Several managed native forest stands have been reforested with conifer trees in Europe during recent centuries. These habitat alterations have influenced ground-dwelling invertebrates. We studied carabid beetle assemblages from a native beech forest (70-y-old), and a recently established (5-y-old), a young (15-y-old), a middle-aged (30-y-old) and a mature (50-y-old) Norway spruce plantation by pitfall trapping to explore the effect of reforestation on carabid beetles. The total number of carabid species, and the forest species were highest in the beech forest. The number of open-habitat species was highest in the youngest, relatively open monoculture. Ordination also confirmed changes in carabid composition with change in the studied habitats. Newly proposed forest affinity indices, based on species specificity, fidelity, and on a combination of specificity and fidelity were significantly higher in beech forest than in spruce plantations. We found these affinity indices especially useful in revealing the ecological character of the studied carabid assemblages. Regression analyses showed that leaf litter cover, herbs, shrubs, canopy closure and prey abundance were related to the structure of carabid-beetle assemblages.
Allegro, G. and R. Sciaky. 2002. Assessing the potential role of ground beetles (Coleoptera, Carabidae) as bioindicators in poplar stands, with a newly proposed ecological index (FAI). Forest Ecol. Manag. 175:275-284.
'Assessing the potential role of ground beetles (Coleoptera, Carabidae) as bioindicators in poplar stands, with a newly proposed ecological index (FAI) ' () 175 Forest Ecol. Manag. : 275 -284 .
Baars, M.A. and Th.S. Van Dijk. 1984. Population dynamics of two carabid beetles at a Dutch heathland. II. Egg production and survival in relation to density. J. Anim. Ecol. 53:389-400.
'Population dynamics of two carabid beetles at a Dutch heathland. II. Egg production and survival in relation to density ' () 53 J. Anim. Ecol. : 389 -400 .
Basu, A., I. R. Harris, and S. Basu. 1997. Minimum distance estimation: The approach using density-based distances. In: G. S. Maddala and C. R. Rao, (eds), Handbook of Statistics, volume 15, pp. 21-48. North-Holland.
Minimum distance estimation: The approach using density-based distances , () 21 -48 .
Baguette, M. and S. Gérard. 1993. Effects of spruce plantations on carabid beetles in southern Belgium. Pedobiologia 37:129-140.
'Effects of spruce plantations on carabid beetles in southern Belgium ' () 37 Pedobiologia : 129 -140 .
Bonham, K.J., R. Mesibov and R. Bashford. 2002. Diversity and abundance of some ground-dwelling invertebrates in plantation vs. native forests in Tasmania, Australia. Forest Ecol. Manag. 158:237-247.
'Diversity and abundance of some ground-dwelling invertebrates in plantation vs. native forests in Tasmania, Australia ' () 158 Forest Ecol. Manag. : 237 -247 .
Brunsting, M.H. and H.J.L. Heessen. 1984. Density regulation in the carabid beetle Pterostichus oblongopunctatus. J. Anim. Ecol. 53:751-760.
'Density regulation in the carabid beetle ' () 53 Pterostichus oblongopunctatus. J. Anim. Ecol. : 751 -760 .
Bryan, K.M. and S.D. Wratten. 1984. The responses of polyphagous predators to spatial heterogeneity: aggregation by carabid and staphylinid beetles to their cereal aphid prey. Ecol. Entomol. 9:251-259.
'The responses of polyphagous predators to spatial heterogeneity: aggregation by carabid and staphylinid beetles to their cereal aphid prey ' () 9 Ecol. Entomol. : 251 -259 .
Butterfield, J. 1997. Carabid community succession during forestry cycle in conifer plantations. Ecography 20:614-625.
'Carabid community succession during forestry cycle in conifer plantations ' () 20 Ecography : 614 -625 .
Butterfield, J., M.L. Luff, M. Baines and M.D. Eyre. 1995. Carabid beetle communities as indicators of conservation potential in upland forests. Forest Ecol. Manag. 79:63-77.
'Carabid beetle communities as indicators of conservation potential in upland forests ' () 79 Forest Ecol. Manag. : 63 -77 .
Desender, K., A. Ervynck and G. Tack. 1999. Beetle diversity and historical ecology of woodlands in Flanders. Belg. J. Zool. 129:139-156.
'Beetle diversity and historical ecology of woodlands in Flanders ' () 129 Belg. J. Zool. : 139 -156 .
Digweed, S.C., C.R. Currie, H.A. Cárcamo and J.R. Spence. 1995. Digging out the digging - in effect of pitfall traps: influences of depletion and disturbance on catches of ground beetles (Coleoptera: Carabidae). Pedobiologia 39, 561-576.
'Digging out the digging - in effect of pitfall traps: influences of depletion and disturbance on catches of ground beetles (Coleoptera: Carabidae) ' () 39 Pedobiologia : 561 -576 .
