Biodiversity and Conservation

, Volume 23, Issue 13, pp 3287–3300 | Cite as

Isolation, patch size and matrix effects on bird assemblages in forest reserves

  • Mikko Mönkkönen
  • Ari Rajasärkkä
  • Petri Lampila
Original Paper


Habitat fragmentation is one of the most studied topics in ecology but our knowledge is still limited particularly concerning matrix effects on species distribution in a human-dominated landscape. We tested the ability of random sampling hypothesis, colonization–extinction dynamics and matrix-related concepts to explain the variation in species richness, total bird density and community composition in old-forest bird assemblages in two contrasting landscapes. We collected data on breeding bird abundances from 66 old-growth forest reserves in NE Finland and six larger areas in adjacent Russian Karelia using the line transect method. In Finland, protected old-forest patches are embedded in a matrix dominated by young regeneration stands. In Russia, study areas were situated in continuous, old forest dominated landscapes. Bird assemblages in old-forest patches embedded in human-modified matrix in Finland were not random samples from Russian bird assemblages. In the Finnish assemblages, species richness was lower and total bird density higher. Species richness declined with increasing distance (isolation) to Russia. Bird assemblages in large forest reserves in Finland close to Russia were structurally more similar to assemblages in the continuous reference landscape than those in small and more distant reserves. The results support the idea that several mechanisms related to colonisation–extinction dynamics and to matrix resource availability influence species distribution in fragmented landscapes but in species-specific ways. We conclude that even though small and isolated protected areas may foster high relative bird species density their ecological integrity is compromised, and therefore, improving matrix quality around reserves may lead to considerable conservation benefits.


Bird density Colonization–extinction dynamics Conservation planning Landscape neighborhood effects Species richness Theory of island biogeography 



We want to thank all those skilled ornithologists who have participated in line transect censuses over the years. We are grateful to the Interreg Karjala IIIa project, University of Oulu (PL) and Academy of Finland (to MM; project #138032) for funding. This paper was peer-reviewed in Peerage of Science, and we are grateful to Peer#673 for constructive comments. Moreover, we are grateful to two anonymous referees for their constructive criticism.

Supplementary material

10531_2014_780_MOESM1_ESM.pdf (44 kb)
Supplementary material 1 (PDF 43 kb)


