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Home , Ballota nigra, Centaurea nigra, Centaurea scabiosa, Filipendula ulmaria, Knautia arvensis, Melilotus altissimus

APPENDIX 1. Table 1. Sources of meta-data for analysis of bumblebee forage preferences

No. of Top 5 forage Top 5 forage plants Site Year of forage (Descending order) (Descending order) No. Author study Paper / Report title Site Location Habitat types plants Long-tongued * Short-tongued * 1 Carvell, C 1999 Habitat use and conservation of bumblebees (Bombus Salisbury Plain Unimproved 20 Trifolium pratense vernus spp.) under different grassland management regimes Training Area, calcareous Trifolium pratense Wiltshire grassland Echium vulgare nigra Lotus corniculatus Echium vulgare Stachys officinalis arvensis 2 Carvell, C 2000a Studies of the distribution and habitat requirements of Castlemartin Range, Unimproved 8 minor Rhinanthus minor and other bumblebees at Pembrokeshire, SW mesotrophic and Trifolium pratense Lotus corniculatus Castlemartin Range, Pembrokeshire. Wales coastal grassland Centaurea nigra officinalis Melilotus officinalis Trifolium repens Rubus caesius Centaurea nigra 3 Carvell, C 2000b Studies of the distribution and habitat requirements of Kenfig NNR, Mesotrophic and 22 Trifolium pratense Rubus caesius Bombus sylvarum and other bumblebees at Kenfig Glamorgan, S Wales dune grassland Anthyllis vulneraria repens National Nature Reserve, Glamorgan Rhinanthus minor Thymus polytrichus Rubus caesius Rhinanthus minor Echium vulgare Echium vulgare 4 Dicks, L. 1999 The structure and functioning of plant-insect Shelfanger, Norfolk Ancient hay 13 Centaurea nigra Centaurea nigra visitor communities. meadow Lathyrus pratensis Leucanthemum vulgare Cirsium palustre Cirsium palustre Trifolium repens Hypochaeris radicata Lotus corniculatus Lotus corniculatus 5 Dicks, L. 1999 The structure and functioning of plant-insect flower Hickling Broad NNR, Restored hay 12 Lotus corniculatus Lotus corniculatus visitor communities. Norfolk meadow Lathyrus pratensis Lathyrus pratensis Vicia cracca Vicia cracca Leucanthemum vulgare Leucanthemum vulgare Vicia sativa Hypochaeris radicata 6 Hughes, L. 1998 The Great Yellow Bumblebee, Bombus distinguendus ; South Uist, Western Machair dune 11 Trifolium pratense Trifolium repens aspects of habitat use, phenology and conservation onIsles, Scotland grasslands Trifolium repens Senecio jacobaea the Machair of the Outer Hebrides. Vicia cracca Odontites vernus Prunella vulgaris Trifolium pratense Senecio jacobaea Leontodon autumnalis 7 Macdonald, M. 1998 Observations on foraging machair on S. Uist and Western Is les, Machair dune 9 Anthyllis vulneraria Trifolium repens Barra (unpublished) Scotland grasslands Trifolium pratense Anthyllis vulneraria Trifolium repens Potentilla palustris Vicia cracca Trifolium pratense Rhinanthus minor Rhinanthus minor 8 Macdonald, M. 1997 Feeding ecology of some Bombus and Psithyrus Strathpeffer, Highland Garden and 63 Pulmonaria spp. Borago officinalis bumblebees in northern Scotland. Region, Scotland farmland Vicia sepium Pulmonaria spp. Napeta x faassenii Rubus fruticosus Salvia officinalis Chamerion angustifolium Digitalis purpurea Rubus idaeus

