ECOGRAPHY 27: 417 /426, 2004

Ground beetle species (Coleoptera, Carabidae) associations with land cover variables in northern England and southern Scotland
M. D. Eyre and M. L. Luff

Eyre, M. D. and Luff, M. L. 2004. Ground beetle species (Coleoptera, Carabidae) associations with land cover variables in northern England and Scotland. / Ecography 27: 417 /426. Distribution data concerning 172 ground beetle species derived from 1145 pitfall trap sites in northern England and southern Scotland were used to assess the relationship between species distribution and 12 satellite-derived land cover variables at the regional scale. A number of species were strongly associated with one cover type and negatively with others. The major variation was for preferences for covers in upland or lowland parts of the region. Other distinct preferences for some species were covers such as those at the coast whilst a number of common species showed no strong preference for any cover variable. The synthesis of ground beetle species distribution and satellitederived cover data is discussed in relation to environmental assessment and change. M. D. Eyre (m.d.eyre@ncl.ac.uk) and M. L. Luff, Centre for Life Science Modelling, School of Biology, The University, Newcastle upon Tyne, U.K. NE1 7RU.

Ground beetle research within European landscapes tends to be confined to one land cover type or limited by activities such as farming. The distribution of ground beetle species and assemblages in wooded habitats have received considerable attention with investigations concerned with both broadleaved woodland (Assmann 1999, Desender et al. 1999, Judas et al. 2002, Moore et al. 2002) and conifer forests (Butterfield 1997, Jukes et al. 2001, Koivula and Niemela 2002). Work on the ground beetles of grasslands tends to be linked with other invertebrate groups (Niemela and Baur 1998, So derstrom et al. 2001, Vessby et al. 2002), as does work on upland areas (Dennis et al. 2002, Eyre et al. 2003a). More limited areas within the landscape such as riverine sediments and corridors (Eyre et al. 2001, Bonn et al. 2002), saltmarshes (Elkaim and Rybarczyk 2000, Irmler et al. 2002) and urban and post-industrial land (Kielhorn et al. 1999, Andersen 2000, Bra ndle et al. 2000) have also been assessed with regard to ground beetle distribution. Research at a larger scale has focused on the effects of the composition of the agricultural land-

scape (Helenius et al. 2001, Kinnunen et al. 2001, Basedow 2002, Pena et al. 2003) whilst Niemela et al. (2002) investigated ground beetle species distribution across urban-rural gradients in three countries. Satellite-derived land cover data has been applied consistently to assess vegetation distribution and change and has been used in investigations into changes in such covers as rain forests (Wittmann et al. 2002), boreal woodlands (Masek 2001), deserts (Muldavin et al. 2001), tundra (Walker 2000), wetlands (Townsend and Walsh 2001) and coasts (Donoghue and Mironnet 2002) and to assess landscape fragmentation (Petit and Lambin 2002). The use of this cover data has expanded into applications concerned with habitats of mammals (Velazquez et al. 2001, Jepsen et al. 2002), birds (Groom and Grubb 2002, Daily et al. 2001) and vertebrates (Cassidy et al. 2001). Invertebrate habitat assessments using satellite imagery has concentrated on disease vectors (Dister et al. 1997, Daniel et al. 1998, Moncayo et al. 2000) and pests of grasslands, forests and crops (Dreiser 1994, Grilli and Gorla 1997, Brewster et al. 1999, Radeloff et

Accepted 12 January 2004 Copyright # ECOGRAPHY 2004 ISSN 0906-7590

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al. 2000). There has been some assessment of woodland invertebrate taxa distribution (Va sa nen and Helio vaara 1994, Chust et al. 2000) and some conservation assessments with butterflies and dragonflies (Fuller et al. 1998, Moilanen and Hanski 1998, Debinski et al. 1999, Wessels et al. 2000). Eyre et al. (2003b) classified British 10 km grid squares according to pooled ground beetle species lists and related this to satellite-derived land cover data. The incorporation of invertebrate data with this sort of cover data has concentrated on the identification of habitat types for various assemblages, with little work carried out on the assessment of the relationship of individual species with specific land cover types. The present work, therefore, uses ground beetle species data from 1145 sites in northern England and southern Scotland and 12 land cover types derived from Landsat satellite imagery in order to investigate relationship of each species to each cover type.
Fig. 1. Map showing the distribution of the 259 1 km squares in northern England and southern Scotland containing the 1145 pitfall sites, with national grid 100 km numbers.

Methods
Data
Species The occurrence of 172 species from 1145 pitfall sites in 259 1 km national grid squares in northern England and southern Scotland were used for analysis. The data were generated between 1985 and 2002. Some of the data has been previously used in a number of publications concerned with the distribution of species assemblages (Gardner 1991, Luff et al. 1992, Butterfield et al. 1995) but they have not been used in work relating individual species to specific land cover variables. The 1145 sites were in the 259 1 km national grid squares shown in Fig. 1. Ground beetle presence/ absence species lists, for each site, were generated using the pitfall trapping method outlined by Luff (1996), with 5 /10 traps (8.5 cm diameter, 10 cm deep, part filled with preservative) used between May and September. Nomenclature follows Luff (1998). Land cover The Landsat Thematic Mapper-based land cover data were released in the Countryside Information System (CIS) (Anon. 1995) as a list of the number of hectares of cover per 1 km national grid square (%) of 17 cover types and unclassified pixels. The data were derived from images generated between 1987 and 1990 (Fuller et al. 1994). The 17 land classes in the CIS were reduced to 12 classes for these analyses and Table 1 shows the classes used and an indication of altitudinal distribution. Three sea and coast covers were merged to form a coastal cover; urban, suburban and inland bare ground were 418

combined to produce an urban cover type; the shrub heath and shrub/grass heath were combined to produce one shrub heath cover.

Table 1. The Countryside Information System (CIS) land cover classes and the derived land classes used in the analyses, with indications of altitudinal distribution. CIS cover Bog (herbaceous) Bracken Sea/estuary Coastal bare ground (beach/ mudflats/cliffs) Saltmarsh Coniferous/evergreen woodland Deciduous/mixed wood Grass moor/dune grass/rough pasture Inland water Pasture/meadow/amenity grass Rough grass/marsh Shrub heath Shrub/grass heath Tilled land (arable crops) Suburban/rural development Urban development Inland bare ground Unclassified Derived covers Bog (mainly upland) Bracken (mainly upland)

Coastal (lowland) Coniferous woodland (mainly upland) Deciduous wood (mainly lowland) Heath grassland (upland) Inland water (upland and lowland) Managed grassland (mainly lowland) Rough grass (mainly lowland) Shrub heath (upland) Tilled land (lowland) Urban (lowland) /

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Analysis
CANOCO Ver. 4 (Ter Braak and Smilauer 1998) was used to analyse the 1145 ground beetle species lists with the proportional land cover data from appropriate 1 km squares. The 1 km land class data was used since it was this was the smallest scale satellite-derived cover data available. This version of CANOCO produces a speciesenvironment table showing the weighted averages with respect to each standardised environmental variable, in this case the twelve cover types. This table indicates the relationship, or lack thereof, between cover types and species. An indication of which were the most important land covers was provided using automatic forward selection of the 12 cover variables within CANOCO with their significance calculated using Monte Carlo permutation tests.

