Abstract
We studied the effects of fragmentation (edge effects and patch size) and trampling (path cover) on carabid beetle assemblages in urban woodland patches in Helsinki, Finland. We expected that (1) open habitat and generalist species would benefit and forest species would suffer from increased woodland fragmentation, and (2) most carabid species would respond negatively to increased levels of trampling. A total of 2088 carabid individuals representing 37 species were collected. A cluster analysis distinguished sites in the interior of large woodland patches, with low or moderate path cover, from the other sites. The other sites did not cluster meaningfully, suggesting increased variation in the carabid fauna with increasing human impact. All species and ecological species-groups decreased with increasing distance from the edge toward the woodland interior and with increasing patch size. This pattern is in accordance with our expectation for open habitat and generalist species but opposite to what we expected for forest species. The reason for these surprising results may be that (1) the species we collected are not true forest interior species, (2) urban woodland edges are optimal habitats for many forest carabids, or (3) edges are actually sub-optimal, and high catches simply reflect increased activity of beetles moving away from the edge. Trampling did not have an overall negative effect on carabids as hypothesized. Species associated with moist forest habitat responded as predicted: they decreased in abundance with increasing path cover. Furthermore, open habitat species decreased with increasing path cover but more straightforward than we had predicted. Model elaboration, by dropping the highly trampled sites from the analyses, suggested that our data of high trampling may be too scarce: the results without these sites were more in accordance with our predictions than with the full dataset.
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References
Alaruikka D, Kotze DJ, Matveinen K, Niemelä J (2002) Carabid beetle and spider assemblages along a forested urban-rural gradient in southern Finland. Journal of Insect Conservation 6:195–206.
Baars MA (1979) Catches in pitfall traps in relation to mean densities of carabid beetles. Oecologia 41: 25–46.
Beall G (1942) The transformation of data from entomological field experiments so that the analysis of variance becomes applicable. Biometrika 32:243–262.
Bhuju DR, Ohsawa M (1998) Effects of nature trails on ground vegetation and understory colonization of a patchy remnant forest in an urban domain. Biological Conservation 85:123–135.
Breslow NE, Clayton DG (1993) Approximate inference in generalized linear mixed models. Journal of the American Statistical Association 88:9–25.
Clarke GM (1987) Statistics and experimental design, 2nd Edition. A series of student texts in contemporary biology. The Camelot Press Ltd. Southampton.
Clarke KR, Warwick RM (1994) Change in marine communities: An Approach to Statistical Analysis and Interpretation. Bowne Press Ltd., UK.
Didham RK, Hammond PM, Lawton JH, Eggleton P, Stork NE (1998) Beetle species responses to tropical forest fragmentation. Ecological Monographs 68:295–323.
Duffey E (1975) The effects of human trampling on the fauna of grassland litter. Biological Conservation 7:255–274.
Florgård C (2000) Long-term changes in indigenous vegetation preserved in urban areas. Landscape and Urban Planning 52:101–116.
Frankie GW, Ehler LE (1978) Ecology of insects in urban environments. Annual Review of Entomology 23:367–387.
Gandhi KJK, Spence JR, Langor DW, Morgantini LE (2001) Fire residuals as habitat reserves for epigaeic beetles (Coleoptera: Carabidae and Staphylinidae). Biological Conservation 102:131–141.
Goodwin BJ, Fahrig L (2002) Effect of landscape structure on the movement behaviour of a specialized goldenrod beetle, Trirhabda borealis. Canadian Journal of Zoology 80:24–35.
Grandchamp AC, Niemelä J, Kotze J (2000) The effects of trampling on assemblages of ground beetles (Coleoptera, Carabidae) in urban forests in Helsinki, Finland. Urban Ecosystems 4:321–332.
Greenslade PJM (1964) Pitfall trapping as a method for studying populations of Carabidae (Coleoptera). Journal of Animal Ecology 33:301–310.
Halme E, Niemelä J (1993) Carabid beetles in fragments of coniferous forest. Annales Zoologici Fennici 30:17–30.
Heliölä J, Koivula M, Niemelä J (2001) Distribution of carabid beetles (Coleoptera, Carabidae) across a Boreal forest-clearcut ecotone. Conservation Biology 15:370–377.
Iida S, Nakashizuka T (1995) Forest fragmentation and its effects on species diversity in sub-urban coppice forests in Japan. Forest Ecology and Management 73:197–210.
Kinnunen H, Holopainen J, Pakkala T (1999) Landscape level patterns of carabid beetle communities in a forest-field mosaic. In: Kinnunen H (ed) Search of spatial scale—carabid beetle communities in agricultural landscapes, chapter 5, 13pp. Ph.D. Dissertation, Department of Ecology and Systematics, University of Helsinki, Finland.
Koivula M (2002a) Alternative harvesting methods and boreal carabid beetles (Coleoptera, Carabidae). Forest Ecology and Management 167:103–121.
Koivula M (2002b) Boreal carabid-beetle (Coleoptera, Carabidae) assemblages in thinned uneven-aged and clear-cut spruce stands. Annales Zoologici Fennici 39:131–149.
Koivula M, Kotze DJ, Hiisivuori L, Rita H (2003) Pitfall trap efficiency: do trap size, collecting fluid and vegetation structure matter? Entomologica Fennica 14:1–14.
Koivula M, Kukkonen J, Niemelä J (2002) Boreal carabid-beetle (Coleoptera, Carabidae) assemblages along the clear-cut originated succession gradient. Biodiversity and Conservation 11:1269–1288.
Kotze DJ, O'Hara RB (2003) Species decline—but why? Explanations of carabid beetle (Coleoptera, Carabidae) declines in Europe. Oecologia 135:138–148.
