In general, large-scale human land use strongly affects Capercaillie populations in Finland. Thus, it seems obvious that large spatial scales should be considered in the conservation and management of Capercaillie. The amount of forest is important, especially in the areas of strong human impact, i.e. in the areas where forest landscapes have been altered and the spatial and age-class structure have been modified (II, III). In particular, the forest management around lekking areas should consider larger spatial scales than before (III). It is not enough to perform carefully planned cuttings inside the lekking areas (inside the radius of 1000 m, e.g.
Helle et al. 1994) if the forest cover at the surroundings (at the radius of 3000 m) has already been cut to a level that is too low for the lekking areas to persist.
I would recommend that the management of Capercaillie is performed at two levels (i.e. multi-level planning, see also Suchant &
Braunisch 2004, Braunisch &
Suchant 2007). The first priority should be to secure the large regional-scale forest cover (II, III).
The second priority should be to promote fine-grained mosaic-like structure of forests inside the separate forest patches, in particular close to lekking areas (III, see also Miettinen 2009). Along with the improving lekking area persistence,
these targets might also enhance the amount and quality of the Bilberry-rich brood rearing habitats (e.g.
Lakka & Kouki 2009). However, some additional local-scale measures like small predator control might be needed to further improve Capercaillie breeding success (e.g.
Kurki et al. 1997).
The current guidelines for forest management practices in Finland (Heinonen et al. 2005, Metsätalouden kehittämiskeskus Tapio 2006) pay attention to Capercaillie, too. The results of this thesis, however, add new aspects to these guidelines. For instance, the Finnish state enterprise Metsähallitus plans forestry on regional scale, and promotes the overall variability of forests (Heinonen et al. 2005).
However, the practical instructions for Capercaillie lekking site management consider only an area up to 1000 m from the lekking centre (Heinonen et al. 2005). This will not guarantee that larger-scale connectivity between leks is actually maintained. Thus, it is advised that larger-scale considerations shall be included in the management of Capercaillie lekking sites in the future. The management of local Capercaillie populations is especially challenging in private forests, where planning across forest stands and among several land owners is encouraged (Metsätalouden
kehittämiskeskus Tapio 2006), but rarely implemented.
Capercaillie is often considered an old-forest specialist, but this relationship is nowadays hard to detect, at least in such a straightforward manner as has traditionally been possible (e.g.
Swenson & Angelstam 1993).
However, because of the obvious restrictions to study the phenomena connected to older forests in our present-day ‘young forest landscapes’ (IV), I would be cautious about giving specific management recommendations considering forest age. It is possible, as noted already e.g. by Helle et al.
(1989), that it is not the forest age per se that is important for Capercaillie, but some other characteristics that are connected to older forests. These could include multi-layered forest structure with versatile cover and food, and the presence of large timber trees (see also Miettinen et al. 2009). How well Capercaillie can maintain viable populations in younger forests is a question that still needs to be resolved, considering both the lekking site viability (IV), and the other life-history stages of Capercaillie (I).
How should Capercaillie populations be managed in practice? Lindén et al. (2000) have already proposed a plan of large-scale forest corridors or
‘forest bridges’ through Finland.
These are 50 km- wide forest belts, with approximately one-third consisting of mature (> 40–50
years) forest, spanning from eastern Finland towards the west and south (Lindén et al. 2000). I suggest that functioning lekking sites are included as ‘nodes’ in these forest bridges and that the bridges are then built around the lekking areas. These measures would most likely enhance Capercaillie lekking area persistence (e.g. III), and thus benefit the local population viability (e.g. Helle et al.
1994, Wegge & Rolstad 2002).
Conserving the connectivity of forests on a large spatial scale could also offer a precaution to minimize risks connected to future range shifts stemming from climate change (Thomas & Lennon 1999).
To implement the new management guidelines and to create and maintain large-scale forest corridors, regional-level forestry planning and cooperation between forest owners will be needed. In principle, the implementation of the forest corridors probably requires that some modifications in rotation times and landscape patterns are addressed in specific regions.
However, the idea behind the ‘forest bridges’ is not to put the forests under a strict conservation regime, but to ensure that both the overall connectivity of the forests and the amount of older forest stay high enough in a landscape (Lindén et al.
2000). Conservation areas may complement these corridors, but mostly it is a question of managed forests. I anticipate that implementation of ‘Capercaillie programme’ requires better interaction between forest owners,
regional forestry authorities and game researchers. For example, results from game monitoring schemes could be more tightly included and applied in forestry plans. To support private land owners’ biodiversity-friendly forest use, tax reductions or other incentives are probably also needed.
For instance, the Forest Biodiversity Programme (METSO) is designed to offer a tool kit for enhancing private forest owners’ willingness to voluntarily protect or manage their forests in a biodiversity-friendly way (Ministry of the Environment 2010).
Capercaillie has several characteristics which make it highly attractive as a flagship species in the northern boreal forests. Firstly, the species is generally highly valued among land owners and hunters (e.g. Leinonen & Ermala 1995), guaranteeing its yearly abundance estimation through wildlife triangle counts performed by thousands of volunteers (Forsman et al. 2010).
Secondly, Capercaillie has umbrella-species characteristics, i.e. the preservation of Capercaillie will benefit also other wildlife (Pakkala et al. 2003). Lastly, Capercaillie has a special status in the Finnish culture:
it is traditionally considered as a focal symbol of healthy forests (Lindén 2002c). Thus, it is easy to see that Capercaillie deserves a special status in the Finnish biota.
