Habitat characteristics of Capercaillie Tetrao urogallus display grounds in southeastern Norway

Habitat composition of 39 capercaillie Tetrao urogallus display grounds was compared with available and unused habitats in two coniferous forest areas in southeastern Norway. Macro-habitat was described with respect to forest successional stage and topography. Micro-habitat was described with respect to microtopography, plant association, and forest stand structure. Totally, 28 variables were measured. A pronounced macro-habitat preference was found for elevated areas with forests older than 60-70 years. Micro–habitat composition varied considerably. However, a slight but significant preference was found for heterogeneous, open-spaced understory, and for visibility of more than 30 m. Suitable micro-habitats were abundant, and distribution of display grounds were mainly limited by macro-habitat characteristics, i.e. elevated, old forest areas.     

Activity patterns of Capercaillie, Tetrao urogallus, during winter

Winter activity patterns of 16 radio-marked capercaillie, Tetrao urogallus , were studied during 1981–83 at Varaldskogen, SE Norway. Activity was confined to the light hours of the day, closely following the photoperiod. Diel distribution showed a major peak near sunset during every month. A second peak was found in the morning. This peak was comparable to the evening peak in early and late winter, but nearly disappeared in midwinter. Ambient temperature rather than photoperiod seemed to be the important proximate factor responsible for the changes in morning activity. By postponing activity to the afternoon, capercaillie avoids energy expenditure during the coldest morning hours.
Total daily activity (TDA) during the period November-April averaged about 3 h for both sexes. TDA of cocks was U-shaped with the lowest values (2.0 h) in December-January, whereas TDA of hens was fairly stable during November-March with mean value of 2.7 h, increasing abruptly to 4 h in April. TDA during December-January is believed to express the time used for feeding in pine trees. The significantly higher TDA of hens (26%) compared with cocks during this period may be explained by a higher relative heat loss of hens (body weights: hens 2.0 kg, cocks 4.3 kg). The increased level of activity during early and late winter among cocks was probably due to display and related territorial behaviour, whereas increased activity of hens during April probably was caused by a shift from feeding in pines to more time-consuming, selective feeding on the ground     

Capercaillie Tetrao urogallus in Scotland - demography of a declining population

Capercaillie Tetrao urogallus numbers declined in Scottish forests between 1992 and 1997, although the rate (mean 16% per year, 95% CL 1–30%) differed among forests. Hens declined more quickly than cocks despite the sex ratio in chicks favouring females. A probable explanation is that adult cocks lived longer. Juvenile survival was less than adult survival, largely because juveniles were more likely to die from hitting fences. Although poor reproduction was probably the main cause of the decline, this might not have occurred had fence strikes been substantially fewer.     

Winter tourism increases stress hormone levels in the Capercaillie Tetrao urogallus

Montane and alpine habitats in Europe remained relatively undisturbed until the beginning of the last century. Today, outdoor recreation activities are a major economic factor in alpine regions. Many tourism areas coincide with winter habitats of shy and endangered species. The Western Capercaillie Tetrao urogallus has suffered from rapid population declines during recent decades over much of its range. In central Europe, many Capercaillie are exposed to intensive human recreation activities in their habitats, which may contribute to this decline. However, little is known about their susceptibility to human recreation activities. This study assessed whether human recreation in winter evokes physiological stress responses in several populations of Capercaillie. During two winters, we sampled 1130 Capercaillie droppings in Germany and Switzerland of populations at various distances from winter recreation activities and measured concentrations of faecal corticosterone metabolites. Capercaillie in relatively dense and homogeneous mountain forests dominated by Norway Spruce Picea abies showed markedly increased stress hormone levels closer to locations with winter recreation activity. However, this physiological response to human recreation was not detectable in forests dominated by various pine species and a heterogeneous structure. Capercaillie may be particularly sensitive to recreation because any factor affecting their fine‐tuned physiological and behavioural adaptations to survive under harsh winter conditions may lead to harmful fitness costs.     

Simple and Effective Methods of Freezing Capercaillie (Tetrao urogallus L.) Semen

