Assessing how bird richness across ecological guilds of mountain-breeding birds responds to changes in forest attributes may inform the development of sustainable management strategies. In this study, we surveyed forest structure and bird communities across 148 sampling units in France and Italy to identify threshold values for stand variables that influence bird species richness. The sampling covered an altitudinal range from 805 to 1555 m a.s.l. and was mainly dominated by mountain beech and Alpine coniferous forests. We modelled species richness of overall bird communities and individual ecological guilds in response to structural and compositional attributes and topographic variables using generalized modelling. Bird species richness was negatively associated with stand-level total tree density. However, higher silver fir density was associated with greater species richness across most bird guilds, which preferred mixed woods and heterogeneous stand structure. An increase in deadwood volume had a significant positive effect on all guilds, confirming the importance of old-growth forest features for forest birds. Threshold values indicating a shift in species richness of bird guilds were identified only for some stand structural attributes (i.e., total basal area and deadwood volume). Our results suggest that the conservation of biodiversity in forest bird communities may benefit from management strategies that maintain mixed, relatively open canopies and promote deadwood retention to ensure the availability of associated habitats. Our approach lays the groundwork for further research, applying this methodology to additional variables and biodiversity metrics.
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Portaccio A, Paillet Y, Chojnacki L, Trentanovi G, Campagnaro T, Burrascano S, Sitzia T (2026). Bird response to forest structure and composition and implications for sustainable mountain forest management. iForest 19: 18-27. - doi: 10.3832/ifor4921-018
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Mirko Di Febbraro
Paper history
Received: Jun 06, 2025
Accepted: Dec 16, 2025
First online: Jan 11, 2026
Publication Date: Feb 28, 2026
Publication Time: 0.87 months
© SISEF - The Italian Society of Silviculture and Forest Ecology 2026
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(1)
Augusto L, Dupouey J-L, Ranger J (2003)Effects of tree species on understory vegetation and environmental conditions in temperate forests. Annals of Forest Science 60: 823-831.
CrossRef |
Gscholar
(2)
Ausden M (2004)Habitat management. In: “Bird Ecology and Conservation: A Handbook of Techniques” (Sutherland WJ, Newton I, Green RE eds). Oxford University Press, Oxford, UK, pp. 329-369.
Gscholar
(3)
Barbati A, Marchetti M, Chirici G, Corona P (2014)European forest types and Forest Europe SFM indicators: tools for monitoring progress on forest biodiversity conservation. Forest Ecology and Management 321: 145-157.
CrossRef |
Gscholar
(4)
Balakrishnan N, Colton T, Everitt B, Piegorsch W, Ruggeri F, Teugels J (2020)Generalized Poisson distributions. Wiley StatsRef: Statistics Reference Online, John Wiley and Sons, Ltd., website.
CrossRef |
Gscholar
(5)
Batáry P, Fronczek S, Normann C, Scherber C, Tscharntke T (2014)How do edge effect and tree species diversity change bird diversity and avian nest survival in Germany’s largest deciduous forest? Forest Ecology and Management 319: 44-50.
CrossRef |
Gscholar
(6)
Birčák T, Reif J (2015)The effects of tree age and tree species composition on bird species richness in a Central European montane forest. Biologia 70 (11): 1528-1536.
CrossRef |
Gscholar
(7)
Bledy M, Vacek S, Brabec P, Vacek Z, Cukor J, Aerny J, Sevčík R, Brynychová K (2024)Silver fir (
Abies alba Mill.): review of ecological insights, forest management strategies, and climate change’s impact on European forests. Forests 15 (6): 998.
CrossRef |
Gscholar
(8)
Bouvet A, Paillet Y, Archaux F, Tillon L, Denis P, Gilg O, Gosselin F (2016)Effects of forest structure, management and landscape on bird and bat communities. Environmental Conservation 43: 148-160.
CrossRef |
Gscholar
(9)
Braunisch V, Roder S, Coppes J, Froidevaux JS, Arlettaz R, Bollmann K (2019)Structural complexity in managed and strictly protected mountain forests: effects on the habitat suitability for indicator bird species. Forest Ecology and Management 448: 139-149.
CrossRef |
Gscholar
(10)
Brooks ME, Kristensen K, Van Benthem KJ, Magnusson A, Berg CW, Nielsen A, Skaug HJ, Maechler M, Bolker BM (2017)glmmTMB balances speed and flexibility among packages for zero-inflated generalized linear mixed modeling. The R Journal 9 (2): 378-400.