Unfortunately, the future of this magnificent bird is far from being secured. Based on my results, the most important step forward in securing the Capercaillie is to think
beyond the forest stand level, that is, to consider a management unit covering tens or hundreds of square kilometers. This is a challenge that should be addressed in the near future in order to maintain the viability of Capercaillie populations in Finland.
Acknowledgements
First of all, I thank my supervisors Harto Lindén and Pekka Helle for being the best supervisors one can wish for.
Harto, you were the driving force behind the whole project and you took me underneath your wings from the very beginning. The ways you have supported me growing into an independent researcher have been so numerous that I cannot mention them all here. What has probably meant the most for me is your curious and open attitude towards the world and people, and your bit twisted sense of humour. You have always supported my ambitions, no matter how weird they are, and kept perfect balance between guidance and freedom. I’ve really enjoyed our journey together, thank you.
Pekka, because of the distance between Helsinki and Oulu you were forced to the role of ‘the second’ supervisor. However, I feel that your input has been priceless in balancing the sometimes strong opinions me and Harto had (even balancing between opinions in some cases). Your calm and reflective presence has been like cool and soothing water for me — you must have noticed that I might get a bit restive sometimes… You also offered me the most unforgettable nature experience ever by taking me in the middle of a lekking site in
Taivalkoski. Thank you.
During this project several people have helped me without counting hours. My warmest thanks to my co-authors: Ari Nikula, Jani Pellikka, Andy Lindén, Jonas Knape, Kai Norrdahl, Petri Suorsa, and Pentti Valkeajärvi. Special thanks to Ari:
you would deserve a credit being my third supervisor, thank you for your selfless help and support! Warm thanks also to my thesis advisory committee members Mikko Mönkkönen and Hanna Kokko. And thank you Jon Swenson and Jari Kouki for your thorough and timely review of this thesis.
From the Finnish Game and Fisheries Research Institute (RKTL), I want to thank especially Katja Holmala, Kaija Pönni-Susiluoto, Oili Vuorimies, Marcus Wikman, Nina Peuhkuri, Ilpo Kojola, Vesa Ruusila, and Eero Helle for their support and friendship. To represent ‘Ärkkis’ has meant a lot for me, and I am humbly
thankful for the opportunity. Thank you also to all those people, mainly hunters, who year after year collect the precious wildlife triangle data in Finland.
From my other home, the department, I would like to thank especially my
custodian Veijo Kaitala, my mentor Esa Ranta († 29.8.2008), always helpful Ilkka Teräs, and the Finnish School in Wildlife Biology, Conservation and Management (LUOVA), particularly Ilkka Hanski and Anni Tonteri. Without LUOVA funding and support I would not be at this point now. Other funding agents for my project
were Riistanhoito-Säätiö, Research Foundation of the University of Helsinki, Chancellor’s Travel Grant, and Finnish Concordia Fund.
I want to thank my friends at the University for your smiling faces, supportive comments, interesting lunch discussions, and parties. It’s been just amazing to work in the atmosphere you have created. Special thanks to Marja Järvenpää, Milla Niemi, Edward Kluen, Maaike de Heij, Wouter Vahl, Jostein Starrfelt, Christoph Meier, Maria Delgado, Christopher Wheat, Heini Kujala, Joona
Lehtomäki, Jussi Lehtonen, Katja Heubel, Phillip Gienapp, Mike Fowler, and the rest of the Integrative Ecology Unit, IKP. Thank you also the people in my ‘side-projects’, especially Minttu Heimovirta and Katja Bargum at Yle, and the whole Nisse-tiimi. You were all very important in keeping me attached to ‘the real world’ during this project.
Thank you Silja Lahti, Iina Reinikka, Piia Nordberg, Minna Markkanen, Liina Väänänen, Noora Huldén, Marja Hakala-Yatkin, Päivi Lehtonen, and Riitta
Koivikko for... being there. I have been blessed with so many close friends in my life. Thank you for sharing your moments with warm tea pots, cold beer classes, supportive shoulders, and laughter, with me. You never doubted my ability to cope this or other major projects in my life. Let’s always keep our friendship alive!
I sincerely thank my family: mum, dad, Eko, Henna, and Vilma, for their love and support. Iskä: kiitin sinua jo gradussa kaikesta siitä innosta ja ilosta luontoa ja sen ihmeitä kohtaan, jonka olet jotenkin onnistunut täysimääräisenä välittämään minulle. Kiitos uudelleen, ja aina. Rakastan sinua. Äiti: Sama koskee tietysti sinua. Se, miten rakastettu ja hyväksytty koen olevani, on suurin lahja minkä olen koskaan voinut sinulta ja isältä saada. Ilman siitä kumpuavaa luottamusta itseeni ja elämään mikään haaste ei onnistuisi – toivottavasti tämä saavutus on siitä todiste. Rakastan sinua.
Last, I thank my beloved men of which one is a bit hairier than the other. Thank you, Mosku, for keeping me active both mentally and physically. You have
definitely given me some grey hair, but it has been worth it (and I guess your fur has got a bit of grey as well). In contrast, you Esa, have the ability to blow away all my worries. You are the bedrock of my life. Thank you for everything,
rakastan sinua.
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