A continuous decline in the number and range of capercaillie (Tetrao urogallus L.) in many European countries can be observed, mostly due to habitat destruction by human activity, unecological forestry management, and increased density of natural predators. Ex situ in vitro gene banks provide a unique opportunity to preserve the genetic material for future generations. Simple and effective cryopreservation methods for capercaillie semen are discussed. Semen was collected from seven males kept in the Capercaillie Breeding Centre at Forestry Wisła in Poland. Within five minutes after collection, ejaculates were diluted with EK diluent, then divided into two parts, and subjected to two freezing procedures: in pellets and in straws. In fresh semen, ejaculate clearness, viscosity, color and volume, as well as sperm concentration, motility and morphology, were evaluated, while in frozen-thawed semen only motility and morphology of sperm were determined. Fertilizing ability of thawed semen was examined for samples frozen in straws. Significant (P<0.05) differences between individual males were found in relation to the majority of fresh semen traits: ejaculate volume averaged 102.1 µL (varying from 49.0 to 205.0); average sperm concentration was 632.5 x10⁶ mL^-1 (178.8–1257.1); percentage of live normal cells varied from 39.2 to 70.3% (58.7% on an average); percentage of motile cells ranged from 76.0 to 85.7%) and motility parameters were male dependent, as well. Both cryopreservation methods had a negative effect on morphology and motility of frozen-thawed semen; however, the straw method yielded 60.7% and the pellet method 42.5% of live cells in total in thawed semen (P<0.05), while the number of live normal (intact) cells was similar (22.4 and 22.2%, respectively). Egg fertility varied between 77.8 and 91.7% (average 84.4%). Both freezing procedures seem to be effective in obtaining acceptable viability and high fertilizing potency of thawed sperm and can be used to create a gene bank of capercaillie semen.     

The bacterial microbiota in the ceca of Capercaillie (Tetrao urogallus) differs between wild and captive birds

The diet of wild capercaillie differs strongly between seasons. Particularly during winter, when energy demands are high and the birds forage solely on coniferous needles, microbial fermentations in the ceca are considered to contribute significantly to the energy requirement and to the detoxification of the resinous diet. Here, we present the first cultivation-independent analysis of the bacterial community in the cecum of capercaillie, using the 16S rRNA gene as a molecular marker. Cloning and fingerprinting analyses of cecum feces show distinct differences between wild and captive birds. While certain lineages of Clostridiales, Synergistetes, and Actinobacteria are most prevalent in wild birds, they are strongly reduced in individuals raised in captivity. Most striking is the complete absence of Megasphaera and Synergistes species in captive capercaillie, which are characterized by a large abundance of Gammaproteobacteria closely related to members of the genus Anaerobiospirillum, bacteria that are commonly connected with intestinal dysfunction. The community profiles of cecum content from wild birds differed between summer and winter season, and the cecum wall may be an important site for bacterial colonization. Our results corroborate the hypothesis that the bacterial community in the ceca of tetraonid birds changes in response to their highly specialized seasonal diets. Moreover, we propose that the observed differences in community profiles between wild and captive capercaillie reflects a disturbance in the bacterial microbiota that compromises the performance of the cecum and may be responsible for the high mortality of captive birds released into nature.     

Rain, breeding success and distribution of Capercaillie Tetrao urogallus and Black Grouse Tetrao tetrix in Scotland

The number of chicks reared by Capercaillie was inversely related to the number of days with rain during and just after their hatching. However, rain had no detectable effect on the breeding success of Black Grouse on the same study area. The breeding distribution of Capercaillie in Scotland is more limited than that of Black Grouse, which extends into rainier areas. It is speculated that sexual selection for large size in Capercaillie has indirectly caused their chicks to be more vulnerable to rain and, consequently, has limited their distribution to less rainy areas.     

Nesting habitat,clutch size and nest failure of Capercaillie Tetrao urogallus in Scotland

CapsuleThe mean clutch size was 7.3, and habitat and year did not significantly affect nest failure rate.     

Habitat selection and diet of Western Capercaillie Tetrao urogallus in an atypical biogeographical region

Ecological features and conservation requirements of populations at the latitudinal limits of a species’ geographical range frequently differ from those in other parts of the range. Identifying such differences is key to implementing effective conservation strategies for threatened range‐edge populations especially, in the context of rapid global warming, at the lower‐latitude range edge. We studied habitat selection and diet of the endangered Cantabrian Capercaillie Tetrao urogallus cantabricus in a recently discovered population at the southernmost edge of the sub‐species’ range. This is the only Western Capercaillie population in the Mediterranean biogeographical region. We combined non‐systematic surveys based on questionnaires, reports and field sampling with data from radiotracking to assess habitat selection. Diet was surveyed by micro‐histological methods from droppings collected in the new population, which inhabits Pyrenean Oak Quercus pyrenaica forests and Scots Pine Pinus sylvestris plantations, and in two Cantabrian populations inhabiting Eurosiberian forests. Capercaillie preferred large (> 500 ha) and medium‐sized (100–500 ha) Pyrenean Oak forest fragments and large Scots Pine plantations. Forest fragments smaller than 100 ha and non‐forested habitats were always avoided. Diet differed markedly between Mediterranean and Eurosiberian populations. Bilberry Vaccinium myrtillus is common in the diet of most Capercaillie populations but was scarce in the study area and so was rare in the diet of the new population. Instead, Rockrose Halimium lasianthum was described for the first time as a major food resource for the Capercaillie and was consumed in autumn and winter. Pine needles were also heavily consumed in winter. We document for the first time the strong preference of Capercaillie for Pyrenean Oak forests and a moderately high consumption of the leaves, buds and acorns of this tree species throughout the year. Habitat selection and diet of this Mediterranean population differ from those of the core Cantabrian and other populations. Our results suggest a wider environmental tolerance (phenotypic plasticity) in the species than previously recognized. We advocate specific protection for this unique range‐edge Capercaillie population and its Pyrenean Oak forest habitat.     