Results
The relationships between individual species and the 12 land cover variables, shown as weighted averages, are given in Table 2. Seven species were related to the bog variable. One was Agonum gracile Sturm, a wetland species, but the others tended to be upland species (e.g. Carabus arvensis Herbst, C. glabratus Paykull, Patrobus assimilis Chaudoir), more usually associated with heath grassland. The species positively associated with the bracken variable were also associated with other upland covers, especially heath grassland and shrub heath (e.g. Bembidion nigricorne Gyllenhal, Miscodera arctica (Paykull), Notiophilus germinyi Fauvel) but there were also positive relationships with coastal species (e.g. Bembidion minimum (Fabricius), Dicheirotrichus gustavi Crotch), reflecting the presence of bracken on well-drained coastal grasslands. Some species associated in this study with conifer woodland were those more usually found in wetland sites (Agonum gracile , Bembidion nitidulum (Marsham)) or associated with covers found next to coniferous plantations in the region, especially shrub heath (e.g. Bradycellus collaris (Paykull)) but there were also strong associations with species known to inhabit conifer woods (Calathus micropterus (Duftschmid), Trechus rubens (Fabricius)). A number of species usually found in woodland sites were positively related to the deciduous woodland variable (Abax parallelepipedus (Piller & Mitterpacher), Agonum assimile (Paykull), Calathus piceus (Marsham), Leistus fulvibarbis Dejean) and a number of species were negatively related to this variable, especially coastal species (e.g. Broscus cephalotes (Linnaeus), Calathus mollis (Marsham)). There were differences in the associations of species preferring the main upland land cover types. There were those with a clear relationship with heath grass (e.g. Amara lunicollis Schioedte, Carabus arvensis , C. glabraECOGRAPHY 27:4 (2004)

tus , Pterostichus versicolor (Sturm)) and other with shrub heath (e.g. Bradycellus ruficollis (Stephens), Miscodera arctica , Olisthopus rotundatus (Paykull), Trichocellus cognatus (Gyllenhal)). However some species were found to be strongly associated with both these covers (Agonum ericeti (Panzer), Carabus nitens Linnaeus, Dyschirius globosus (Herbst)). Some species were positively related to managed grassland (Carabus granulatus Linnaeus, Pterostichus minor (Gyllenhal), P. vernalis (Panzer)) and some to the tilled land (Bembidion quadrimaculatum (Linnaeus), Stomis pumicatus (Panzer), Trechus micros (Herbst)) but there are a number (e.g. Agonum dorsale (Pontoppidan), Bembidion aeneum Germar, B. guttula (Fabricius), B. tetracolum Say, Clivina fossor (Linnaeus), Pterostichus melanarius (Illiger)) associated with both of these covers. Species related to the urban variable tended to be also related to coastal (e.g. Broscus cephalotes , Calathus cinctus Motschulsky, Dromius melanocephalus Dejean), to tilled land (e.g. Bembidion obtusum Serville, Harpalus rubripes (Duftschmid), H. rufipes (De Geer)) or to both (e.g. Dromius notatus Stephens, Harpalus affinis (Schrank)) and reflected the lowland land cover preferences of these species. Of the species positively associated with rough grass, some were more usually associated with uplands (Agonum ericeti , Cymindis vaporariorum (Linnaeus)), some with wetlands (Agonum gracile , A. marginatum (Linnaeus)) and some with coastal (Amara lucida (Duftschmid), A. spreta Dejean, Broscus cephalotes , Calathus mollis ). There were some wetland species positively related to inland water (Agonum micans Nicolai, Trichocellus placidus (Gyllenhal)) but others with no obvious relationship to water (Abax parallelepidedus , Harpalus rufipes , H. rufibarbis (Fabricius)). The largest weighted averages were for species associated with the coast (e.g. Amara lucida. A. spreta , Broscus cephalotes , Calathus mollis ). Differences between species with preferences for either upland or lowland covers were emphasised by the number of species negatively related with cover variables. Heath grassland and shrub heath, mainly upland covers, had the most species with strong negative relationships, with a considerable number negatively related to the mainly lowland covers of tilled land and managed grassland. A number of species usually recorded from riverine sediments were associated with several cover variables. Bembidion atrocoeruleum Stephens and B. tibiale (Duftschmid) were related to managed grassland, B. punctulatum Drapiez and B. schueppeli Dejean to tilled land and Agonum albipes (Fabricius), Amara fulva (Mueller) and Asaphidion pallipes (Duftschmid) to both deciduous woodland and managed grassland. These associations appear to reflect the landscape through which the rivers flow. Twenty species had no obvious association with any cover variable. These were all common species in the region (e.g. Calathus fuscipes 419

