Abstract
The effects of habitat area and fragmentation are confounded in many studies. Since a reduction in habitat area alone reduces patch size and increases patch isolation, many studies reporting fragmentation effects may really be documenting habitat-area effects. We designed an experimental landscape system in the field, founded on fractal neutral landscape models, to study arthropod community responses to clover habitat in which we adjusted the level of fragmentation independently of habitat area. Overall, habitat area had a greater and more consistent effect on morphospecies richness than fragmentation. Morphospecies richness doubled between 10 and 80% habitat, with the greatest increase occurring up to 40% habitat. Fragmentation had a more subtle and transient effect, exhibiting an interaction at intermediate levels of habitat only at the start of the study or in the early-season (June) survey. In these early surveys, morphospecies richness was higher in clumped 40–50% landscapes but higher in fragmented landscapes at 60–80% habitat. Rare or uncommon species are expected to be most sensitive to fragmentation effects, and we found a significant interaction with fragmentation at intermediate levels of habitat for these types of morphospecies in early surveys. Although the effects of fragmentation are expected to amplify at higher trophic levels, all trophic levels exhibited a significant fragmentation effect at intermediate levels of habitat in these early surveys. Predators/parasitoids were more sensitive to habitat area than herbivores, however. Thus, our results confirm that habitat area is more important than fragmentation for predicting arthropod community responses, at least in this agricultural system.
Similar content being viewed by others
References
Andrén H (1994) Effects of habitat fragmentation on birds and mammals in landscapes with different proportions of suitable habitat: a review. Oikos 71:355–366
Bender DJ, Tischendorf L, Fahrig L (2003) Evaluation of patch isolation metrics for predicting animal movement in binary landscapes. Landscape Ecol 18:17–39
Bolger DT, Suarez AV, Crooks KR, Morrison SA, Case TJ (2000) Arthropods in urban habitat fragments in southern California: area, age and edge effects. Ecol Appl 10:1230–1248
Collinge SK (2000) Effects of grassland fragmentation on insect species loss, colonization, and movement patterns. Ecology 81:2211–2226
Collinge SK (2009) Ecology of fragmented landscapes. Johns Hopkins University Press, Baltimore
Crist TO, Ahern RG (1999) Effects of habitat patch size and temperature on the distribution and abundance of ground beetles (Coleoptera: Carabidae) in an old field. Environ Entomol 28:681–689
Cronin JT (2003) Patch structure, oviposition behavior, and the distribution of parasitism risk. Ecol Monogr 73:283–300
Davies K, Margules C, Lawrence JF (2000) Which traits of species predict population declines in experimental forest fragments? Ecology 81:1450–1461
Debinski DM, Holt RD (2000) A survey and overview of habitat fragmentation experiments. Conserv Biol 14:342–355
Ewers RM, Didham RK (2006) Confounding factors in the detection of species responses to habitat fragmentation. Biol Rev 81:117–142
Ewers RM, Didham RK (2007) The effect of fragment shape and species’ sensitivity to habitat edges on animal population size. Conserv Biol 21:926–936
Fahrig L (1997) Relative effects of habitat loss and fragmentation on population extinction. J Wildl Manag 61:603–610
Fahrig L (2003) Effects of habitat fragmentation on biodiversity. Annu Rev Ecol Syst 34:487–515
Gardner RH, Milne BT, Turner MG, O’Neill RV (1987) Neutral models for the analysis of broad-scale landscape pattern. Landscape Ecol 1:19–28
Golden DM, Crist TO (1999) Experimental effects of habitat fragmentation on old-field canopy insects: community, guild and species responses. Oecologia 118:371–380
Grez A, Zaviezo T, Tischendorf L, Fahrig L (2004) A transient, positive effect of habitat fragmentation on insect population densities. Oecologia 141:444–451
Haynes KJ, Crist TO (2009) Insect herbivory in an experimental agroecosystem: the relative importance of habitat area, fragmentation, and the matrix. Oikos 118:1477–1486
Holt RD, Lawton JH, Polis GA, Martinez ND (1999) Trophic rank and the species–area relationship. Ecology 80:1495–1504
Kruess A, Tscharntke T (1994) Habitat fragmentation, species loss and biological control. Science 264:1581–1584
Kruess A, Tscharntke T (2000) Species richness and parasitism in a fragmented landscape: experiments and field studies with insects on Vicia sepium. Oecologia 122:129–137
Lawton JH (1995) Ecological experiments with model systems. Science 269:328–331
McGarigal K, Cushman SA (2002) Comparative evaluation of experimental approaches to the study of habitat fragmentation effects. Ecol Appl 12:335–345
Öckinger E, Schweiger O, Crist TO, Debinski DM, Krauss J, Kuussaari M, Petersen JD, Pöyry J, Settele J, Summerville KS, Bommarco R (2010) Life-history traits predict species responses to habitat area and isolation: a cross-continental synthesis. Ecol Lett 13:969–979
Roland J, Taylor PD (1997) Insect parasitoid species respond to forest structure at different scales. Nature 386:710–713
Steffan-Dewenter I, Tscharntke T (2002) Insect communities and biotic interactions on fragmented calcareous grasslands—a mini review. Biol Conserv 104:275–284
Summerville KS, Crist TO (2001) Effects of experimental habitat fragmentation on patch use by butterflies and skippers (Lepidoptera). Ecology 82:1360–1370
Thies C, Tscharntke T (1999) Landscape structure and biological control in agroecosystems. Science 285:893–895
Tscharntke T, Steffan-Dewenter I, Kruess A, Thies C (2002) Characteristics of insect populations on habitat fragments: a mini-review. Ecol Res 17:229–239
Van Nouhuys S (2005) Effects of habitat fragmentation at different trophic levels in insect communities. Ann Zool Fennici 42:433–447
Wiens JA, Stenseth NC, Van Horne B, Ims RA (1993) Ecological mechanisms and landscape ecology. Oikos 66:369–380
With KA (1997) The application of neutral landscape models in conservation biology. Conserv Biol 11:1069–1080
With KA, King AW (1999a) Dispersal success in fractal landscapes: a consequence of lacunarity thresholds. Landscape Ecol 14:73–82
With KA, King AW (1999b) Extinction thresholds in fractal landscapes. Conserv Biol 13:314–326
With KA, Cadaret SJ, Davis C (1999) Movement responses to patch structure in experimental fractal landscapes. Ecology 80:1340–1353
With KA, Pavuk DM, Worchuck JL, Oates RK, Fisher JL (2002) Threshold effects of landscape structure on biological control in agroecosystems. Ecol Appl 12:52–65
Zabel J, Tscharntke T (1998) Does fragmentation of Urtica habitats affect phytophagous and predatory insects differentially? Oecologia 116:419–425
Acknowledgments
This research was supported by a grant from the National Science Foundation (DEB-9610159). We appreciate the assistance of the dozen or so undergraduates who helped to establish and maintain this experimental system. We thank L. Murray for statistical advice, J. R. Nechols and two anonymous reviewers for their comments on the manuscript.
Author information
Authors and Affiliations
Corresponding author
Electronic supplementary material
Below is the link to the electronic supplementary material.
Rights and permissions
About this article
Cite this article
With, K.A., Pavuk, D.M. Habitat area trumps fragmentation effects on arthropods in an experimental landscape system. Landscape Ecol 26, 1035–1048 (2011). https://doi.org/10.1007/s10980-011-9627-x
Received:
Accepted:
Published:
Issue Date:
DOI: https://doi.org/10.1007/s10980-011-9627-x