A first test of unattended,acoustic recorders for monitoring Capercaillie Tetrao urogallus lekking activity

Capsule: Automated acoustic recording can be used as a valuable survey technique for Capercaillie Tetrao urogallus leks, improving the quality and quantity of field data for this endangered bird species. However, more development work and testing against traditional methods are needed to establish optimal working practices.

Aims: This study aims to determine whether Capercaillie vocalizations can be recognized in lek recordings, whether this can be automated using readily available software, and whether the number of calls resulting varies with location, weather conditions, date and time of day.

Methods: Unattended recording devices and semi-automated call classification software were used to record and analyse the display calls of Capercaillie at three known lek sites in Scotland over a two-week period.

Results: Capercaillie calls were successfully and rapidly identified within a data set that included the vocalizations of other bird species and environmental noise. Calls could be readily recognized to species level using a combination of unsupervised software and manual analysis. The number of calls varied by time and date, by recorder/microphone location at the lek site, and with weather conditions. This information can be used to better target future acoustic monitoring and improve the quality of existing traditional lek surveys.

Conclusion: Bioacoustic methods provide a practical and cost-effective way to determine habitat occupancy and activity levels by a vocally distinctive bird species. Following further testing alongside traditional counting methods, it could offer a significant new approach towards more effective monitoring of local population levels for Capercaillie and other species of conservation concern.     

Viability and Management of an Endangered Capercaillie (Tetrao urogallus) Metapopulation in the Jura Mountains, Western Switzerland

The populations of Capercaillie (Tetrao urogallus), the largest European grouse, have seriously declined during the last century over most of their distribution in western and central Europe. In the Jura mountains, the relict population is now isolated and critically endangered (about 500 breeding adults). We developed a simulation software (TetrasPool) that accounts for age and spatial structure as well as stochastic processes, to perform a viability analysis and explore management scenarios for this population, capitalizing on a 24 years-long series of field data. Simulations predict a marked decline and a significant extinction risk over the next century, largely due to environmental and demographic stochasticity (average values of life-history parameters would otherwise allow stability). Variances among scenarios mainly stem from uncertainties about the shape and intensity of density dependence. Uncertainty analyses suggest to focus conservation efforts on enhancing, not only adult survival (as often advocated for long-lived species), but also recruitment. The juvenile stage matters when local populations undergo extinctions, because it ensures connectivity and recolonization. Besides limiting human perturbations, a silvicultural strategy aimed at opening forest structure should improve the quality and surface of available patches, independent of their size and localization. Such measures are to be taken urgently, if the population is to be saved.     

Habitat selection and diet of the Capercaillie Tetrao urogallus in Abernethy Forest,Strathspey, Scotland

Capsule Woodland characteristics are important for fully grown birds and for broods.

Aims To describe habitat selection and diet of the Capercaillie in ancient native pinewoods and Scots Pine Pinus sylvestris plantations, in order to provide a guide for forest management.

Methods Tree selection was studied in winter by locating droppings under trees along transects, and comparing those trees used by Capercaillie with those not used. Habitat selection by broods was studied by following radiotagged chicks or females with broods. Diet was described from the analysis of droppings.

Results In winter, Capercaillie preferred Scots Pines with wide crowns and large branches, and little surrounding Heather Calluna vulgaris, but avoided forest tracks. Female Capercaillie occurred in stands with a higher density of pines compared to males. The diet of fully grown birds comprised primarily Scots Pine needles between October and April. Scots Pine pollen cones were eaten during May, June and July. Bilberry Vaccinium myrtillus leaves and fruit were eaten in summer and autumn. Habitats used by broods had a higher percentage of Vaccinium sp. and smaller Scots Pines compared with random sites. Chicks hatch at the beginning of June, and in their first two weeks, their diet comprised Formicidae, Lepidoptera larvae and Coleoptera. In July and August, Lepidoptera larvae were rarely eaten but a higher percentage of chicks took vegetable material (particularly Bilberry fruit). In June, smaller numbers of invertebrates were generally trapped in mires of wooded bogs compared with areas of Heather and Vaccinium sp. in old woodland. Areas of Vaccinium sp. had the highest numbers of Formicidae, Coleoptera and Diptera, and especially Lepidoptera larvae. Numbers of Lepidoptera larvae declined during June and July as larvae pupated, explaining the reduction of Lepidoptera larvae in the diet of older chicks.