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Bog (/0.11 (/0.32 (/0.28 (/0.34 0.36 0.40 1.05 (/0.23 (/0.29 (/0.40 (/0.15 (/0.28 (/0.30 (/0.25 (/0.39 (/0.01 (/0.26 (/0.40 (/0.47 0.32 (/0.21 (/0.24 (/0.06 (/0.23 (/0.47 (/0.36 (/0.41 (/0.37 (/0.36 (/0.38 (/0.28 (/0.33 (/0.37 0.34 (/0.33 (/0.35 (/0.27 (/0.15 (/0.41 (/0.26 (/0.40 (/0.19 (/0.36 (/0.18 (/0.27 (/0.37 (/0.33 (/0.35 (/0.31 0.62 0.19 0.77 0.05 (/0.56 (/0.34 (/0.34 (/0.28 0.61 0.13 (/0.11 (/0.13 (/0.18 0.20 (/0.24 (/0.19 0.30 0.73 (/0.65 (/0.44 0.20 0.18 0.03 0.34 (/0.65 (/0.55 0.90 0.57 1.61 (/0.11 (/0.13 0.20 0.21 (/0.30 (/0.53 0.62 0.08 0.16 0.16 0.40 (/0.57 0.47 (/0.33 0.07 0.32 0.75 0.43 0.09 0.29 0.63 0.15 0.21 0.20 (/0.21 0.02 (/0.05 0.08 0.96 (/0.21 0.01 0.20 (/0.01 0.14 (/0.07 0.24 (/0.03 0.08 (/0.21 (/0.07 0.07 0.02 0.34 (/0.03 (/0.21 0.04 0.00 0.44 0.26 0.00 0.00 0.01 0.07 0.29 0.12 0.19 0.22 0.06 0.37 0.12 (/0.17 (/0.01 (/0.21 (/0.05 0.07 0.32 (/0.05 0.37 0.24 0.04 (/0.35 (/0.41 (/0.47 (/0.22 (/0.03 (/0.19 0.46 (/0.47 (/0.27 (/0.19 (/0.25 (/0.15 (/0.27 (/0.35 (/0.10 (/0.15 (/0.39 0.03 0.15 (/0.26 (/0.19 (/0.38 (/0.41 (/0.09 0.25 (/0.31 (/0.40 (/0.32 (/0.20 (/0.18 (/0.17 (/0.51 (/0.07 (/0.18 (/0.34 (/0.31 (/0.30 (/0.51 0.13 1.33 (/0.15 1.37 0.07 (/0.30 (/0.44 0.04 (/0.34 (/0.29 0.18 (/0.25 (/0.09 (/0.27 0.25 0.06 0.93 (/0.23 (/0.08 (/0.23 0.16 (/0.21 (/0.13 (/0.14 (/0.10 (/0.02 (/0.08 (/0.23 (/0.33 (/0.02 0.46 (/0.11 (/0.31 0.55 (/0.33 (/0.28 (/0.30 (/0.29 (/0.29 (/0.25 (/0.22 (/0.05 (/0.20 1.54 (/0.18 (/0.26 (/0.17 (/0.24 (/0.25 (/0.04 (/0.31 (/0.28 (/0.28 0.68 (/0.32 (/0.22 (/0.25 (/0.12 (/0.13 (/0.32 (/0.40 (/0.54 (/0.62 0.83 0.46 0.27 (/0.55 (/0.59 (/0.49 0.04 (/0.49 (/0.53 (/0.31 (/0.36 0.03 (/0.45 (/0.54 (/0.63 0.92 (/0.26 (/0.39 (/0.50 (/0.44 (/0.67 (/0.37 (/0.48 (/0.57 (/0.45 (/0.50 (/0.41 (/0.20 (/0.57 0.29 (/0.26 (/0.49 (/0.43 (/0.26 (/0.62 (/0.27 (/0.58 (/0.04 (/0.29 0.08 (/0.52 (/0.57 (/0.51 (/0.33 (/0.36 (/0.28 (/0.48 (/0.47 (/0.50 0.78 0.01 (/0.02 (/0.40 (/0.44 (/0.51 (/0.33 (/0.40 (/0.39 (/0.45 (/0.48 (/0.25 (/0.44 (/0.51 (/0.50 (/0.18 (/0.27 (/0.44 (/0.51 (/0.39 (/0.50 (/0.36 (/0.51 (/0.50 (/0.51 (/0.51 (/0.45 (/0.48 (/0.51 (/0.16 (/0.48 (/0.49 (/0.43 (/0.20 (/0.50 0.14 (/0.49 (/0.50 1.82 (/0.28 (/0.51 (/0.49 (/0.30 (/0.46 (/0.47 0.15 0.69 0.42 0.68 (/0.76 (/0.24 (/0.68 (/0.38 0.09 0.97 0.42 0.12 0.24 0.37 (/0.09 0.16 0.25 0.40 (/0.96 (/0.23 0.33 0.55 0.34 0.00 (/0.98 (/0.58 (/0.20 0.31 0.79 0.22 0.50 1.06 0.55 (/0.23 1.06 1.05 0.61 0.28 0.54 0.16 (/0.45 0.44 (/0.35 (/0.24 0.29 0.88 0.40 0.38 0.60 0.24 0.04 0.14 0.13 (/0.42 (/0.24 0.22 0.88 1.08 (/0.20 (/0.11 0.86 0.46 0.49 1.22 0.16 0.42 0.30 2.52 (/0.33 (/0.02 0.09 0.61 0.71 2.63 1.79 0.44 0.47 0.31 0.91 0.34 (/0.29 (/0.10 (/0.38 (/0.11 0.23 0.25 0.31 (/0.44 (/0.19 1.33 (/0.14 (/0.33 0.28 1.02 (/0.36 0.34 (/0.08 0.05 0.37 0.61 0.91 0.91 (/0.74 (/0.33 (/0.31 0.38 0.64 0.54 0.05 0.32 0.57 0.45 0.21 0.01 0.50 0.64 (/0.31 (/0.57 0.25 0.51 1.01 0.58 (/0.32 (/0.11 1.43 0.93 0.44 0.40 0.62 0.10 1.09 (/0.53 0.09 0.24 0.57 0.50 1.33 0.21 0.25 0.70 (/0.87 0.08 0.88 0.74 0.59 0.84 0.61 (/0.23 0.25 (/0.12 0.33 0.78 0.06 0.68 1.64 0.29 (/0.31 0.04 0.40 0.36 0.09 0.55 (/0.01 0.08 0.14 0.95 0.02 (/0.03 0.09 (/0.42 (/0.22 1.01 0.52 (/0.33 (/0.04 (/0.39 0.18 0.07 0.10 (/0.04 (/0.04 (/0.12 (/0.04 (/0.03 0.04 0.04 (/0.08 0.19 (/0.31 (/0.25 (/0.40 (/0.21 0.01 (/0.19 (/0.04 (/0.05 Bra CoW DeW HeG ShH MGr InW Urb TiL RoG Coa (/0.23 (/0.07 (/0.23 (/0.14 (/0.22 (/0.08 0.07 0.82 (/0.23 0.29 0.06 1.13 0.31 0.00 1.33 0.28 0.45 0.08 4.42 (/0.09 (/0.23 (/0.06 (/0.23 (/0.23 4.49 2.56 (/0.23 0.02 (/0.23 0.88 0.19 (/0.23 (/0.09 (/0.23 (/0.23 (/0.23 (/0.05 (/0.17 (/0.23 (/0.21 2.19 (/0.23 (/0.23 (/0.23 (/0.23 (/0.23 (/0.08 (/0.23 (/0.21

Table 2. Species-environment table showing the weighted averages, with respect to standardised environmental variables (Bra, Bracken; CoW, Coniferous wood; DeW, Deciduous woodland; HeG, Heath grassland; ShH, Shrub heathland; MGr, Managed grassland; InW, Inland water; Urb, Urban; TiL, Tilled land; RoG, Rough grass; Coa, Coastal). The 126 species with more than 10 records are included and averages of 0.50 and above are emboldened.

Species

Abax parallelepipedus Pill & Mitt. Agonum albipes (F.) Agonum assimile (Payk.) Agonum dorsale (Pont.) Agonum ericeti (Panz.) Agonum fuliginosum (Panz.) Agonum gracile Sturm Agonum marginatum (L.) Agonum micans Nicol. Agonum moestum (Dufts.) Agonum muelleri (Herbst) Amara aenea (De Geer) Amara apricaria (Payk.) Amara aulica (Panz.) Amara bifrons (Gyll.) Amara communis (Panz.) Amara familiaris (Dufts.) Amara fulva (Muell.) Amara lucida (Dufts.) Amara lunicollis Schioedte Amara ovata (F.) Amara plebeja (Gyll.) Amara praetermissa (Sahl.) Amara similata (Gyll.) Amara spreta Dejean Amara tibialis (Payk.) Asaphidion curtum Heyden Asaphidion flavipes (L.) Asaphidion pallipes (Dufts.) Badister bipustulatus (F.) Bembidion aeneum Germar Bembidion atrocoeruleum Steph. Bembidion biguttatum (F.) Bembidion bruxellense Wesm. Bembidion decorum (Zenk. in Panz.) Bembidion femoratum Sturm Bembidion guttula (F.) Bembidion lampros (Herbst) Bembidion litorale (Oliv.) Bembidion mannerheimei Sahl. Bembidion minimum (F.) Bembidion monticola Sturm Bembidion nigricorne Gyll. Bembidion nitidulum (Marsh.) Bembidion obtusum Serv. Bembidion punctulatum Drap. Bembidion quadrimaculatum (L.) Bembidion schueppeli Dejean Bembidion tetracolum Say

ECOGRAPHY 27:4 (2004)