CrossRef |
Gscholar
(11)
Bull EL, Parks C G, Torgersen TR (1997)Trees and logs important to wildlife in the interior Columbia River basin. General Technical Report PNW-GTR-391, USDA Forest Service, Pacific NW Research Station, Portland, OR, USA, pp. 55.
Gscholar
(12)
Bunce RGH, Bogers MMB, Evans D, Halada L, Jongman RHG, Mücher CA, Bauch B, De Blust G, Parr TW, Olsvig-Whittaker L (2013)The significance of habitats as indicators of biodiversity and their links to species. Ecological Indicators 33: 19-25.
CrossRef |
Gscholar
(13)
Burrascano S, De Andrade RB, Paillet Y, Odor P, Antonini G, Bouget C, Campagnaro T, Gosselin F, Janssen P, Persiani AM, Nascimbene J, Sabatini FM, Sitzia T, Blasi C (2018)Congruence across taxa and spatial scales: are we asking too much of species data? Global Ecology and Biogeography 27: 980-990.
CrossRef |
Gscholar
(14)
Chamberlain DE, Negro M, Caprio E, Rolando A (2013)Assessing the sensitivity of alpine birds to potential future changes in habitat and climate to inform management strategies. Biological Conservation 167: 127-135.
CrossRef |
Gscholar
(15)
Corkery I, Irwin S, Quinn JL, Keating U, Lusby J, O’Halloran J (2020)Changes in forest cover result in a shift in bird community composition. Journal of Zoology 310 (4): 306-314.
CrossRef |
Gscholar
(16)
Del Castillo J, Pérez-Casany M (2005)Overdispersed and underdispersed Poisson generalizations. Journal of Statistical Planning and Inference 134 (2): 486-500.
CrossRef |
Gscholar
(17)
Díaz IA, Armesto JJ, Reid S, Sieving KE, Willson MF (2005)Linking forest structure and composition: avian diversity in successional forests of Chiloé Island, Chile. Biological Conservation 123 (1): 91-101.
CrossRef |
Gscholar
(18)
Drapeau P, Nappi A, Imbeau L, Saint-Germain M (2009)Standing deadwood for keystone bird species in the eastern boreal forest: managing for snag dynamics. The Forestry Chronicle 85: 227-234.
CrossRef |
Gscholar
(19)
Duflot R, Heinrichs S, Balducci L, Chianucci F, Hofmeister J, Paillet Y, Trentanovi G, Archaux F, Boch S, Bouget C, Dvolák D, Fischer M, Gosselin F, Gosselin M, Gossner MM, Holá E, Hošek J, Jung K, Palice Z, Renner SC, Weisser WW, Nagel TA, Burrascano S, Schall P (2025)Sustainable forest planning: Assessing biodiversity effects of Triad zoning based on empirical data and virtual landscapes. Proceedings of the National Academy of Sciences USA 122 (39): e2512683 122.
CrossRef |
Gscholar
(20)
Farwell LS, Wood PB, Dettmers R, Brittingham MC (2020)Threshold responses of songbirds to forest loss and fragmentation across the Marcellus-Utica shale gas region of central Appalachia, USA. Landscape Ecology 35: 1353-1370.
CrossRef |
Gscholar
(21)
Fauth JE, Bernardo J, Camara M, Resetarits Jr WJ, Van Buskirk J, McCollum SA (1996)Simplifying the jargon of community ecology: a conceptual approach. The American Naturalist 147 (2): 282-286.
CrossRef |
Gscholar
(22)
Fong Y, Huang Y, Gilbert PB, Permar SR (2017)chngpt: threshold regression model estimation and inference. BMC Bioinformatics 18 (1): 454.
CrossRef |
Gscholar
(23)
Forest Europe (2020)State of Europe’s Forests 2020 (Köhl M, Linser S, Prins K eds). Forest Europe Liaison Unit, Bratislava, Slovakia, pp. 394.
Gscholar
(24)
Fuller RJ (2003)Bird life of woodland and forest. Cambridge University Press, Cambridge, UK, pp. 244.
Gscholar
(25)
Fuller RJ, Robles H (2018)Conservation strategies and habitat management for European forest birds. In: “Ecology and Conservation of Forest Birds” (Mikusinski G, Roberge JM, Fuller RJ eds). Cambridge University Press, Cambridge, UK, pp. 455-507.
Gscholar
(26)
Geitzenauer M, Blondet M, De Koning J, Ferranti F, Sotirov M, Weiss G, Winkel G (2017)The challenge of financing the implementation of Natura 2000 - Empirical evidence from six European Union Member States. Forest Policy and Economics 82: 3-13.