Conclusions Habitat management for Capercaillie should aim to provide old Scots Pine forest with a Vaccinium-rich field layer. Within Scotland, this may entail increasing the tree density in ancient native pinewoods but decreasing it in pine plantations. The removal or closure of tracks might increase the amount of woodland available to Capercaillie, by reducing human disturbance.     

Timing of breeding in Black Grouse Tetrao tetrix and Capercaillie Tetrao urogallus and distribution of insect food for the chicks

In Perthshire (Scotland), Black Grouse Tetrao tetrix broods less than 3 weeks old preferred herb-rich grassy flushes with Bog Myrtle Myrica gale , whereas in the northern Pennines (England), all broods were found on grass-dominated moorland or enclosed rough grazing habitats chiefly of grass and rushes. In both areas, broods used habitats with taller vegetation that had more invertebrates, particularly moth caterpillars and sawfly larvae (Sym-phyta, Hymenoptera). Within preferred habitats, broods were found in vegetation containing more flies (Diptera), plant bugs (Hemiptera) and ichneumon flies (Hymenoptera) than at randomly chosen sites. The main hatch of Black Grouse chicks in Perthshire coincided with the peak availability of moth caterpillars preferred by chicks. Broods in the Pennines hatched 1 week later than those in Scotland, and this later breeding in the south may be an adaptation to utilize the later peak biomass of sawfly larvae, the chief insect prey there. Data collected for Capercaillie Tetrao urogallus suggest a similar degree of synchronization between their peak hatch and the availability of moth caterpillars on Bilberry Vaccinium myrtillus.     

Comparative Examination of Capercaillie (Tetrao urogallus L.) Behaviour Responses and Semen Quality to Two Methods of Semen Collection

Artificial insemination (AI) is very helpful in solving the reproductive and biodiversity problems observed in small, closed avian populations. The successful production of fertilized eggs using AI is dependent on the collection of good quality semen. Two methods of male sexual stimulation and semen collection from captive kept capercaillie (Tetrao urogallus L.), one of the most seriously endangered grouse species in Europe, are compared in this study. Ejaculates were obtained either with the use of a dummy female or by the dorso-abdominal massage method. Differences in the individual responses of the males to the two methods of semen collection as well as in their semen quality were noted. Only sperm concentration (432.4 x 10⁶ mL^-1 with dummy female and 614.5 x 10⁶ mL^-1 for massage method) was significantly affected by capercaillie stimulation method. Sperm motility and morphology were not affected (P≥0.05). Thus, for semen collection from captive kept capercaillie both methods can be used successfully. The dummy female can be an alternative to dorso-abdominal massage method, commonly used for semen collection from domesticated bird species.     

Juvenile sex ratio in relation to breeding success in Capercaillie Tetrao urogallus and Black Grouse T. tetrix

We examined the relationship between breeding success and juvenile sex ratio in two gamebirds with differing degrees of sexual size dimorphism. We used breeding success as an indicator of the level of environmental stress the birds had experienced. Capercaillie Tetrao urogallus and Black Grouse T. tetrix wing samples were collected from hunters in northern Sweden during 1977-86. In both species, annual juvenile sex ratio (percentage of males) increased with annual breeding success, i.e. with reduced environmental stress. We attribute this to the higher energy requirements during growth, vulnerability to environmental stress, and mortality of juvenile males compared with females, especially under adverse conditions. Moreover, the relationship between breeding success and juvenile sex ratio was stronger in Capercaillie than in Black Grouse. This reflects the higher degree of sexual size dimorphism in Capercaillie, and consequent larger mortality difference between juvenile males and females.     

Estimating Population Size for Capercaillie (Tetrao urogallus L.) with Spatial Capture-Recapture Models Based on Genotypes from One Field Sample

We conducted a survey of an endangered and cryptic forest grouse, the capercaillie Tetrao urogallus, based on droppings collected on two sampling occasions in eight forest fragments in central Switzerland in early spring 2009. We used genetic analyses to sex and individually identify birds. We estimated sex-dependent detection probabilities and population size using a modern spatial capture-recapture (SCR) model for the data from pooled surveys. A total of 127 capercaillie genotypes were identified (77 males, 46 females, and 4 of unknown sex). The SCR model yielded atotal population size estimate (posterior mean) of 137.3 capercaillies (posterior sd 4.2, 95% CRI 130–147). The observed sex ratio was skewed towards males (0.63). The posterior mean of the sex ratio under the SCR model was 0.58 (posterior sd 0.02, 95% CRI 0.54–0.61), suggesting a male-biased sex ratio in our study area. A subsampling simulation study indicated that a reduced sampling effort representing 75% of the actual detections would still yield practically acceptable estimates of total size and sex ratio in our population. Hence, field work and financial effort could be reduced without compromising accuracy when the SCR model is used to estimate key population parameters of cryptic species.