Dufrěne, M. and P. Legendre. 1997. Species assemblages and indicator species: the need for a flexible asymmetrical approach. Ecol. Monogr. 67:345-366.
'Species assemblages and indicator species: the need for a flexible asymmetrical approach ' () 67 Ecol. Monogr. : 345 -366 .
Elek, Z., T. Magura and B. Tóthmérész. 2005. Effect of canopy closure of a young Norway spruce plantation on ground beetles. In: G.L. Lövei and S. Toft (eds), European Carabidology 2003. Proceedings of the 11th European Carabidologists' Meeting. DIAS Report, No. 114. Flakkebjerg, pp. 89-97.
, , .
Fahy, O. and M. Gormally. 1998. A comparison of plant and carabid beetle communities in an Irish oak woodland with a nearby conifer plantation and clearfelled site. Forest Ecol. Manag. 110:263-273.
'A comparison of plant and carabid beetle communities in an Irish oak woodland with a nearby conifer plantation and clearfelled site ' () 110 Forest Ecol. Manag. : 263 -273 .
Grüm, L. 1971. Spatial differentiation of the Carabus L. (Carabidae, Coleoptera) mobility. Ekologika Polska 19:1-34.
'Spatial differentiation of the Carabus L. (Carabidae, Coleoptera) mobility ' () 19 Ekologika Polska : 1 -34 .
Guillemain, M., M. Loreau and T. Daufresne. 1997. Relationships between the regional distribution of carabid beetles (Coleoptera, Carabidae) and the abundance of their potential prey. Acta Oecol. 18:465-483.
'Relationships between the regional distribution of carabid beetles (Coleoptera, Carabidae) and the abundance of their potential prey ' () 18 Acta Oecol. : 465 -483 .
Humphrey, J.W., C. Hawes, A.J. Peace, R. Ferris-Kaan and M.R. Jukes. 1999. Relationship between insect diversity and habitat characteristics in plantation forests. Forest Ecol. Manag. 113:11-21.
'Relationship between insect diversity and habitat characteristics in plantation forests ' () 113 Forest Ecol. Manag. : 11 -21 .
Hurka, K. 1996. Carabidae of the Czech and Slovak Republics. Kabourek, Zlin.
Ings, T.C. and S.E. Hartley. 1999. The effect of habitat structure on carabid communities during the regeneration of a native Scottish forest. Forest Ecol. Manag. 119:123-136.
'The effect of habitat structure on carabid communities during the regeneration of a native Scottish forest ' () 119 Forest Ecol. Manag. : 123 -136 .
Jukes, M.R., A.J. Peace and R. Ferris. 2001. Carabid beetle communities associated with coniferous plantations in Britain: the influence of site, ground vegetation and stand structure. Forest Ecol. Manag. 148:271-286.
'Carabid beetle communities associated with coniferous plantations in Britain: the influence of site, ground vegetation and stand structure ' () 148 Forest Ecol. Manag. : 271 -286 .
Koivula, M. 2002. Boreal carabid-beetle (Coleoptera, Carabidae) assemblages in thinned uneven-aged and clear-cut spruce stands. Ann. Zool. Fenn. 39:131-149.
'Boreal carabid-beetle (Coleoptera, Carabidae) assemblages in thinned uneven-aged and clear-cut spruce stands ' () 39 Ann. Zool. Fenn. : 131 -149 .
Koivula, M., J. Kukkonen and J. Niemelä. 2002. Boreal carabid-beetle (Coleoptera, Carabidae) assemblages along the clear-cut originated succession gradient. Biodivers. Conserv. 11:1269-1288.
'Boreal carabid-beetle (Coleoptera, Carabidae) assemblages along the clear-cut originated succession gradient ' () 11 Biodivers. Conserv. : 1269 -1288 .
Koivula, M. and J. Niemelä. 2003. Gap felling as a forest harvesting method in boreal forests: responses of carabid beetles (Coleoptera, Carabidae). Ecography 26:179-187.
'Gap felling as a forest harvesting method in boreal forests: responses of carabid beetles (Coleoptera, Carabidae) ' () 26 Ecography : 179 -187 .
Kotze, D. J. and M.J. Samways. 1999. Invertebrate conservation at the interface between the grassland matrix and natural Afromontane forest fragments. Biodivers. Conserv. 8:1339-1363.
'Invertebrate conservation at the interface between the grassland matrix and natural Afromontane forest fragments ' () 8 Biodivers. Conserv. : 1339 -1363 .
Legendre, P. and L. Legendre. 1998. Numerical Ecology. Elsevier, Amsterdam.
Numerical Ecology , ().
Loreau, M. 1988. Determinants of the seasonal pattern in the niche structure of a forest carabid community. Pedobiologia 31:75-87.