CrossRef |
Gscholar
(27)
Gil-Tena A, Saura S, Brotons L (2007)Effects of forest composition and structure on bird species richness in a Mediterranean context: implications for forest ecosystem management. Forest Ecology and Management 242: 470-476.
CrossRef |
Gscholar
(28)
Gregory RD, Gibbons DW, Donald PF (2004)Bird census and survey techniques. In: “Bird Ecology and Conservation: A Handbook of Techniques” (Sutherland WJ, Newton I, Green RE eds). Oxford University Press, Oxford, UK, pp. 17-56.
Gscholar
(29)
Gregory RD, Van Strien A, Vorisek P, Gmelig Meyling AW, Noble DG, Foppen RP, Gibbons DW (2005)Developing indicators for European birds. Philosophical Transactions of the Royal Society B: Biological Sciences 360: 269-288.
CrossRef |
Gscholar
(30)
Gregory RD, Skorpilova J, Vorisek P, Butler S (2019)An analysis of trends, uncertainty and species selection shows contrasting trends of widespread forest and farmland birds in Europe. Ecological Indicators 103: 676-687.
CrossRef |
Gscholar
(31)
Hagan JM, Whitman AA (2006)Biodiversity indicators for sustainable forestry: simplifying complexity. Journal of Forestry 104 (4): 203-210.
CrossRef |
Gscholar
(32)
Hobson KA, Bayne E (2000)Breeding bird communities in boreal forest of Western Canada: consequences of unmixing the mixed woods. The Condor 102: 759-769.
CrossRef |
Gscholar
(33)
Kebrle D, Hofmeister J, Kodet V, Hošek J (2023)Key structural factors and their thresholds for promoting bird diversity in spruce-dominated production forests of central Europe. Forest Ecology and Management 550 (9): 121522.
CrossRef |
Gscholar
(34)
Kleiber C, Zeileis A (2020)Package aer - Applied econometrics with R. R package version 1(4), CRAN, pp. 206.
Online |
Gscholar
(35)
Korlan M, Holmes R, Recher H, Adamík P, Kropil R (2013)Convergence in foraging guild structure of forest breeding bird assemblages across three continents is related to habitat structure and foraging opportunities. Community Ecology 14 (1): 89-100.
CrossRef |
Gscholar
(36)
Larrieu L, Cabanettes A, Gonin P, Lachat T, Paillet Y, Winter S, Bouget C, Deconchat M (2014)Deadwood and tree microhabitat dynamics in unharvested temperate mountain mixed forests: a life-cycle approach to biodiversity monitoring. Forest Ecology and Management 334: 163-173.
CrossRef |
Gscholar
(37)
Lindenmayer DB, Claridge AW, Gilmore AM, Michael D, Lindenmayer BD (2002)The ecological roles of logs in Australian forests and the potential impacts of harvesting intensification on log-using biota. Pacific Conservation Biology 8 (2): 121-140.
CrossRef |
Gscholar
(38)
Mazerolle MJ (2019)AICcmodavg: model selection and multimodel inference based on (Q)AIC(c). R package version 2:2-1, website.
CrossRef |
Gscholar
(39)
McCain CM (2009)Global analysis of bird elevational diversity. Global Ecology and Biogeography 18 (3): 346-360.
CrossRef |
Gscholar
(40)
Mikusinski G, Villero D, Herrando S, Brotons L (2018)Macroecological patterns in forest bird diversity in Europe. In: “Ecology and Conservation of Forest Birds” (Mikusinski G, Roberge JM, Fuller RJ eds). Cambridge University Press, Cambridge, UK, pp. 137-182.
Gscholar
(41)
Mitchell MS, Rutzmoser SH, Wigley TB, Loehle C, Gerwin JA, Keyser PD, Lancia RA, Perry RW, Reynolds CJ, Thill RE, Weih R, White D, Wood PB (2006)Relationships between avian richness and landscape structure at multiple scales using multiple landscapes. Forest Ecology and Management 221 (1-3): 155-169.
CrossRef |
Gscholar
(42)
Mollet P, Bollmann K, Braunisch V, Alletaz R (2018)Subalpine coniferous forests of Europe. Avian communities in European high-altitude woodlands. In: “Ecology and Conservation of Forest Birds” (Mikusinski G, Roberge JM, Fuller RJ eds). Cambridge University Press, Cambridge, UK, pp. 231-252.
Gscholar
(43)
Müller J, Bütler R (2010)A review of habitat thresholds for dead wood: a baseline for management recommendations in European forests. European Journal of Forest Research 129: 981-992.
CrossRef |
Gscholar
(44)
Paillet Y, Pernot C, Boulanger V, Debaive N, Fuhr M, Gilg O, Gosselin F (2015)Quantifying the recovery of old-growth attributes in forest reserves: a first reference for France. Forest Ecology and Management 346: 51-64.