Table 2 (Continued ) Bog (/0.32 (/0.39 0.16 (/0.07 (/0.35 (/0.45 (/0.38 (/0.43 (/0.25 0.05 0.51 (/0.43 (/0.23 0.70 0.88 (/0.18 0.00 0.18 0.29 0.33 (/0.24 (/0.37 (/0.27 (/0.01 0.24 (/0.38 (/0.37 (/0.44 (/0.35 0.04 (/0.29 (/0.39 (/0.24 0.04 (/0.27 (/0.38 (/0.30 (/0.29 (/0.31 (/0.24 0.09 0.07 (/0.34 (/0.22 (/0.19 (/0.27 0.17 0.25 (/0.04 0.15 0.58 1.18 (/0.86 (/0.54 (/0.68 (/0.42 (/0.67 (/0.66 (/0.58 0.38 (/0.04 (/0.64 (/0.64 0.49 (/0.25 (/0.48 1.00 (/0.01 (/0.65 (/0.37 (/0.37 (/0.35 0.69 0.58 (/0.16 (/0.59 (/0.45 (/0.11 (/0.47 (/0.17 (/0.65 0.01 (/0.09 (/0.09 (/0.19 (/0.24 (/0.09 0.32 (/0.38 0.08 0.48 (/0.22 0.12 (/0.43 (/0.69 0.45 0.60 (/0.32 (/0.52 (/0.02 (/0.47 (/0.40 (/0.20 0.91 0.10 0.80 (/0.32 (/0.11 (/0.17 0.38 (/0.08 0.17 0.49 (/0.05 (/0.33 0.16 (/0.14 (/0.50 (/0.08 0.59 0.26 0.15 0.67 (/0.35 (/0.32 0.02 (/0.42 1.15 (/0.31 (/0.07 (/0.39 0.55 0.10 (/0.38 (/0.10 0.32 (/0.13 0.01 (/0.26 (/0.35 0.08 (/0.40 0.15 (/0.08 (/0.36 0.67 (/0.21 (/0.31 0.15 0.47 (/0.04 0.73 0.04 (/0.19 0.67 0.51 (/0.06 (/0.30 (/0.27 (/0.30 (/0.27 (/0.17 (/0.22 0.83 (/0.25 (/0.03 (/0.26 0.19 0.14 (/0.13 (/0.16 0.07 0.19 (/0.11 (/0.24 (/0.13 0.25 0.30 (/0.30 (/0.24 (/0.14 (/0.24 (/0.28 (/0.09 (/0.23 (/0.27 0.10 (/0.31 (/0.31 (/0.21 (/0.26 (/0.24 (/0.11 0.17 0.00 (/0.30 (/0.15 (/0.09 0.43 0.14 (/0.22 0.10 (/0.06 (/0.23 0.45 (/0.19 (/0.22 (/0.24 0.59 (/0.62 (/0.19 (/0.40 0.13 (/0.18 (/0.23 (/0.65 0.81 (/0.49 (/0.60 (/0.22 0.03 (/0.45 (/0.19 (/0.16 (/0.04 0.35 0.18 0.25 (/0.42 (/0.50 (/0.18 (/0.34 (/0.07 (/0.42 (/0.03 0.07 (/0.10 (/0.01 1.40 0.30 0.04 (/0.10 0.11 0.83 0.14 0.07 (/0.16 (/0.36 0.24 0.35 (/0.24 (/0.37 0.23 (/0.37 (/0.50 (/0.24 (/0.17 0.15 (/0.09 (/0.43 (/0.60 (/0.56 (/0.46 (/0.37 (/0.08 0.07 (/0.59 (/0.46 0.90 0.98 (/0.21 (/0.08 0.79 0.46 0.39 0.15 (/0.42 (/0.35 0.18 0.18 (/0.61 (/0.51 (/0.51 (/0.55 0.85 (/0.35 (/0.59 (/0.46 0.72 (/0.28 (/0.44 (/0.47 (/0.40 (/0.50 (/0.59 0.00 (/0.03 (/0.60 (/0.23 (/0.31 (/0.20 0.22 0.40 (/0.16 0.14 0.92 (/0.48 1.64 0.17 1.66 (/0.50 (/0.51 (/0.51 1.28 (/0.40 0.33 0.66 (/0.51 (/0.44 (/0.01 0.02 (/0.41 0.22 1.05 0.42 0.22 0.84 (/0.49 (/0.38 (/0.07 0.93 (/0.49 (/0.49 (/0.48 (/0.50 0.87 (/0.33 (/0.49 (/0.41 (/0.24 (/0.51 (/0.51 (/0.51 (/0.51 (/0.22 (/0.50 0.22 (/0.10 (/0.51 2.07 (/0.37 (/0.33 0.40 0.72 (/0.07 1.10 0.10 0.02 (/0.21 0.04 (/0.21 0.31 (/0.11 (/0.11 (/0.20 (/0.02 (/0.09 0.13 (/0.21 0.09 (/0.21 (/0.02 (/0.21 0.11 (/0.21 (/0.07 0.03 (/0.01 0.26 0.02 0.15 (/0.21 (/0.21 (/0.05 (/0.08 0.29 (/0.21 0.35 0.34 0.13 (/0.02 0.59 0.57 0.20 0.30 0.04 (/0.03 (/0.01 (/0.07 0.38 (/0.21 0.03 0.01 (/0.13 (/0.17 (/0.03 (/0.18 (/0.07 (/0.26 (/0.38 0.17 (/0.33 1.93 1.63 1.54 0.29 0.37 (/0.04 (/0.41 1.46 0.25 (/0.48 (/0.50 (/0.22 0.08 (/0.43 (/0.30 (/0.19 0.09 0.06 0.08 (/0.12 (/0.32 1.08 1.04 1.32 0.76 (/0.36 0.82 0.35 1.21 0.22 1.67 1.39 0.65 1.24 0.80 (/0.02 (/0.21 (/0.10 1.98 (/0.32 0.18 (/0.38 (/0.08 (/0.26 0.03 (/0.33 (/0.40 0.09 (/0.83 (/0.34 (/0.84 0.32 (/0.16 (/0.03 (/0.62 0.38 (/0.15 (/0.75 (/0.26 0.66 (/0.81 (/0.84 0.12 (/0.11 (/0.88 (/0.64 (/0.49 (/0.51 0.66 0.63 0.06 (/0.61 0.16 0.35 (/0.31 0.64 (/0.76 0.06 1.06 0.51 (/0.30 0.60 0.48 0.74 0.20 0.29 1.08 (/0.06 0.09 0.04 (/0.84 0.39 0.20 (/0.56 (/0.71 0.22 (/0.76 (/0.61 0.15 (/0.05 (/0.09 (/0.15 0.42 1.00 0.64 0.02 0.04 (/0.09 (/0.06 0.96 (/0.19 0.33 (/0.05 (/0.12 0.00 (/0.13 (/0.04 (/0.11 0.20 (/0.07 (/0.05 (/0.12 2.43 0.17 0.53 0.86 0.56 (/0.05 0.65 0.13 0.21 (/0.18 (/0.41 0.01 0.12 0.36 (/0.02 (/0.23 (/0.16 (/0.03 0.86 (/0.22 (/0.01 (/0.03 0.00 (/0.01 0.05 (/0.07 0.30 Bra CoW DeW HeG ShH MGr InW Urb TiL RoG Coa (/0.23 (/0.23 0.35 (/0.20 1.16 3.58 3.27 1.10 0.26 (/0.06 (/0.23 3.55 (/0.23 (/0.23 (/0.23 (/0.23 (/0.06 (/0.23 (/0.19 (/0.17 (/0.22 (/0.23 (/0.23 (/0.11 (/0.22 2.84 1.40 2.87 0.65 (/0.03 0.57 0.47 0.81 (/0.23 (/0.23 0.80 (/0.02 1.34 0.67 (/0.19 (/0.13 (/0.16 2.61 (/0.23 (/0.02 (/0.23 0.10 0.08 0.01 (/0.19 (/0.23

ECOGRAPHY 27:4 (2004)