Kotze DJ, Samways MJ (1999) Invertebrate conservation at the interface between the grassland matrix and natural Afromontane forest fragments. Biodiversity and Conservation 8:1339–1363.
Kotze DJ, Samways MJ (2001) No general edge effects for invertebrates at Afromontane forest/grassland ecotones. Biodiversity and Conservation 10:443–466.
Liddle MJ (1997) Recreation ecology: The Ecological Impact of Outdoor Recreation and Ecotourism. Chapman and Hall, London.
Lehvävirta S (1999) Trampling and urban forest structure. Urban Ecosystems 3:45–56.
Lehvävirta S, Rita H (2002) Natural regeneration of trees in urban woodlands. Journal of Vegetation Science 13:57–66.
Lindroth CH (1985) The carabidae (Coleoptera) of Fennoscandia and Denmark, Part I. Scandinavian Science Press, Copenhagen.
Lindroth CH (1986) The carabidae (Coleoptera) of Fennoscandia and Denmark, Part II. Scandinavian Science Press, Copenhagen.
Lövei GL, Sunderland KD (1996) Ecology and behavior of ground beetles (Coleoptera: Carabidae). Annual Review of Entomology 41:231–256.
Magura T, Tóthmérész B, Molnár T (2001) Forest edge and diversity: carabids along forest-grassland transects. Biodiversity and Conservation 10:287–300.
Malmivaara M, Löfström I, Vanha-Majamaa I (2002) Anthropogenic effects on understorey vegetation in Myrtillus Type Urban Forests in Southern Finland. Silva Fennica 36:367–381.
Murcia C (1995) Edge effects in fragmented forests: Implications for conservation. Trends in Ecology and Evolution 10:58–62.
Niemelä J (1999) Ecology and urban planning. Biodiversity and Conservation 8:119–131.
Niemelä J (2000) Is there a need for a theory of urban ecology? Urban Ecosystems 3:57–65.
Niemelä J (2001) Carabid beetles (Coleoptera, Carabidae) indicating habitat fragmentation: a review. European Journal of Entomology 98:127–132.
Niemelä J, Haila Y, Halme E, Lahti T, Pajunen T, Punttila P (1988) The distribution of carabid beetles in fragments of old coniferous taiga and adjacent managed forest. Annales Zoologici Fennici 25:107–119.
Niemelä J, Haila Y, Punttila P (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.
Niemelä J, Haila Y, Ranta E (1986) Spatial heterogeneity of carabid beetle dispersion in uniform forests on the Åland Islands, SW Finland. Annales Zoologici Fennici 23:289–296.
Niemelä J, Kotze J, Ashworth A, Brandmayr P, Desender K, New T, Penev L, Samways M, Spence J (2000) The search for common anthropogenic impacts on biodiversity: a global network. Journal of Insect Conservation 4:3–9.
Niemelä J, Kotze DJ, Venn S, Penev L, Stoyanov I, Spence J, Hartley D, Montes de Oca E (2002) Carabid beetle assemblages (Coleoptera, Carabidae) across urban-rural gradients: an international comparison. Landscape Ecology 17:387–401.
Niemelä JK, Spence JR (1994) Distribution of forest dwelling carabids (Coleoptera): spatial scale and the concept of communities. Ecography 17:166–75.
Pouyat RV, McDonnell MJ, Pickett STA (1997) Litter decomposition and nitrogen mineralization in oak stands along an urban-rural land use gradient. Urban Ecosystems 1:117–31.
Rainio J, Niemelä J (2003) Ground beetles (Coleoptera: Carabidae) as bioindicators. Biodiversity and Conservation 12:487–506.
Ranney JW, Bruner MC, Levenson JB (1981) The importance of edge in the structure and dynamics of forest islands. In: Burgess RL, Sharpe DM (eds) Forest Island dynamics in man-dominated landscapes pp. 67–95. Springer-Verlag, New York.
Risser PG (1995) The status of the science examining ecotones. BioScience 45:318–325.
Rivers-Moore NA, Samways MJ (1996) Game and cattle trampling, and impacts of human dwellings on arthropods at a game park boundary. Biodiversity and Conservation 5:1545–1556.
Rudnicky JL, McDonnell MJ (1989) Forty-eight years of canopy change in a hardwood-hemlock forest in New York City (USA). Bulletin of the Torrey Botanical Club 116:52–64.
Rykken JJ, Capen DE, Mahabir SP (1997) Ground beetles as indicators of land type diversity in the green mountains of Vermont. Conservation Biology 11:522–530.
Samways MJ (1994) Insect conservation biology. Chapman & Hall, London.
Stork N (ed.) (1990) The role of ground beetles in ecological and environmental studies. Intercept Publications, Andover.
Thacker JRM (1996) Carabidologists and fragmented habitats. Trends in Ecology and Evolution 11:103–104.
Usher MB, Field JP, Bedford SE (1993) Biogeography and diversity of ground-dwelling arthropods in farm woodlands. Biodiversity Letters 1:54–62.
Venables WN, Ripley BD (2001) Modern applied statistics with S-PLUS. Springer Verlag, Berlin.
Venn SJ, Kotze DJ, Niemelä J (2003) Urbanization effects on carabid diversity in boreal forests. European Journal of Entomology 100:73–80.
White GC, Bennetts RE (1996) Analysis of frequency count data using the negative binomial distribution. Ecology 77:2549–2557.
Zar JH (1999) Biostatistical analysis, 4th Edition. Prentice Hall, New Jersey.
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Lehvävirta, S., Kotze, D.J., Niemelä, J. et al. Effects of fragmentation and trampling on carabid beetle assemblages in urban woodlands in Helsinki, Finland. Urban Ecosyst 9, 13–26 (2006). https://doi.org/10.1007/s11252-006-5526-3
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DOI: https://doi.org/10.1007/s11252-006-5526-3