error in the article on black grouse in issue 40 

In the paper Population ecology of black grouse on north-east Scottish moorland in 1956–82 by 

Adam Watson (Grouse News 40: 6-13) an error has been found in the results, heading breeding success. 

On page 11 in the paragraph after Table 7, first line, “correlated negatively” should be “correlated 

positively”, as in the summary, and the negative sign in front of the r value is incorrect and should be 

deleted. The paragraph should be as follows: Breeding success of greyhens was strongly correlated 

positively with total June rainfall (n = 5, r = 0.973, P = 0.0054, adjusted R 2 = 0.928). The incorporation of 

June mean temperature or the number of June rain days reduced the proportion of variation accounted for 

in a multiple regression. 

37


SNIPPETS 

The effect of climate change on grouse populations in Norway 

Different species of grouse and vole may have peak population after peaks in the seed crop of bilberry 

Vaccinium myrtillus because of reduced levels of feeding deterrents in bilberry plants. In this paper we 

predicted that grouse reproduction may depend also on temperatures in June–September in the 2 previous 

years, because bilberry plants will be less exhausted after a high seed crop in or after warm summers, and 

thus rebuild their chemical defence more quickly. After peak years of bilberry, the populations of 

capercaillie Tetrao urogallus and bank vole Myodes glareolus in southern Norway were negatively 

related to summer temperatures in the previous year or previous 2 years. In five areas in Norway chick 

production in willow ptarmigan Lagopus lagopus was negatively related to summer temperatures in the 2 

previous years. Densities of capercaillie and black grouse Tetrao tetrix were more likely to peak in vole 

peak years at high altitudes in eastern Norway in the autumn, where summer temperatures are low. The 

paper concludes that high summer temperatures may limit grouse reproduction through the effect on 

bilberry plants and that a warm climate thus adversely affects population levels of grouse. 

Selås, V., Sonerud, G.A., Framstad, E., Kålås, J.A., Kobro, S., Pedersen, H.B., Spidsø, T.K. and Wiig, Ø. 

2010. Climate change in Norway: warm summers limit grouse reproduction. - Population Ecology 

53: 361-371. 

Tor Kristian Spidsö, Editor Grouse News, TKS.Grouse@gmail.com 

Amazing Species required 

Rachel Roberts 

Dear All 

Many of you last year kindly assisted me in contributing to the profiles of threatened species for daily 

publication on the new IUCN Species of the Day website. The website was very successful and we 

finished the end of the year with a dedicated following of over 4,000 users on the social networking site 

Twitter. We also attracted interest from a UK publishing company, which is now producing a wonderful 

coffee table book of all of last year‟s profiles which will be available later this year. 

On the back of this success we have kept the website on, but under the new name of „Amazing 

Species‟. As I am now working on this alone, the profiles are weekly instead of daily but the format is 

still exactly the same. http://www.iucnredlist.org/amazing-species. 

I would be extremely grateful if I could ask for your support again this year, and to nominate one or 

two threatened species that you wish to see profiled on the website. We would prefer the species to be in 

any of the RL categories, but if you are in a group lacking verified assessments then please do not let this 

prevent you nominating species. View this as a great way to raise the profile of your species, as we do get 

lots of visits to the site and are frequently told how interesting and useful it is especially as an education 

tool. 

As I am now running this single-handedly, it would be wonderful if you could assist me by writing the 

short text required. But, if this is not feasible, then please just send me the names of the species you wish 

to profile and hopefully I will get around to writing it for you to review. I also need a high resolution 

photo and permission granted to use that photo. Thank you so much for your ongoing support on this, it is 

hugely appreciated. 

Instructions for Amazing Species text – 170 words: 

Intro: Scientific and common name and Red List category if it has one, and distribution and range 

with a brief description. Example: Delacour‟s Langur, Trachypithecus delacouri, is listed as „Critically 

Endangered‟ on the IUCN Red List of Threatened Species TM . Found only in north-central Vietnam, 

Delacour‟s Langur is one of the rarest and most threatened primates on Earth. 

Middle Paragraph: Threats to the species. 

End Paragraph: Conservation measures being undertaken or that should be undertaken. 

Rachel Roberts, Executive Assistant IUCN Species Survival Commission, The Innovation Centre, 

University of Bath, Carpenter House, First Floor, Broad Quay, Bath, BA1 1UD, UK, 

sscchairoffice@iucn.org. http://www.iucnredlist.org/species-of-the-day. 

38


Japanese rock ptarmigan 

Japanese rock ptarmigan Lagopus muta japonica (Photos Hiroshi Nakamura) 

39

Grouse News 41.pdf - Galliformes-sg.org

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