'Determinants of the seasonal pattern in the niche structure of a forest carabid community ' () 31 Pedobiologia : 75 -87 .
Lövei, G. L. 2005. Generalised entropy indices have a long history in ecology - a comment. Communty Ecology 6:245-247.
'Generalised entropy indices have a long history in ecology - a comment ' () 6 Communty Ecology : 245 -247 .
Lövei, G. and K.D. Sunderland. 1996. Ecology and behavior of ground beetles (Coleoptera: Carabidae). Annu. Rev. Entomol. 41:231-256.
'Ecology and behavior of ground beetles (Coleoptera: Carabidae) ' () 41 Annu. Rev. Entomol. : 231 -256 .
Luff, M.L. 1975. Some features influencing the efficiency of pitfall traps. Oecologia 19:345-357.
'Some features influencing the efficiency of pitfall traps ' () 19 Oecologia : 345 -357 .
Mader, H.J. 1984. Animal habitat isolation by roads and agricultural fields. Biol. Conserv. 29:81-96.
'Animal habitat isolation by roads and agricultural fields ' () 29 Biol. Conserv. : 81 -96 .
Magura, T. 2002. Carabids and forest edge: spatial pattern and edge effect. Forest Ecol. Manag. 157:23-37.
'Carabids and forest edge: spatial pattern and edge effect ' () 157 Forest Ecol. Manag. : 23 -37 .
Magura, T., V. Ködöböcz and Zs. Bokor. 2001a. Effects of forestry practices on carabids (Coleoptera: Carabidae) - Implication for nature management. Acta Phytopathol. Entomol. Hung. 36:179-188.
'.Effects of forestry practices on carabids (Coleoptera: Carabidae) - Implication for nature management ' () 36 Acta Phytopathol. Entomol. Hung. : 179 -188 .
Magura, T., B. Tóthmérész and Zs. Bordán. 1997. Comparison of the carabid communities of a zonal oak-hornbeam forest and pine plantations. Acta Zool. Acad. Sci. Hung. 43:173-182.
'Comparison of the carabid communities of a zonal oak-hornbeam forest and pine plantations ' () 43 Acta Zool. Acad. Sci. Hung. : 173 -182 .
Magura, T., B. Tóthmérész and Zs. Bordán. 2000a. Effects of nature management practice on carabid assemblages (Coleoptera: Carabidae) in a non-native plantation. Biol. Conserv. 93:95-102.
'.Effects of nature management practice on carabid assemblages (Coleoptera: Carabidae) in a non-native plantation ' () 93 Biol. Conserv. : 95 -102 .
Magura, T., B. Tóthmérész and Z. Elek. 2002. Impacts of non-native spruce reforestation on ground beetles. Eur. J. Soil Biol. 38:291-295.
'Impacts of non-native spruce reforestation on ground beetles ' () 38 Eur. J. Soil Biol. : 291 -295 .
Magura, T., B. Tóthmérész and Z. Elek. 2003. Diversity and composition of carabids during a forestry cycle. Biodiv. Conserv. 12:73-85.
'Diversity and composition of carabids during a forestry cycle ' () 12 Biodiv. Conserv. : 73 -85 .
Magura, T., B. Tóthmérész and Z. Elek. 2005. Impacts of leaf-litter addition on carabids in a conifer plantation. Biodiv. Conserv. 14:475-491.
'Impacts of leaf-litter addition on carabids in a conifer plantation ' () 14 Biodiv. Conserv. : 475 -491 .
Magura, T., B. Tóthmérész and T. Molnár. 2000b. Spatial distribution of carabids along grass-forest transects. Acta Zool. Acad. Sci. Hung. 46:1-17.
'.Spatial distribution of carabids along grass-forest transects ' () 46 Acta Zool. Acad. Sci. Hung. : 1 -17 .
Magura, T., B. Tóthmérész and T. Molnár. 2001b. Forest edge and diversity: carabids along forest-grassland transects. Biodiv. Conserv. 10:287-300.
'.Forest edge and diversity: carabids along forest-grassland transects ' () 10 Biodiv. Conserv. : 287 -300 .
Mátyás, Cs. 1996. Forestry Ecology. Mezőgazda Kiadó, Budapest. (in Hungarian)
Moffatt, C., S. McNeill and A. J. Morton. 2005. The quantification of habitat architecture for explanations of arthropod assemblage patterns: a comparison of two methods. Community Ecology 5:83-92.
'The quantification of habitat architecture for explanations of arthropod assemblage patterns: a comparison of two methods ' () 5 Community Ecology : 83 -92 .
Molnár, T., T. Magura and B. Tóthmérész. 2001. Ground beetles (Carabidae) and edge effect in oak-hornbeam forest and grassland transects. Eur. J. Soil Biol. 37:297-300.
