European Journal of Forest Research

, Volume 135, Issue 3, pp 539–549 | Cite as

Human-sensitive bryophytes retreat into the depth of forest fragments in central European landscape

  • Jeňýk HofmeisterEmail author
  • Jan Hošek
  • Marek Brabec
  • Aleš Tenčík
Original Paper


An importance of the edge effect and other environmental factors on bryophyte communities has been repeatedly evaluated in boreal forests whereas only rarely in forest remnants in traditionally inhabited landscape of central Europe. Adopting the design of a previous study dealing with vascular plants, we examined whether species richness and composition of bryophyte communities in 23 forest fragments (0.1–255 ha) in a representative upland agricultural landscape in central Bohemia coincides with the same environmental factors as vascular plants and if so, whether congruently or not. According to generalized additive models, bryophyte species richness and composition were significantly related to forest vegetation type, stand basal area and soil pH, analogous to vascular plants. Distribution of human-sensitive bryophyte species was further associated with slope aspect and an unlimited long-range edge effect, shown by increased occurrence of human-sensitive bryophytes along the entire distance gradient from the forest edge (0–477 m). Communities of bryophytes and vascular plants were highly congruent in species richness, species turnover and even in number of species with similar colonization ability. We conclude that requirements of human-sensitive bryophytes are fulfilled only in the core areas of large forest fragments. The small size of most current forest fragments and the range of the edge effect suggest that spatial patterns of bryophyte species composition within forest fragments have been substantially altered in the entire region of central Europe.


Colonization Forest continuity Fragmentation Forest management Fragment size 



We would like to thank P. Myšina and J. Hošková for collecting soil samples and other field environmental data. We are grateful to E. Holá and J. Kučera for help with determination of some bryophytes, Z. Drhovská for rectification of cadastral maps, and to T. Svoboda, R. Filippi, and V. Treml for processing of synthetic maps and related datasets. This research was funded by the Ministry of Environment of the Czech Republic (Grant Projects VaV SM 6/69/05 and SP 2d3/139/07).

Supplementary material

10342_2016_953_MOESM1_ESM.docx (62 kb)
Supplementary material 1 (DOCX 61 kb)