Species

Bembidion tibiale (Dufts.) Bradycellus collaris (Payk.) Bradycellus harpalinus Serv. Bradycellus ruficollis (Steph.) Bradycellus verbasci (Dufts.) Broscus cephalotes (L.) Calathus cinctus Mots. Calathus erratus (Sahl.) Calathus fuscipes (Goeze) Calathus melanocephalus (L.) Calathus micropterus (Dufts.) Calathus mollis (Marsh.) Calathus piceus (Marsh.) Carabus arvensis Herbst Carabus glabratus Payk. Carabus granulatus L. Carabus nemoralis Muell. Carabus nitens L. Carabus problematicus Herbst Carabus violaceus L. Cicindela campestris L. Clivina collaris (Herbst) Clivina fossor (L.) Cychrus caraboides L. Cymindis vaporariorum (L.) Dicheirotrichus gustavi Crotch Dromius linearis (Oliv.) Dromius melanocephalus Dejean Dromius notatus Steph. Dyschirius globosus (Herbst) Elaphrus cupreus Dufts. Elaphrus riparius (L.) Harpalus affinis (Schrank) Harpalus latus (L.) Harpalus rubripes (Dufts.) Harpalus rufibarbis (F.) Harpalus rufipes (De Geer) Harpalus tardus (Panz.) Leistus ferrugineus (L.) Leistus fulvibarbis Dejean Leistus rufescens (F.) Loricera pilicornis (F.) Metabletus foveatus (Four.) Miscodera arctica (Payk.) Nebria brevicollis (F.) Nebria gyllenhali (Schoen.) Nebria salina Fair. & Lab. Notiophilus aquaticus L. Notiophilus biguttatus (F.) Notiophilus germinyi Fauvel Notiophilus palustris (Dufts.)

421

422
Bog (/0.34 (/0.17 0.70 (/0.20 0.46 1.01 (/0.27 0.48 (/0.14 (/0.14 (/0.21 0.18 0.01 0.40 (/0.25 (/0.21 0.11 (/0.31 (/0.10 (/0.23 (/0.10 (/0.25 (/0.09 (/0.24 0.05 (/0.29 (/0.53 (/0.21 0.85 (/0.28 0.52 0.78 (/0.52 0.89 (/0.02 (/0.39 (/0.39 0.27 (/0.13 0.88 (/0.32 (/0.26 1.15 (/0.62 0.03 (/0.56 (/0.27 (/0.40 (/0.17 (/0.47 (/0.16 (/0.31 0.99 (/0.23 (/0.48 (/0.08 (/0.43 (/0.49 0.11 (/0.45 0.02 0.08 0.02 (/0.06 (/0.08 (/0.40 0.10 0.04 (/0.47 0.05 (/0.11 (/0.04 (/0.06 0.12 (/0.50 (/0.06 (/0.35 (/0.01 (/0.50 1.58 0.19 (/0.32 0.45 (/0.26 (/0.50 0.19 (/0.18 (/0.43 (/0.29 (/0.19 (/0.22 (/0.09 (/0.40 (/0.35 0.13 (/0.50 (/0.31 (/0.21 0.47 0.10 0.01 (/0.25 1.70 (/0.38 (/0.04 0.67 0.18 (/0.29 0.26 (/0.18 (/0.39 0.27 (/0.14 (/0.33 (/0.34 (/0.09 (/0.15 0.01 (/0.27 (/0.27 0.28 (/0.45 (/0.28 (/0.42 0.19 (/0.10 (/0.21 (/0.25 0.84 (/0.28 (/0.31 0.12 0.18 0.20 0.20 (/0.20 (/0.12 0.03 0.01 (/0.11 (/0.30 0.10 0.25 0.08 (/0.09 0.07 (/0.23 (/0.14 0.00 (/0.28 0.10 0.03 2.12 (/0.06 (/0.13 0.47 (/0.26 (/0.30 (/0.49 0.39 (/0.51 (/0.57 1.06 (/0.46 0.22 0.26 (/0.09 0.02 (/0.02 (/0.36 0.35 (/0.23 (/0.53 0.30 0.00 (/0.11 0.12 0.00 (/0.25 0.57 (/0.27 0.34 0.39 (/0.57 (/0.51 0.27 (/0.45 0.07 0.44 (/0.42 0.15 0.53 0.71 (/0.10 0.16 (/0.21 0.43 0.77 (/0.04 0.35 0.44 0.22 (/0.14 0.12 (/0.60 0.24 (/0.63 0.04 (/0.01 (/0.18 (/0.21 0.05 (/0.16 (/0.21 (/0.10 (/0.10 0.08 (/0.03 0.02 0.04 0.03 (/0.19 0.07 (/0.06 (/0.21 0.05 0.10 0.18 0.03 (/0.05 0.07 (/0.05 (/0.21 0.61 0.40 (/0.71 (/0.83 0.55 (/0.80 (/0.80 0.98 (/0.76 0.29 0.55 0.16 (/0.06 0.06 (/0.50 0.58 0.06 (/0.68 1.22 0.17 1.14 0.01 0.25 (/0.06 0.81 (/0.85 0.56 0.09 (/0.31 (/0.15 (/0.15 (/0.14 (/0.02 (/0.25 (/0.02 (/0.09 0.14 0.57 0.04 0.08 0.00 (/0.03 0.25 0.15 (/0.06 0.16 0.06 (/0.10 (/0.03 (/0.30 0.08 (/0.29 (/0.14 Bra CoW DeW HeG ShH MGr InW Urb TiL RoG Coa 0.77 (/0.20 (/0.23 (/0.23 (/0.23 (/0.23 (/0.22 (/0.23 (/0.19 (/0.06 0.25 (/0.08 (/0.01 (/0.19 (/0.11 0.05 (/0.23 (/0.20 (/0.18 (/0.23 (/0.15 0.13 (/0.23 (/0.19 (/0.23 (/0.08

Table 2 (Continued )

Species

Notiophilus substriatus Water. Olisthopus rotundatus (Payk.) Patrobus assimilis Chaud. Patrobus atrorufus (Stroem) Pterostichus adstrictus Esch. Pterostichus aethiops (Panz.) Pterostichus cristatus (Dufour) Pterostichus diligens (Sturm) Pterostichus madidus (F.) Pterostichus melanarius (Ill.) Pterostichus minor (Gyll.) Pterostichus niger (Schall.) Pterostichus nigrita (Payk.) Pterostichus rhaeticus Heer Pterostichus strenuus (Panz.) Pterostichus vernalis (Panz.) Pterostichus versicolor (Sturm) Stomis pumicatus (Panz.) Synuchus nivalis (Panz.) Trechus micros (Herbst) Trechus obtusus Erich. Trechus quadristriatus (Schrank) Trechus rubens (F.) Trechus secalis (Payk.) Trichocellus cognatus (Gyll.) Trichocellus placidus (Gyll.)

ECOGRAPHY 27:4 (2004)