CrossRef |
Gscholar
(45)
Ram D, Axelsson AL, Green M, Smith HG, Lindström A (2017)What drives current population trends in forest birds - forest quantity, quality or climate? A large-scale analysis from northern Europe. Forest Ecology and Management 385: 177-188.
CrossRef |
Gscholar
(46)
R Core Team (2018)R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria.
Online |
Gscholar
(47)
Reif J, Skálová AJ, Vermouzek Z, Voríšek P (2022)Long-term trends in forest bird populations reflect management changes in Central European forests. Ecological Indicators 141: 109137.
CrossRef |
Gscholar
(48)
Rigal S, Dakos V, Alonso H, Aunis A, Benko Z, Brotons L, Chodkiewicz T, Chylarecki P, de Carli E, del Moral J C, Domsa C, Escandell V, Fontaine B, Foppen R, Gregory R, Harris S, Herrando S, Husby M, Ieronymidou C, Jiguet F, Kennedy J, Klvanová A, Kmecl P, Kuczynski L, Kurlavičius P, Kålås J A, Lehikoinen A, Lindström A, Lorrillière R, Moshoej C, Nellis R, Noble D, Eskildsen D P, Paquet J, Pélissié M, Pladevall C, Portolou D, Reif J, Schmid H, Seaman B, Szabo Z D, Szép T, Florenzano GT, Teufelbauer N, Trautmann S, van Turnhout C, Vermouzek Z, Vikstroem T, Vorísek I, Weiserbs A, Devictor V (2023)Farmland practices are driving bird population decline across Europe. Proceeding of the National Academy of Sciences USA 120 (21): e2216573120
CrossRef |
Gscholar
(49)
Roberge JM, Angelstam P (2006)Indicator species among resident forest birds: a cross-regional evaluation in northern Europe. Biological Conservation 130: 134-147.
CrossRef |
Gscholar
(50)
Roberge JM, Virkkala R, Mönkkönen M (2018)Boreal forest bird assemblages and their conservation. In: “Ecology and Conservation of Forest Birds” (Mikusinski G, Roberge JM, Fuller RJ eds). Cambridge University Press, Cambridge, UK, pp. 183-230.
Gscholar
(51)
Saetre P, Saetre LS, Brandtberg PO, Lundkvist H, Bengtsson J (1997)Ground vegetation composition and heterogeneity in pure Norway spruce and mixed Norway spruce-birch stands. Canadian Journal of Forest Research 27 (12): 2034-2042.
CrossRef |
Gscholar
(52)
Sánchez S, Cuervo JJ, Moreno E (2012)Vegetation structure in beech-fir forests: effects on the avian community. Revue d’Écologie (Terre et Vie) 67: 213-222.
CrossRef |
Gscholar
(53)
Seibold S, Richter T, Geres L, Seidl R, Martin R, Mitesser O, Senf C, Griem L, Müller J (2024)Soundscapes and airborne laser scanning identify vegetation density and its interaction with elevation as main driver of bird diversity and community composition. Diversity and Distributions 30 (12): 3905.
CrossRef |
Gscholar
(54)
Smyčková M, Koutecky T, Ujházyová M, Ujházy K, Verheyen K, Volarík D, Sebesta J, Friedl M, Malis F, Hofmeister J (2024)Herb layer species richness declines with heterogeneity of the forest structure in primary beech-dominated forests while proportion of forest specialists increases. Forest Ecology and Management 556: 121728.
CrossRef |
Gscholar
(55)
Uhl B, Krah FS, Baldrian P, Brandl R, Hagge J, Müller J, Thorn S, Vojtech T, Bässler C (2022)Snags, logs, stumps, and microclimate as tools optimizing deadwood enrichment for forest biodiversity. Biological Conservation 270 (5): 109569.
CrossRef |
Gscholar
(56)
Wesolowski T, Martin K (2018)Tree holes and hole-nesting birds in European and North-American forests. In: “Ecology and Conservation of Forest Birds” (Mikusinski G, Roberge JM, Fuller RJ eds). Cambridge University Press, Cambridge, UK, pp. 79-134.
Gscholar
(57)
Zuur AF, Ieno EN (2016)A protocol for conducting and presenting results of regression-type analyses. Methods in Ecology and Evolution 7: 636-645.
CrossRef |
Gscholar
(58)
Zuur AF, Ieno EN, Elphick CS (2009)A protocol for data exploration to avoid common statistical problems. Methods in Ecology and Evolution 1: 3-14.
CrossRef |
Gscholar