  1. Andrén H (1994) Fragmentation on birds and mammals in landscapes with different proportions of suitable habitat: a review. Oikos 71:355–366CrossRefGoogle Scholar
  2. Angelstam P, Boutin S, Schmiegelow FKA, Villard M-A, Drapeau P, Host G, Innes J, Isachenko G, Kuuluvainen T, Mönkkönen M, Niemelä J, Niemi G, Roberge J-M, Spence J, Stone D (2004) Targets for boreal forest biodiversity conservation—a rationale for macroecological research and adaptive management. Ecol Bull 51:487–509Google Scholar
  3. Bray JR, Curtis JT (1957) An ordination of the upland forest communities of Southern Wisconsin. Ecol Monogr 27:325–349CrossRefGoogle Scholar
  4. Brook BW, Sodhi NS, Bradshaw CJA (2008) Synergies among extinction drivers under global change. Trends Ecol Evol 23:453–460PubMedCrossRefGoogle Scholar
  5. Brotons L, Mönkkönen M, Martin JL (2003a) Are fragments islands? Landscape context and density-area relationships in boreal forest birds. Am Nat 162:343–357PubMedCrossRefGoogle Scholar
  6. Brotons L, Mönkkönen M, Huhta E, Nikula A, Rajasärkkä A (2003b) Effects of landscape structure and forest reserve location on old growth forest bird distribution in northern Finnish forest reserves. Landsc Ecol 18:377–393CrossRefGoogle Scholar
  7. Clifford P, Richardson S, Hémon D (1988) Assessing the significance of correlation between two spatial processes. Biometrics 45:123–134CrossRefGoogle Scholar
  8. Connor EF, McCoy ED (1979) The statistics and biology of species-area relationship. Am Nat 113:791–833CrossRefGoogle Scholar
  9. Driscoll DA, Lindenmayer DB (2009) Empirical tests of metacommunity theory using an isolation gradient. Ecol Monogr 79:485–501CrossRefGoogle Scholar
  10. Driscoll DA, Banks SC, Barton PS, Lindenmayer DB, Smith AL (2013) Conceptual domain of the matrix in fragmented landscapes. Trends Ecol Evol 28:605–613PubMedCrossRefGoogle Scholar
  11. Dunning JB, Danielson BJ, Pulliam HR (1992) Ecological processes that affect populations in complex landscapes. Oikos 65:169–175CrossRefGoogle Scholar
  12. Edenius L, Elmberg J (1996) Landscape level effects of modern forestry on bird communities in north Swedish boreal forests. Landsc Ecol 11:325–338CrossRefGoogle Scholar
  13. Ehrlich PR, Pringle RM (2008) Where does biodiversity go from here? A grim business-as-usual forecast and a hopeful portfolio of partial solutions. Proc Natl Acad Sci USA 105:11579–11586PubMedCentralPubMedCrossRefGoogle Scholar
  14. Esseen P, Ehnström B, Ericson L, Sjöberg K (1997) Boreal forests. Ecol Bull 46:16–47Google Scholar
  15. Estades CF, Temple SA (1999) Deciduous-forest bird communities in a fragmented landscape dominated by exotic pine plantations. Ecol Appl 9:573–585CrossRefGoogle Scholar
  16. Evans KL, Greenwood JJD, Gaston KJ (2005) Dissecting the species–energy relationship. Proc R Soc B Biol Sci 272:2155–2163CrossRefGoogle Scholar
  17. Ewers RM, Didham RK (2006) Confounding factors in the detection of species responses to habitat fragmentation. Biol Rev 81:117–142PubMedCrossRefGoogle Scholar
  18. Fahrig L (2001) How much habitat is enough? Biol Cons 100:65–74CrossRefGoogle Scholar
  19. Finnish Forest Research Institute (2011) Finnish Statistical Yearbook of Forestry. Metla, Vantaa.
  20. Gustafson EJ, Gardner RH (1996) The effect of landscape heterogeneity on the probability of patch colonization. Ecology 77:94–107CrossRefGoogle Scholar
  21. Haila Y (1983) Land birds on northern islands: a sampling metaphor for insular colonization. Oikos 41:334–351CrossRefGoogle Scholar
  22. Hanski I, Gilpin M (1991) Metapopulation dynamics: brief history and conceptual domain. Biol J Linnean Soc 42:3–16CrossRefGoogle Scholar
  23. Honkanen M, Roberge JM, Rajasärkkä A, Mönkkönen M (2010) Disentangling the effects of area, energy and habitat heterogeneity on boreal forest bird species richness in protected areas. Glob Ecol Biogeogr 19:61–71CrossRefGoogle Scholar
  24. Järvinen O, Väisänen RA (1976) Finnish line transect censuses. Ornis Fenn 53:115–118Google Scholar
  25. Järvinen O, Väisänen RA (1983) Correction coefficients for line transect censuses of breeding birds. Ornis Fenn 60:97–104Google Scholar
  26. Krebs CJ (1999) Ecological methodology. Addison Weasley Longman, Menlo ParkGoogle Scholar
  27. Lampila P, Mönkkönen M, Desrochers A (2005) Demographic responses by birds to forest fragmentation. Conserv Biol 19:1537–1546CrossRefGoogle Scholar