(Goeze), C. melanocephalus (Linnaeus), Carabus violaceus Linnaeus, Cychrus caraboides Linnaeus, Leistus rufescens (Fabricius), Loricera pilicornis (Fabricius), Nebria brevicollis (Fabricius), Notiophilus biguttatus (Fabricius), Pterostichus madidus (Fabricius), P. niger (Schaller), Trechus obtusus Erichson) and tend to be found in both upland and lowland areas of the region (Luff 1998). The constrained ordination (CANOCO) biplot resulted in low eigenvalues and cumulative percentage variances, indicative of a noisy data set (Table 3). However, there were strong species/environment correlations with all three axes, which together accounted for 75% of the variance explained by the environmental data. The biplot of the first and second axes (Fig. 2), accounting for over 60% of this variance, shows the distribution of selected species relative to the major land cover variables. Upland species (e.g. Miscodera arctica , Trichocellus cognatus ) were found along the negative axis 1 with the shrub heath, heath grassland, bracken and bog variables. Lowland species (e.g. Pterostichus cristatus (Dufour), Harpalus rufipes ) were associated along the positive axis 1 with managed grassland and tilled land variables whilst Bradycellus verbasci (Duftschmid) was related to the urban variable, also on the positive axis 1. There was no obvious relationship between the negative axis 2 and cover types or species but the coastal variable was strongly related to the positive axis 2, as were a number of species, especially Amara spreta , A. lucida and Broscus cephalotes . Table 4 shows the correlation coefficients of the environmental variables with three axes of the constrained ordination. Table 5 shows the results of the automatic forward selection of the 12 land cover variables, the variance explained by each and their statistical significance. The tilled land variable was very strongly related to axis 1 whilst the coastal variable was associated with axis 2, with both explaining the same amount of variance. Axis 1 was also positively related to the urban, managed grassland, coastal and deciduous woodland variables, all generally lowland cover types, and negatively with shrub heath, heath grassland, bog and bracken, all upland covers. Of these cover types, shrub heath cover explained about half of that of tilled land and coastal but the others explained less because their associations with other cover types. The urban
Table 3. Eigenvalues, cumulative explained variance (%), cumulative species environment relationship (%) and species/ environment correlations, for the rst three constrained ordination axes. Axis 1 Eigenvalues Cumulative % variance Cumulative species/environment relationship Species environment correlations 0.402 4.2 35.7 0.890 Axis 2 0.280 7.1 60.5 0.853 Axis 3 0.167 8.8 75.3 0.813

variable was also related to the positive axis 2, opposite the negatively related managed grassland and tilled land covers. Axis 3 was related to differences between the upland covers of shrub heath cover and heath grassland.

Discussion
The results showed that species preferences for land cover variables differed considerably. A number of species were strongly associated with, for instance, the cover types of heath grassland and shrub heath and with the coastal variable whilst with a number of the more abundant species (Luff 1998), there were no strong relationships with any of the land cover variables. Within ground beetle species assemblage classifications covering a range of habitat and land cover types (Turin et al. 1991, Luff et al. 1992, Eyre et al. 1996, Turin 2000) some species were confined to one group or habitat type whilst others occurred in most of the groups in a classification. Major differences in species land cover preferences were shown between upland and lowland cover types using the satellite-derived cover data. This pattern was also observed within regional or national ground beetle assemblage habitat classifications (Luff et al. 1989, Eyre and Luff 1990a, Eyre and Luff 1990b). Differences between ground beetle morphology and life traits have been shown for species in the agricultural landscape (Ribera et al. 1999, 2001), associated with land cover disturbance. The strong relationship between a number of very habitat specific species, especially those of riverine sediments (Eyre et al. 2001), with some of the land cover variables, particularly managed grassland and tilled land, indicates a need for some care in interpretation of results. Sediment species were associated with particular land covers depending on where they were within catchments. The Landsat-derived land cover data used here with ground beetles has also been applied by Griffiths et al. (2000) in work on plant species distribution and by South et al. (2001) to predict mammal distribution in Britain. Other applications arising from the generation of satellite-derived cover data has included predictions of plant species diversity (Gould 2000, Wessels et al. 2000, Luoto et al. 2002a). Land cover change needs monitoring as does its effects on invertebrate species, for instance on butterflies (Maes and Van Dyck 2001). Kerr et al. (2001) and Luoto et al. (2002b) applied satellite land cover data to the assessment of butterfly species distribution over large areas whilst Pearlstine et al. (2002) used this type of cover data to investigate both butterfly and ant species distribution. Eyre et al. (2003b) have shown a strong relationship between British ground beetle species assemblages and satellite-derived land covers using data at the national 10 km grid square scale. These beetle data are usually used 423

ECOGRAPHY 27:4 (2004)

Fig. 2. Constrained ordination biplot showing the distribution of ground beetle species in relation to the eight major land cover variables (Am, Agonum marginatum ; Al, Amara lucida ; As, Amara spreta ; At, Amara tibialis ; Ap, Asaphidion pallipes ; Bb, Bembidion bruxellense ; Bm, Bembidion minimum ; Bco, Bradycellus collaris ; Br, Bradycellus rucollis ; Bv, Bradycellus verbasci ; Bce, Broscus cephalotes ; Cc, Cicindela campestris ; Cm, Calathus melanocephalus ; Cv, Cymindis vaporariorum ; Dg, Dyschirius globosus ; Dm, Dromius melanocephalus ; Hr, Harpalus rupes ; Ma, Miscodera arctica ; Mf, Metabletus foveatus ; Or, Olisthopus rotundatus ; Pa, Pterostichus adstrictus ; Pc, Pterostichus cristatus ; Tr, Trechus rubens ; Tc, Trichocellus cognatus ).

for atlas production (Luff 1998) but Telfer et al. (2002) showed that the information derived from national recording schemes can be applied at other scales, including the 1 km national grid square in a regional context. Blackburn and Gaston (2002) point out all scales are useful in providing information in macroecological studies with, for instance, the 1 km scale land
Table 4. Correlations between the land cover variables and the rst three axes of the constrained ordination. Cover variable Tilled land Coastal Shrub heath Urban Managed grassland Coniferous woodland Bracken Deciduous woodland Heath grassland Rough grass Bog Inland water Axis 1 0.674 0.333 (/0.526 0.446 0.398 (/0.110 (/0.292 0.304 (/0.533 0.084 (/0.345 0.104 Axis 2 (/0.337 0.736 0.205 0.315 (/0.367 (/0.058 0.202 (/0.302 0.018 0.157 (/0.002 (/0.079 Axis 3 0.081 (/0.135 0.573 0.019 (/0.035 (/0.046 0.248 0.160 (/0.440 (/0.141 (/0.277 (/0.011

cover data and ground beetle site data providing different insights than the 10 km pooled cover and beetle lists used by Eyre et al. (2003b). The ability to relate ground beetle species distribution to remote-sensed land cover data is likely to increase the possibility of using these species in monitoring broader environmental change. Satellite-derived land cover data are likely to be the only way to assess land cover and change at both the regional and national scale, as well as the only way to compare covers in different countries. The 1 km scale cover data employed in this work may have been appropriate for investigations such as those of Niemela et al. (2002) in assessing ground beetle species distribution across urban-rural gradients in three countries. The use of ground beetle data in wider environmental assessments and policy decisions has been advocated (McCracken and Bignal 1998, Niemela 1999, 2000) whilst their use in conservation assessments has also proved possible (Eyre et al. 1996, Eyre and Luff 2002). Satellite land cover data, with concomitant ground beetle survey work, may provide input into assessments of environmental change brought about by both climatic change and policy decisions.