  1. Alignier A, Deconchat M (2011) Variability of forest edge effect on vegetation implies reconsideration of its assumed hypothetical pattern. Appl Veg Sci 14:67–74CrossRefGoogle Scholar
  2. Bartels SF, Chen HYH (2013) Interactions between overstorey and understorey vegetation along an overstorey compositional gradient. J Veg Sci 24:543–552CrossRefGoogle Scholar
  3. Bergamini A, Pauli D, Peintinger M, Schmid B (2001) Relationships between productivity, number of shoots and number of species in bryophytes and vascular plants. J Ecol 89:920–929CrossRefGoogle Scholar
  4. Cadenasso ML, Pickett STA (2001) Effect of edge structure on the flux of species into forest interiors. Conserv Biol 15:91–97CrossRefGoogle Scholar
  5. Cadenasso ML, Traynor MM, Pickett STA (1997) Functional location of forest edges: gradients of multiple physical factors. Can J For Res 27:774–782CrossRefGoogle Scholar
  6. Cleavitt NL (2002) Stress tolerance of rare and common moss species in relation to their occupied environments and asexual dispersal potential. J Ecol 90:785–795CrossRefGoogle Scholar
  7. Danihelka J, Chrtek J Jr, Kaplan Z (2012) Checklist of vascular plants of the Czech Republic. Preslia 84:647–811Google Scholar
  8. De Cáceres M, Jansen F (2015) Relationship between species and groups of sites. R-package ‘indicspecies’, version 1.7.4Google Scholar
  9. De Cáceres M, Legendre P (2009) Associations between species and groups of sites: indices and statistical inference. Ecology 90:3566–3574CrossRefPubMedGoogle Scholar
  10. De Frenne P, Baeten L, Graae BJ, Brunet J, Wulf M, Orczewska A, Kolb A, Jansen I, Jamoneau A, Jacquemyn H, Hermy M, Diekmann M, De Schrijver A, De Sanctis M, Decocq G, Cousins SAO, Verheyen K (2011) Interregional variation in the floristic recovery of post-agricultural forests. J Ecol 99:600–609Google Scholar
  11. de Mendiburu F (2014) Statistical procedures for agricultural research. R-package ‘agricolae’, version 1.1-7Google Scholar
  12. Dierßen K (2001) Distribution, ecological amplitude and phytosociological characterization of european bryophytes, 1st edn. Cramer in der Gebrüder Borntraeger Verlagsbuchhandlung, BerlinGoogle Scholar
  13. Graae BJ, Sunde PB (2000) The impact of forest continuity and management on forest floor vegetation evaluated by species traits. Ecography 23:720–731CrossRefGoogle Scholar
  14. Halpern CB, Dovčiak M, Urgenson LS, Evans SA (2014) Substrates mediate responses of forest bryophytes to a gradient in overstory retention. Can J For Res 44:855–866CrossRefGoogle Scholar
  15. Harper KA, MacDonald SE, Burton PJ, Chen J, Brosofske KD, Saunders SC, Euskirchen ES, Roberts D, Jaiteh MS, Esseen PA (2005) Edge influence on forest structure and composition in fragmented landscapes. Conserv Biol 19:768–782CrossRefGoogle Scholar
  16. Harrell FE (2014) Hmisc. R-package, version 3.14-4Google Scholar
  17. Harrison S, Bruna E (1999) Habitat fragmentation and large-scale conservation: What do we know for sure? Ecography 22:225–232CrossRefGoogle Scholar
  18. Hastie T, Tibshirani R (1990) Generalized additive models. Chapman and Hall, New YorkGoogle Scholar
  19. Hermy M, Honnay O, Firbank L, Grashof-Bokdam C, Lawesson JE (1999) An ecological comparison between ancient and other forest plant species of Europe, and the implications for forest conservation. Biol Conserv 91:9–22CrossRefGoogle Scholar
  20. Hochberg Y (1988) A sharper Bonferroni procedure for multiple tests of significance. Biometrika 75:800–803CrossRefGoogle Scholar
  21. Hofmeister J, Hošek J, Brabec M, Hédl R, Modrý M (2013) Strong influence of long-distance edge effect on herb-layer vegetation in forest fragments in an agricultural landscape. Perspect Plant Ecol 1:293–303CrossRefGoogle Scholar
  22. Hutsemekers V, Dopagne C, Vanderpoorten A (2008) How far and how fast do bryophytes travel at the landscape scale? Divers Distrib 14:483–492CrossRefGoogle Scholar
  23. Hylander K (2005) Aspect modifies the magnitude of edge effects on bryophyte growth in boreal forests. J Appl Ecol 42:518–525CrossRefGoogle Scholar
  24. Ingerpuu N, Vellak K, Kukk T, Pärtel M (2001) Bryophyte and vascular plant species richness in boreo-nemoral moist forest and mires. Biodivers Conserv 10:2153–2166CrossRefGoogle Scholar
  25. Kučera J, Váňa J, Hradílek Z (2012) Bryophyte flora of the Czech Republic: updated checklist and Red List and a brief analysis. Preslia 84:813–850Google Scholar
  26. Lindenmayer DB, Fischer J (2006) Habitat fragmentation and landscape change. An ecological and conservation synthesis, 1st edn. Island Press, WashingtonGoogle Scholar
  27. Löbel S, Rydin H (2009) Dispersal and life-history strategies in epiphyte metacommunities: alternative solutions to survival in patchy dynamics landscapes. Oecologia 161:569–579CrossRefPubMedGoogle Scholar
  28. Löbel S, Rydin H (2010) Trade-offs and habitat constraints in the establishment of epiphytic bryophytes. Func Ecol 24:887–897Google Scholar
  29. Löbel S, Snäll T, Rydin H (2006) Metapopulation processes in epiphytes inferred from patterns of regional distribution and local abundance in fragmented forest landscapes. J Ecol 94:856–868CrossRefGoogle Scholar