  28. Lampila P, Mönkkönen M, Rajasärkkä A (2009) Ability of forest reserves to maintain original fauna—why the Chiffchaff (Phylloscopus collybita abietinus) has disappeared from eastern central Finland? Ornis Fenn 86:71–80Google Scholar
  29. Laurance W et al (2012) Averting biodiversity collapse in tropical forest protected areas. Nature 489:290–294PubMedCrossRefGoogle Scholar
  30. MacArthur RH, Wilson EO (1967) The Theory of Island Biogeography. Princeton University Press, PrincetonGoogle Scholar
  31. Marsden SJ, Whiffin M, Galetti M (2001) Bird diversity and abundance in forest fragments and Eucalyptus plantations around an Atlantic forest reserve, Brazil. Biodiv Cons 10:737–751CrossRefGoogle Scholar
  32. Martensen AC, Pimentel RG, Metzger JP (2008) Relative effects of fragment size and connectivity on bird community in the Atlantic rain forest: implications for conservation. Biol Cons 141:2184–2192CrossRefGoogle Scholar
  33. Moilanen A, Hanski I (1998) Metapopulation dynamics: effects of habitat quality and landscape structure. Ecology 79:2503–2515CrossRefGoogle Scholar
  34. Mönkkönen M, Reunanen P (1999) On critical thresholds in landscape connectivity: a management perspective. Oikos 84:302–306CrossRefGoogle Scholar
  35. Mönkkönen M, Welsh DA (1994) A biogeographical hypothesis on the effects of human caused habitat changes on the forest bird communities of Europe and North America. Ann Zool Fenn 31:61–70Google Scholar
  36. Potapov P, Turubanova S, Zhuravleva I, Hansen M, Yaroshenko A, Manisha A (2012) Forest cover change within the Russian European north after the breakdown of Soviet Union (1990–2005). Intl J For Res. doi: 10.1155/2012/729614 Google Scholar
  37. Prugh LR, Hodges KE, Sinclair ARE, Brashares JS (2008) Effect of habitat area and isolation on fragmented animal populations. Proc Natl Acad Sci USA 105:20770–20775PubMedCentralPubMedCrossRefGoogle Scholar
  38. Ricketts TH (2001) The matrix matters: effective isolation in fragmented landscapes. Am Nat 158:87–99PubMedCrossRefGoogle Scholar
  39. Sisk TD, Haddad NM, Ehrlich PR (1997) Bird assemblages in patchy woodlands: modeling the effects of edge and matrix habitats. Ecol Appl 7:1170–1180CrossRefGoogle Scholar
  40. Syrjänen K, Kalliola R, Puolasmaa A, Mattsson J (1994) Landscape structure and forest dynamics in subcontinental Russian European taiga. Ann Zool Fenn 31:19–34Google Scholar
  41. Trzcinski MK, Fahrig L, Merriam G (1999) Independent effects of forest cover and fragmentation on the distribution of forest breeding birds. Ecol Appl 9:586–593CrossRefGoogle Scholar
  42. Uotila A, Maltamo M, Uuttera J, Isomäki A (2001) Stand structure in seminatural and managed forests in eastern Fennoscandia and Russian Karelia. Ecol Bull 51:149–158Google Scholar
  43. Väisänen RA, Lammi E, Koskimies P (1998) Distribution, numbers and population changes in Finnish breeding birds. Otavan Kirjapaino, KeuruuGoogle Scholar
  44. Villard M-A, Trzcinski MK, Merriam G (1999) Fragmentation effects on forest birds: relative influence of woodland cover and configuration on landscape occupancy. Conserv Biol 13:774–781CrossRefGoogle Scholar
  45. Virkkala R, Rajasärkkä A (2006) Spatial variation of bird species in landscapes dominated by old-growth forests in northern boreal Finland. Biodivers Conserv 15:2143–2162CrossRefGoogle Scholar
  46. Virkkala R, Korhonen KT, Haapanen R, Aapala K (2000) Metsien ja soiden suojelutilanne metsä- ja suokasvillisuusvyöhykkeittäin valtakunnan metsien 8. inventoinnin perusteella. Suomen Ympäristö 395:1–49Google Scholar
  47. Watling JI, Nowakowski AJ, Donnelly MA, Orrock JL (2011) Meta-analysis reveals the importance of matrix composition for animals in fragmented habitat. Glob Ecol Biogeogr 20:209–217CrossRefGoogle Scholar
  48. Wilcove DS, Rothstein D, Dubow J, Phillips A, Losos E (1998) Quantifying threats to imperiled species in the United States. Bioscience 48:607–615CrossRefGoogle Scholar
  49. World Bank (2014) Terrestrial protected areas. [online]. Available from: Accessed 17 June 2014

Copyright information

© Springer Science+Business Media Dordrecht 2014

Authors and Affiliations

  • Mikko Mönkkönen
    • 1
  • Ari Rajasärkkä
    • 2
  • Petri Lampila
    • 3
    • 4
  1. 1.Department of Biological and Environmental SciencesUniversity of JyväskyläJyväskyläFinland
  2. 2.Metsähallitus, Natural Heritage ServicesOuluFinland
  3. 3.Department of BiologyUniversity of OuluOuluFinland
  4. 4.Metsähallitus, Liminka Bay Visitor CentreLiminkaFinland

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