Table 5. The additional variance explained by each land cover variable in the constrained ordination, the variance ratios (F) and the signicance (P), listed in order of automatic forward selection by CANOCO. Cover variable Tilled land Coastal Shrub heath Managed grassland Urban Coniferous woodland Bracken Deciduous woodland Heath grassland Rough grass Bog Inland water Additional variance explained 0.28 0.28 0.15 0.09 0.08 0.05 0.05 0.05 0.03 0.03 0.02 0.02 F 34.76 34.76 19.42 11.89 10.94 6.63 6.42 6.02 4.36 3.45 2.59 2.23 p B/0.005 B/0.005 B/0.005 B/0.005 B/0.005 B/0.005 B/0.005 B/0.005 B/0.005 B/0.010 B/0.030 B/0.025

References
Andersen, J. 2000. What is the origin of the carabid beetle fauna of dry, anthropogenic habitats in western Europe? / J. Biogeogr. 27: 795 /806. Anon. 1995. Countryside Information System, Ver. 5.23. / Dept of Environment, London. Desender, K., Ervynck, A. and Tack, G. 1999. Beetle diversity and historical ecology of woodlands in Flanders. / Belg. J. Zool. 129: 139 /155. Assmann, T. 1999. The ground beetle fauna of ancient and recent woodlands in the lowlands of north-west Germany (Coleoptera, Carabidae). / Biodiv. Conserv. 11: 1499 /1517. Basedow, T. 2002. Changes in agriculture in an area in Northern Germany between the years 1971 and 2000, and the reactions of populations of predatory carabids (Col., Carabidae), of other predators, and of cereal aphids, to these changes. / Z. Panzenkrank. Panzenschutz 109: 1 / 14.
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Blackburn, T. M. and Gaston, K. J. 2002. Scale in macroecology. / Global Ecol. Biogeogr. 11: 185 /189. Bonn, A., Hagen, K. and Wohlgemuth-von Reiche, D. 2002. The signicance of ood regimes for carabid beetle and spider communities in riparian habitats / a comparison of three major rivers in Germany. / River Res. Appl. 18: 43 / 64. Bra ndle, M., Durka, W. and Altmoos, M. 2000. Diversity of surface dwelling beetle assemblages in open-cast lignite mines in Central Germany. / Biodiv. Conserv. 9: 1297 / 1311. Brewster, C. C., Allen, J. C. and Kopp, D. D. 1999. IPM from space: using satellite imagery to construct regional crop maps for studying crop-insect interaction. / Am. Entom. 45: 105 /117. Buttereld, J. 1997. Carabid community succession during the forestry cycle in conifer plantations. / Ecography 20: 614 / 625. Buttereld, J. et al. 1995. Carabid beetle communities as indicators of conservation potential in upland forests. / For. Ecol. Manage. 79: 63 /77. Cassidy, K. M. et al. 2001. Using current protection status to assess conservation priorities. / Biol. Conserv. 97: 1 /20. Chust, G. et al. 2000. The effects of the landscape pattern on arthropod assemblages: an analysis of scale-dependence using satellite data. / Belg. J. Entomol. 2: 99 /110. Daily, G. C., Ehrlich, P. R. and Sanchez-Azofeifa, G. A. 2001. Countryside biogeography: use of human-dominated habitats by the avifauna of southern Costa Rica. / Ecol. Appl. 11: 1 /13. Daniel, M. et al. 1998. Predictive map of Ixodes ricinus highincidence habitats and a tick-borne encephalitis risk assessment using satellite data. / Exp. Appl. Acar. 22: 417 /433. Debinski, D. M., Kindscher, K. and Jakubauskas, M. E. 1999. Remote sensing and GIS-based model of habitats and biodiversity in the Greater Yellowstone ecosystem. / Int. J. Rem. Sens. 20: 3281 /3291. Dennis, P., Aspinall, R. J and Gordon, I. J. 2002. Spatial distribution of upland beetles in relation to landform, vegetation and grazing management. / Basic Appl. Ecol. 3: 183 /193. Dister, S. W. et al. 1997. Landscape characterization of peridomestic risk for Lyme disease using satellite imagery. / Am. J. Trop. Med. Hyg. 57: 687 /692. Donoghue, D. N. M. and Mironnet, N. 2002. Development of an integrated geographical information system prototype for coastal habitat monitoring. / Comp. Geosci. 28: 129 /141. Dreiser, U. 1994. Mapping of desert locust habitats in Africa using landsat thematic mapper data. / GeoJournal 32: 55 / 60. Elkaim, B. and Rybarczyk, H. 2000. Structure of the invertebrate community of the salt marshes of the Bay of Somme (eastern English Channel). / Cah. Biol. Marine 41: 295 / 311. Eyre, M. D. and Luff, M. L. 1990a. A preliminary classication of European grassland habitats using carabid beetles. / In: Stork, N. E. (ed.), The role of ground beetles in ecological and environmental studies. Intercept, Andover, pp. 227 /236. Eyre, M. D. and Luff, M. L. 1990b. An initial classication of the habitats of grassland ground beetles in the United Kingdom. / Entomol. Gaz. 41: 197 /208. Eyre, M. D. and Luff, M. L. 2002. The use of ground beetles (Coleoptera: Carabidae) in conservation assessments of exposed riverine sediment habitats in Scotland and northern England. / J. Insect Conserv. 6: 25 /38. Eyre, M. D., Lott, D. A. and Garside, A. 1996. Assessing the potential for environmental monitoring using ground beetles (Coleoptera: Carabidae) with riverside and Scottish data. / Ann. Zool. Fenn. 33: 157 /163. Eyre, M. D., Luff, M. L. and Phillips, D. A. 2001. The ground beetles (Coleoptera, Carabidae) of exposed riverine sediments in Scotland and northern England. / Biodiv. Conserv. 10: 403 /426.
ECOGRAPHY 27:4 (2004)

Eyre, M. D., Luff, M. L. and Woodward, J. 2003a. Grouse moor management: habitat and conservation implications or invertebrates in southern Scotland. / J. Insect Conserv. 7: 21 /32. Eyre, M. D. et al. 2003b. The relationship between British ground beetles (Coleoptera, Carabidae) and land cover. / J. Biogeogr. 30: 719 /730. Fuller, R. M., Groom, G. B. and Jones, A. R. 1994. The LandCover map of Great Britain / an automated classication of Landsat Thematic Mapper data. / Photogram. Engin. Rem. Sens. 60: 553 /562. Fuller, R. M. et al. 1998. The integration of eld survey and remote sensing for biodiversity assessment: a case study in the tropical forests and wetlands of Sango Bay, Uganda. / Biol. Conserv. 86: 379 /391. Gardner, S. M. 1991. Ground beetle (Coleoptera: Carabidae) communities on upland heath and their association with heathland ora. / J. Biogeogr. 18: 281 /289. Gould, W. 2000. Remote sensing of vegetation, plant species richness, and regional biodiversity hotspots. / Ecol. Appl. 10: 1861 /1870. Grifths, G. H., Lee, J. and Eversham, B. C. 2000. Landscape pattern and species richness; regional scale analysis from remote sensing. / Int. J. Rem. Sens. 21: 2685 /2704. Grilli, M. P. and Gorla, D. E. 1997. The spatio-temporal pattern of Delphacodes kuscheli (Homoptera: Delphacidae) abundance in central Argentina. / Bull. Entomol. Res. 87: 45 / 53. Groom, J. D. and Grubb, T. C. 2002. Bird species associated with riparian woodland in fragmented, temperate-deciduous forest. / Conserv. Biol. 16: 832 /836. Helenius, J. et al. 2001. Ground beetle (Coleoptera, Carabidae) diversity in Finnish arable land. / Agric. Food Sci. Fin. 10: 261 /276. Irmler, U. et al. 2002. Zonation of ground beetles (Coleoptera: Carabidae) and spiders (Araneida) in salt marshes at the North and the Baltic Sea and the impact of the predicted sea level increase. / Biodiv. Conserv. 11: 1129 /1147. Jepsen, J. U. et al. 2002. The importance of prey distribution in habitat use by arctic foxes (Alopex lagopus ). / Can. J. Zool. 80: 418 /429. Judas, M., Dornieden, K. and Strothmann, U. 2002. Distribution patterns of carabid beetle species at the landscape-level. / J. Biogeogr. 29: 491 /508. Jukes, M. R., Peace, A. J. and Ferris, R. 2001. Carabid beetle communities associated with coniferous plantations in Britain: the inuence of site, ground vegetation and stand structure. / For. Ecol. Manage. 148: 271 /286. Kerr, J. T., Southwood, T. R. E. and Cihlar, J. 2001. Remotely sensed habitat diversity predicts buttery species richness and community similarity in Canada. / Proc. Natl Acad. Sci. USA 98: 11365 /11370. Kielhorn, K. H., Keplin, B. and Huttl, R. F. 1999. Ground beetle communities on reclaimed mine spoil: effects of organic matter application and revegetation. / Plant Soil 213: 117 /125. Kinnunen, H., Tiainen, J. and Tukia, H. 2001. Farmland carabid beetle communities at multiple levels of spatial scale. / Ecography 24: 189 /197. Koivula, M. and Niemela , J. 2002. Boreal carabid beetles (Coleoptera, Carabidae) in managed spruce forests / a summary of Finnish case studies. / Silva Fenn. 36: 423 /436. Luff, M. L. 1996. Environmental assessments using ground beetles (Carabidae) and pitfall traps. / In: Eyre, M. D. (ed.), Environmental monitoring, surveillance and conservation using Invertebrates. EMS Publ., Newcastle upon Tyne, pp. 42 /47. Luff, M. L. 1998. Provisional atlas of the ground beetles (Coleoptera, Carabidae) of Britain. / Biological Records Centre, Huntingdon. Luff, M. L., Eyre, M. D. and Rushton, S. P. 1989. Classication and ordination of habitats of ground beetles (Coleoptera,