'Ground beetles (Carabidae) and edge effect in oak-hornbeam forest and grassland transects ' () 37 Eur. J. Soil Biol. : 297 -300 .
Niemelä, J. 1997. Invertebrates and boreal forest management. Conserv. Biol. 11:601-610.
'Invertebrates and boreal forest management ' () 11 Conserv. Biol. : 601 -610 .
Niemelä, J. 1999. Management in relation to disturbance in the boreal forest. Forest Ecol. Manag. 115:127-134.
'Management in relation to disturbance in the boreal forest ' () 115 Forest Ecol. Manag. : 127 -134 .
Niemelä, J., Y. Haila, E. Halme, T. Pajunen and P. Punttila. 1992. Small-scale heterogeneity in the spatial distribution of carabid beetles in the southern Finnish taiga. J. Biogeogr. 19:173-181.
'Small-scale heterogeneity in the spatial distribution of carabid beetles in the southern Finnish taiga ' () 19 J. Biogeogr. : 173 -181 .
Niemelä, J., Y. Haila and P. Punttila. 1996. The importance of small-scale heterogeneity in boreal forests: variation in diversity in forest-floor invertebrates across the succession gradient. Ecography 19:352-368.
'The importance of small-scale heterogeneity in boreal forests: variation in diversity in forest-floor invertebrates across the succession gradient ' () 19 Ecography : 352 -368 .
Niemelä, J., D. Langor and J.R. Spence. 1993. Effects of clear-cut harvesting on boreal ground-beetle assemblages (Coleoptera: Carabidae) in western Canada. Conserv. Biol. 7:551-561.
'Effects of clear-cut harvesting on boreal ground-beetle assemblages (Coleoptera: Carabidae) in western Canada ' () 7 Conserv. Biol. : 551 -561 .
Niemelä, J. and J.R. Spence. 1994. Distribution of forest dwelling carabids (Coleoptera): spatial scale and the concept of communities. Ecography 17:166-175.
'Distribution of forest dwelling carabids (Coleoptera): spatial scale and the concept of communities ' () 17 Ecography : 166 -175 .
Ohlemüller, R., P. Bannister, K. J. M. Dickinson, S. Walker, B. J. Anderson and J. B. Wilson 2004. Correlates of vascular plant species richness in fragmented indigenous forests: assessing the role of local and regional factors. Community Ecology 5:45-54.
'Correlates of vascular plant species richness in fragmented indigenous forests: assessing the role of local and regional factors ' () 5 Community Ecology : 45 -54 .
Ricotta C. 2005. On parametric diversity indices in ecology: A historical note. Communty Ecology 6:241-244.
'On parametric diversity indices in ecology: A historical note ' () 6 Communty Ecology : 241 -244 .
Rutherford, A. 2001. Introducing ANOVA and ANCOVA. Sage Publications, London.
Introducing ANOVA and ANCOVA , ().
Sergeeva, T.K. 1994. Seasonal dynamics of interspecific trophic relations in a carabid beetle assemblage. In: K. Desender et al. (eds), Carabid beetles: ecology and evolution. Kluwer Academic Publishers, Dordrecht, pp. 367-370.
Seasonal dynamics of interspecific trophic relations in a carabid beetle assemblage , () 367 -370 .
Sokal, R.R. and F.J. Rohlf. 1981. Biometry. W. H. Freeman, New York.
Biometry , ().
Spence J.R. and J. Niemelä. 1994. Sampling carabid assemblages with pitfall traps: the madness and the method. Can. Entomol. 126:881-894.
'Sampling carabid assemblages with pitfall traps: the madness and the method ' () 126 Can. Entomol. : 881 -894 .
Sustek, Z. 1981. Influence of clear cutting on ground beetles (Coleoptera, Carabidae) in a pine forest. Comm. Ins. Forestalis 12:243-254.
'Influence of clear cutting on ground beetles (Coleoptera, Carabidae) in a pine forest ' () 12 Comm. Ins. Forestalis : 243 -254 .
Szyszko, J. 1987. How can the fauna of Carabidae be protected in managed pine forest? Acta Phytopathol. Entomol. Hung. 22:293-303.
'How can the fauna of Carabidae be protected in managed pine forest? ' () 22 Acta Phytopathol. Entomol. Hung. : 293 -303 .
Thiele, H.U. 1977. Carabid Beetles in their Environments. Springer Verlag, Berlin.
Carabid Beetles in their Environments , ().
Werner, S.M. and K.F. Raffa. 2000. Effects of forest management practices on the diversity of ground-occurring beetles in mixed northern hardwood forests of the Great Lakes Region. Forest Ecol. Manag. 139:135-155.
'Effects of forest management practices on the diversity of ground-occurring beetles in mixed northern hardwood forests of the Great Lakes Region ' () 139 Forest Ecol. Manag. : 135 -155 .