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The Effect of Field Boundary Type on the Community Structure, Spatial The effect of field boundary type on the community structure, spatial distribution and physiological condition of overwintering arthropods, with special reference to Carabidae and Staphylinidae (Coleoptera). by . Georgianne J.K. Griffiths A thesis submitted to the University ofPlymouth in partial fulfilment for the degree of Doctor of Philosophy ----- - - -------- ------- ! d .. I r:::. r: . .,. / r· .:: t::~ ' \/: , <\ , 'T 11 U r'll \, I.... I ..> I I I 1.,.... • I ..... • ' • ..., ..., l ,-. •- ' D l I 1\ '. f I) 1;- t -.~ , • • .;._ ij .- .,. ~ f ··~,· .- \ '-'..._ .... ~- · "'i' ..., _ J nr) r. ·-·y 1 Ll c. tr\i1 University ofPiymouth at Seale-Hayne In collaboration with The Game Conservancy Trust September 2003 11 The effect of field boundary type on the community structure, spatial distribution and physiological condition of overwintering arthropods, with special reference to Carabidae and Staphylinidae (Coleoptera). Georgianne J. K. Griffiths ABSTRACT The potential of different field boundary types in lowland farmland to contribute to arthropod biodiversity and sustainable agriculture was investigated. Field boundaries, categorised according to nationally applicable definitions, were found to represent ecologically differing habitats based on their woody abundance and the frequency of young and mature emergent trees. These habitat characteristics were determining factors in the community structure and composition of overwintering epigeal arthropods. Hedgerows supported the most species rich carabid and staphylinid assemblage. Degraded hedgerow boundaries supported the most equitable carabid community, and provided a refuge for carabid species with poor dispersal power to a greater extent that hedgerows or post and wire fences. The grassy and natural regeneration vegetation associated with post and wire boundaries supported high densities of all taxa particularly overwintering carabid and staphylinid polyphagous predators. A subset of all field boundary types was required for complete species representation, indicating that maximising the heterogeneity of field boundary habitats represented at the farm-scale will enhance arthropod biodiversity in farmland. Carabidae and StaphyHnidae actively selected overwintering sites and the physiological condition of polyphagous predators was generally high. It was concluded that heterogeneous distributions in field boundaries were more likely to be the result of differential microhabitat selection rather than differential survival overwinter. This indicated that favourable overwintering microhabitats occurred in all field boundary types. Generally, overwintering survival did not appear to be a regulating factor in the population dynamics of polyphagous predators. Margins adjacent to pre-existing boundaries may contribute to enhanced densities and physiological condition of some polyphagous predators, both over winter and in early spnng. The results were discussed in relation to field boundary management and agri­ environment policy. 111 CONTENTS CHAPTER ONE: INTRODUCTION 1 1.1 Field boundaries in lowland farmland 1 1.1.1 Classification and stock 1 1.1 .2 Origin 2 1.1.3 Function 5 1.1.4 Policy 5 1.2 Field boundaries and artbropod communities 6 1.2.1 Arthropod habitat preferences and dispersal ability 7 1.2.2 Botanical composition 9 1.2.3 Habitat structure and microclimate 13 1.2.4 Management 16 1.2.5 Field boundary network 19 1.2.6 Landscape structure 23 1.3 Field boundaries as overwintering babitat for polyphagous predators 24 1.3.1 Conservation biological control 25 1.3.2 Polyphagous predators 26 1.3.3 Factors influencing polyphagous predator survival overwinter 29 1.3. 4 Overwintering habitats of polyphagous predators 31 1.4 Aims 37 CHAPTER TWO: DESCRIPTION OF FIELD BOUNDARY TYPES 40 2.1 Introduction 40 2.1.1 Aims 47 2.2 Metbods 47 2.2.1 Field surveys 47 2.2.2 Analysis 52 2.3 Results 53 2.3.1 Woody composition 53 2.3.2 Herbaceous composition 57 2.3.3 Habitat condition 65 2.3.4 Structure 68 2.3.5 Factor analysis 71 2.4 Discussion 75 CHAPTER THREE: COMMUNITY STRUCTURE AND 78 COMPOSffiON OF ARTHROPODS OVERWINTERING IN DIFFERENT FIELD BOUNDARY TYPES AT THE FARM-SCALE 3.1 Introduction 78 3. 1. 