  30. Löbel S, Snäll T, Rydin H (2012) Epiphytic bryophytes near forest edges and on retention trees: reduced growth and reproduction especially in old-growth-forest indicator species. J Appl Ecol 49:1334–1343CrossRefGoogle Scholar
  31. Łuczaj L, Sadowska B (1997) Edge effect in different groups of organisms: vascular plant, bryophyte and fungi species richness across a forest-grassland border. Folia Geobot 32:343–353CrossRefGoogle Scholar
  32. Márialigeti S, Németh B, Tinya F, Ódor P (2009) The effects of stand structure on ground-floor bryophyte assemblages in temperate mixed forests. Biodivers Conserv 18:2223–2241CrossRefGoogle Scholar
  33. Mežaka A, Brūmelis G, Piterāns A (2012) Tree and stand-scale factors affecting richness and composition of epiphytic bryophytes and lichens in deciduous woodland key habitats. Biodivers Conserv 21:3221–3241CrossRefGoogle Scholar
  34. Miller NG, McDaniel SF (2004) Bryophyte dispersal inferred from colonization of an introduced substratum on Whiteface Mountain, New York. Am J Bot 91:1173–1182CrossRefPubMedGoogle Scholar
  35. Murcia C (1995) Edge effects in fragmented forests: implications for conservation. Trends Ecol Evol 10:58–62CrossRefPubMedGoogle Scholar
  36. Ódor P, Heilmann-Clausen J, Christensen M, Aude E, van Dort KW, Piltaver A, Siller I, Veerkamp MT, Walleyn R, Standovár T, van Hees AFM, Kosec J, Matocec N, Kraigher H, Grebenc T (2006) Diversity of dead wood inhabiting fungi and bryophytes in semi-natural beech forests in Europe. Biol Conserv 131:58–71CrossRefGoogle Scholar
  37. Oksanen J, Blanchet FG, Kindt R, Legendre P, Minchin PR, O’Hara RB, Simpson GL, Solymos P, Stevens MHH, Wagner H (2012) Vegan: community ecology package. R-package, version 2.0-5Google Scholar
  38. Pärtel M, Helm A, Ingerpuu N, Reier Ü, Tuvi EL (2004) Conservation of Northern European plant diversity: the correspondence with soil pH. Biol Conserv 120:525–531CrossRefGoogle Scholar
  39. Pellissier V, Bergès L, Nadeltcheva T, Schmitt MC, Avon C, Cluzeau C, Dupouey JL (2013) Understorey plant species show long-range spatial patterns in forest patches according to distance-to-edge. J Veg Sci 24:9–24CrossRefGoogle Scholar
  40. Perhans K, Appelgren L, Jonsson F, Nordin U, Söderström B, Gustafsson L (2009) Retention patches as potential refugia for bryophytes and lichens in managed forest landscapes. Biol Conserv 142:1125–1133CrossRefGoogle Scholar
  41. Pharo EJ, Zartman CE (2007) Bryophytes in a changing landscape: the hierarchical effects of habitat fragmentation on ecological and evolutionary processes. Biol Conserv 135:315–325CrossRefGoogle Scholar
  42. R Development Core Team (2012) R: a language and environment for statistical computing. R foundation for statistical computing. Vienna, Austria.
  43. Ries L, Fletcher RJ Jr, Battin J, Sisk TD (2004) Ecological responses to habitat edges: mechanisms, models, and variability explained. Ann Rev Ecol Evol Syst 35:491–522CrossRefGoogle Scholar
  44. Roberge JM, Bengtsson SBK, Wulff S, Snäll T (2011) Edge creation and tree dieback influence the patch-tracking metapopulation dynamics of a red-listed epiphytic bryophyte. J Appl Ecol 48:650–658CrossRefGoogle Scholar
  45. Snäll T, Ehrlén J, Rydin H (2005) Colonization–extinction dynamics of an epiphyte metapopulation in a dynamic landscape. Ecology 86:106–115CrossRefGoogle Scholar
  46. Stewart KJ, Mallik AU (2006) Bryophyte responses to microclimatic edge effects across riparian buffers. Ecol Appl 16:1474–1486CrossRefPubMedGoogle Scholar
  47. Vanderpoorten A, Engels P, Sotiaux A (2004) Trends in diversity and abundance of obligate epiphytic bryophytes in a highly managed landscape. Ecography 27:567–576CrossRefGoogle Scholar
  48. Verheyen K, Honnay O, Motzkin G, Hermy M, Foster DR (2003) Response of forest plant species to land-use change: a life-history trait-based approach. J Ecol 91:563–577CrossRefGoogle Scholar
  49. Vockenhuber EA, Scherber C, Langenbruch C, Meißner M, Seidel D, Tscharntke T (2011) Tree diversity and environmental context predict herb species richness and cover in Germany’s largest connected deciduous forest. Perspect Plant Ecol 13:111–119CrossRefGoogle Scholar
  50. Westphal C, Härdtle W, von Oheimb G (2004) Forest history, continuity and dynamics naturalness. In: Honnay O, Verheyen K, Bossuyt B, Hermy M (eds) Forest biodiversity: lesson from history for conservation, IUFRO research series, vol 10. CAB International, Wallingford, pp 205–220Google Scholar
  51. Wood SN (2006) Generalized additive models. An introduction with R. Chapman and Hall, Boca RatonGoogle Scholar

Copyright information

© Springer-Verlag Berlin Heidelberg 2016

Authors and Affiliations

  • Jeňýk Hofmeister
    • 1
    • 2
    Email author
  • Jan Hošek
    • 1
  • Marek Brabec
    • 3
  • Aleš Tenčík
    • 1
  1. 1.Ecological Services, Areál ČOVHořoviceCzech Republic
  2. 2.Department of Biogeochemical and Hydrological Cycles, Global Change Research InstituteCzech Academy of SciencesČeské BudějoviceCzech Republic
  3. 3.Department of Nonlinear Modeling, Institute of Computer ScienceCzech Academy of SciencesPragueCzech Republic

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