425

Carabidae) in north-east England. / J. Biogeogr. 16: 121 / 130. Luff, M. L., Eyre, M. D. and Rushton, S. P. 1992. Classication and prediction of grassland habitats using ground beetles (Coleoptera, Carabidae). / J. Environ. Manage. 35: 301 / 315. Luoto, M., Toivonen, T. and Heikkinen, R. K. 2002a. Prediction of total and rare plant species richness in agricultural landscapes from satellite images and topographic data. / Landscape Ecol. 17: 195 /217. Luoto, M., Kuussaari, M. and Toivonen, T. 2002b. Modelling buttery distribution based on remote sensing data. / J. Biogeogr. 29: 1027 /1037. Maes, D. and Van Dyck, H. 2001. Buttery diversity loss in Flanders (north Belgium): Europes worst case scenario? / Biol. Conserv. 99: 263 /276. Masek, J. G. 2001. Stability of boreal forest stands during recent climate change: evidence from Landsat satellite imagery. / J. Biogeogr. 28: 967 /976. McCracken, D. I. and Bignal, E. M. 1998. Applying the results of ecological studies to land-use policies and practices. / J. Appl. Ecol. 35: 961 /967. Moilanen, A. and Hanski, I. 1998. Metapopulation dynamics: effects of habitat quality and landscape structure. / Ecology 79: 2503 /2515. Moncayo, A. C., Edman, J. D. and Finn, J. T. 2000. Application of geographic information technology in determining risk of eastern equine encephalomyelitis virus transmission. / J. Am. Mosquito Cont. Ass. 16: 28 /35. Moore, J. D. et al. 2002. Effects of two silvicultural practices on soil fauna abundance in a northern hardwood forest, Quebec, Canada. / Can. J. Soil Sci. 82: 105 /113. Muldavin, E. H., Neville, P. and Harper, G. 2001. Indices of grassland biodiversity in the Chihuahuan Desert ecoregion derived from remote sensing. / Conserv. Biol. 15: 844 /855. Niemela , J. 1999. Ecology and urban planning. / Biodiv. Conserv. 8: 119 /131. Niemela , J. 2000. Biodiversity monitoring for decision-making. / Ann. Zool. Fenn. 37: 307 /317. Niemela , J. and Baur, B. 1998. Threatened species in a vanishing habitat: plants and invertebrates in calcareous grasslands in the Swiss Jura mountains. / Biodiv. Conserv. 7: 1407 /1416. Niemela , J. et al. 2002. Carabid beetle assemblages (Coleoptera, Carabidae) across urban-rural gradients: an international comparison. / Landscape Ecol. 17: 387 /401. Pearlstine, L. G. et al. 2002. Assessing state-wide biodiversity in the Florida Gap analysis project. / J. Environ. Manage. 66: 127 /144. Pena, N. M. de la. et al. 2003. Landscape context and carabid beetles (Coleoptera: Carabidae) communities of hedgerows in western France. / Agr. Ecosyst. Environ. 94: 59 /72. Petit, C. C. and Lambin, E. F. 2002. Impact of data integration technique on historical land- use/land-cover change: comparing historical maps with remote sensing data in the Belgian Ardennes. / Landscape Ecol. 17: 117 /132.

Radeloff, V. C., Mladenoff, D. J. and Boyce, M. S. 2000. The changing relation of landscape patterns and jack pine budworm populations during an outbreak. / Oikos 90: 417 /430. Ribera, I. et al. 2001. Effect of land disturbance and stress on species traits of ground beetle assemblages. / Ecology 82: 1112 /1129. Ribera, I. et al. 1999. A comparative study of the morphology and life traits of Scottish ground beetles (Coleoptera, Carabidae). / Ann. Zool. Fenn. 36: 21 /37. So derstrom, B. et al. 2001. Plants, insects and birds in seminatural pastures in relation to local habitat and landscape factors. / Biodiv. Conserv. 10: 1839 /1863. South, A. B. et al. 2001. Reintroduction of the European beaver (Castor ber) to Norfolk, UK: a preliminary modelling analysis. / J. Zool. 254: 473 /479. Telfer, M. G., Preston, C. D. and Rothery, P. 2002. A general method for measuring relative change in range size from biological atlas data. / Biol. Conserv. 107: 99 /109. Ter Braak, C. J. F. and Smilauer, P. 1998. CANOCO reference manual and users guide to Canoco for Windows: software for Canonical Community Ordination (Ver. 4). / Centre for Biometry, Wageningen. Townsend, P. A. and Walsh, S. J. 2001. Remote sensing of forested wetlands: application of multitemporal and multispectral satellite imagery to determine plant community composition and structure in southeastern USA. / Plant Ecol. 157: 129 /149. Turin, H. 2000. De Nederlandse Loopkevers. Verspreiding en Oecologie [The Netherlands ground beetles. Distribution and ecology]. / KNNV Uitgeverij, Utrecht. Turin, H. et al. 1991. Ecological characterisation of carabid species (Coleoptera, Carabidae) in the Netherlands from thirty years of pitfall sampling. / Tijds. Entomol. 134: 279 / 304. Va sa nen, R. and Helio vaara, K. 1994. Assessment of insect occurrence in boreal forests based on satellite imagery and eld measurements. / Acta For. Fenn. 243: 1 /39. Velazquez, A. et al. 2001. Effects of landscape changes on mammalian assemblages at Izta- Popo Volcanoes, Mexico. / Biodiv. Conserv. 10: 1059 /1075. Vessby, K. et al. 2002. Species-richness correlations of six different taxa in Swedish seminatural grasslands. / Conserv. Biol. 16: 430 /439. Walker, D. A. 2000. Hierarchical subdivision of Arctic tundra based on vegetation response to climate, parent material and topography. / Glob. Change Biol. 6: 19 /34. Wessels, K. J., Reyers, B. and Jaarsveld, A. S. van 2000. Incorporating land cover information into regional biodiversity assessments in South Africa. / Anim. Conserv. 3: 67 /79. Wittmann, F., Anhuf, D. and Junk, W. J. 2002. Tree species distribution and community structure of central Amazonian varzea forests by remote-sensing techniques. / J. Trop. Ecol. 18: 805 /820.

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