1 Community structure and taxonomic composition 79 IV 3.1.2 Functional composition of Carabidae and Staphylinidae 80 3.1.3 Field boundary factors influencing arthropod communities 83 3.1.4 Field boundary classification based on Carabidae and Staphylinidae 84 3.1.5 Farm-scale representation ofCarabidae and Staphylinidae 84 3.1.6 Aims 86 3.2 Methods 86 3.2.1 Field survey 86 3.2.2 Analysis 88 3.3 Results 91 3.3.1 Community structure and taxonomic composition 91 3.3.2 Functional composition of Carabidae and Staphylinidae 106 3.3.3 Field boundary factors influencing arthropod communities 110 3.3.4 Field boundary classification based on Carabidae and Staphylinidae 121 3.3.5 Farm-scale representation ofCarabidae and Staphylinidae 123 3.4 Discussion 126 CHAPTER FOUR: THE DISTRIBUTION OF OVERWINTERJNG 131 COLEOPTERA IN DIFFERENT FIELD BOUNDARY FEATURES 4.1 Introduction 131 4.1.1 Aims 134 4.2 Methods 135 4.2.1 Field work 135 4.2.2 Analysis 140 4.3 Results 143 4.3.1 Habitat characteristics 143 4.3.2 Density and richness 149 4.3.3 Coleopteran composition in relation to features 159 4.3.4 Habitat characteristics influencing composition 161 4.4 Discussion 171 CHAPTER FIVE: THE PHYSIOLOGICAL CONDIDON OF 175 CARABIDAE OVERWINTERJNG IN DIFFERENT FIELD BOUNDARY TYPES 5.1 Introduction 175 5.1.1 Aims 179 5.2 Methods 179 5.2.1 Field surveys 179 5.2.2 Analysis 183 5.3 Results 184 5.4 Discussion 193 CHAPTER SIX: OVERWINTERING MICROHABITAT SELECTION 198 BY CARABIDAE AND STAPHYLINIDAE IN HEDGEROW AND GRASSY MARGIN V 6.1 Introduction 198 6.1.1 Aims 203 6.2 Methods 203 6.2.1 Marking 203 6.2.2 Field boundary sites 206 6.2.3 Sampling 211 6.2.4 Analysis 212 6.3 Results 213 6.3.1 Marking 213 6.3.2 Sampling 213 6.3 .3 Microhabitat of arenas 216 6.3.4 Spatial distribution of overwintering species 222 6.3.5 Distance moved 234 6.4 Discussion 236 CHAPTER SEVEN: DISCUSSION AND CONCLUSIONS 242 REFERENCES 249 APPENDIX ONE: PUBLICATION 277 VI LIST OF TABLES 1.1.1 Definitions of field boundary types. 3 1.1.2 Estimates offield boundary stock in England and Wales. 4 2.1.1 Indicator species of field boundary herbaceous communities. 44 2.2.1 Description of landscape, structural and botanical variables recorded 50 at each field boundary. 2.3.1 Relative abundance (%) of woody species present in a 30m sample 54 length offield boundary. 2.3.2 Herbaceous species presence-absence recorded from both sides of 59 a 10m sample length of each field boundary. 2.3.3 Frequency of occurrence of indicator species. 64 2.3.4 Landscape, structural and botanical characteristics of each field 69 boundary. 2.3.5 Mean landscape, structural and botanical characteristics of each field 70 boundary type. 2.3.6 Factor loadings for each field boundary characteristic and curnmulative 73 variance explained by each factor. 2.3.7 Mean scores for Factors 1 to 4 for each field boundary type. 74 2 3.3.1 Density (m- ) of higher arthropod taxa emerging from overwintering 95 at each field boundary. 2 3.3.2 Density (m- ) ofColeoptera families emerging from overwintering 96 at each field boundary. 2 3.3.3 Density (m- ) ofCarabid species emerging from overwintering at each 97 field boundary. 2 3.3.4 Density (m- ) ofStaphylinid taxa emerging from overwintering at each 99 field boundary. 3.3.5 Rarefraction estimates of richness (SR) for each taxonomic group, 105 with observed abundance (N) and richness (S) values, for each field boundary type. 3.3.6 Partial-CCA examining variation in community structure: summary 114 inertia and eigenvalues for each taxonomic group. 3.3.7 Taxon codes for partial-CCA ordination plots. 116 4.2.1 Description of features sampled in each field boundary-margin 137 combination. 4.2.2 Description of microhabitat characteristics recorded at each feature 139 location. 4.3.1 Microhabitat characteristics recorded at features within PW-M 144 boundaries. 4.3.2 Microhabitat characteristics recorded at features within PW+M 145 boundaries. 4.3.3 Microhabitat characteristics recorded at features within H-M 146 boundaries. 4.3.4 Micro habitat characteristics recorded at features within H+M 147 Vll boundaries. 2 4.3.5 In PW-M: results ofGLM on density (0.04m- ) and richness of 151 Coleoptera, Staphylinidae, Carabidae and B. lampros from five features at two depths. 2 4.3.6 In PW+M: results ofGLM on density (0.04m- ) and richness of 152 Coleoptera, Staphylinidae, Carabidae and B. lampros from six features at two depths. 2 4.3.7 In H-M: results ofGLM on density (0.04m- ) and richness of 153 Coleoptera, Staphylinidae, Carabidae and B. lampros from nine features at two depths. 2 4.3.8 In H+M: results ofGLM on density (0.04m- ) and richness of 154 Coleoptera, Staphylinidae, Carabidae and B. /ampros from ten features at two depths. 4.3.9 CCA ofColeoptera composition explained by microhabitat 165 characteristics: eigenvalues and variation. 4.3.10 CCA ofCarabidae composition explained by microhabitat 166 characteristics: eigenvalues and variation. 4.3.11 CCA ofStaphylinidae composition explained by microhabitat 167 characteristics: eigenvalues and variation. 5.3.1 Results of the GLM performed on fat content(%) and relative fresh 186 2 weight (mg.mm- ) for B. lampros, L. pilicornis, P. cupreus and P. strennus. 5.3.2 Temperatures eq at each boundary-margin combination and GLM 190 statistics. 5.3.3 Results ofGLM on abundance of B. /ampros and P. cupreus emerging 192 from post and wire and hedgerow boundaries, both with and without margins, in early and late spring. 5.3.4 Results oflinear regression analysis of mean abundance of B.
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