21 Appendices Restoration and management of bumblebee habitat on arable farmland: literature review BD 1617

No. of Top 5 forage plants Top 5 forage plants Site Year of forage (Descending order) (Descending order) No. Author study Paper / Report title Site Location Habitat types plants Long-tongued * Short-tongued * 9 Pickett, H 2000 Bumblebee (Bombus) abundance, diversity and Manor Farm, nr Arable farmland, 7 Centaurea cyanus Centaurea cyanus activity in relation to field margin habitat type Malton, North sown field margins Rhinanthus minor Cirsium arvense Yorkshire Cirsium arvense Rhinanthus minor Lotus corniculatus Prunella vulgaris Leucanthemum vulgare 10 Prys-Jones 1978 Ecological studies of foraging and life history in Wicken Fen, Fen meadow 34 Symphytum officinale Centaurea nigra bumblebees Cambridgeshire pratensis Salix cinerea Cirsium palustre Symphytum officinale Centaurea nigra Cirsium palustre Lythrum salicaria Eupatorium cannabinum 11 Saville, NM 1990 The relationship between numbers, bee forage Madingley Wood, Woodland 24 hirsutum repens species and woodland attributes through the season: Cambridgeshire Ajuga reptans Epilobium hirsutum Madingley Wood case study. In 'Bumblebee ecology Cirsium vulgare Cirsium arvense in woodland and arable farmland' Prunella vulgaris Rubus caesius Rubus caesius Cirsium vulgare 12 Saville, NM 1990 Comparative survey of farm woodlands. In Croxton, West Woodland and 21 Carduus acanthoides Rubus fruticosus 'Bumblebee ecology in woodland and arable farmland' Cambridgeshire arable farmland Rubus fruticosus Carduus acanthoides Epilobium hirsutum Epilobium hirsutum Cirsium arvense Cirsium arvense Odontites vernus Cirsium vulgare 13 Williams, PH 1982 Why are there so many species of bumblebees at Dungeness, Kent Dune ridge 13 Teucrium scorodonia Teucrium scorodonia Dungeness? grasslands Echium vulgare Rubus fruticosus Melilotus altissima Echium vulgare Rubus fruticosus Melilotus altissima Dipsacus fullonum Centaurea nigra 14 Williams, PH 1983 Why are there so many species of bumblebees at Shoreham, Kent Arable farmland and 17 nigra Convolvulus arvensis Dungeness? woodland edge Medicago sativa Chamerion angustifolium Rubus fruticosus Origanum vulgare Chamerion angustifolium Clematis vitalba Cirsium vulgare Medicago sativa

* Note that some forage plant species listed here may not have been included in the final analysis because they were only visited at one site.

APPENDIX 1. Table 2. Details of plant ecological characteristics compared with adjusted bumblebee preference indices.

22 Appendices Restoration and management of bumblebee habitat on arable farmland: literature review BD 1617

Ecological characteristics Brief description (see source for full details) Source * Missing Type of data (continuous/ records discontinuous variables)

Taxonomy- Family name Family name e.g. ECPE 0 Discontinuous

Life history Plant life cycles summarised as: Ann (summer or winter annual); Bien (biennial) or Per (monocarpic or Kleyer 0 Discontinuous polycarpic perennial) Flower type Flower type: ACT= Actinomorphic (flower with radial symmetry); ZYG= Zygomorphic (flower with Stace 0 Discontinuous bilateral symmetry) Flower colour Dominant colour of flower Stace 0 Discontinuous

Flower arrangement & Direction of flower entrance pointing:- 1= downwards; 2= upwards in a head or clustered; 3= upwards in P-J 0 Discontinuous orientation a head, composite; 4= upwards, single flower; 5= horizontally; 6= horizontally or downwards; 7= horizontally, down or upwards; 8= horizontally or upwards

Habitat abundance- Arable Abundance in arable habitats: ++ very common and characteristic; + common within habitat; . ECPE 7 Discontinuous Widespread in habitat; - infrequent and uncharacteristic; -- largely absent from habitat

Period of flowering Period of flowering: SPRIS starting in spring, flowering period <3 months; SPRIL starting in spring, Kleyer 0 Discontinuous flowering period >3 months; ESUMS starting in early summer, flowering period <3 months; ESUML starting in early summer, flowering period >3 months; SUMAU flowering in late summer or autumn.

Earliest Flowering time Earliest month of flowering (value 1-12) Ecoflora 0 Continuous

Latest Flowering time Latest month of flowering (value 1-12) Ecoflora 0 Continuous

Length flowering period Number of months in flower (value 2-7) Ecoflora/ Stace 0 Continuous

Floristic diversity Associated floristic diversity: 1= 10 species or fewer; 2= 10.1-14 species; 3= 14.1-18 species; 4= 18.1- ECPE 7 Continuous 22 species; 5= >22 species Height (cm) Plant height: Log transformed for analysis Stace 0 Continuous

Corolla length mm Flower corolla length measured as the part of the corolla tube that can be probed by a bumblebees' B&P (most) or 5 Continuous proboscis (mm) W Change in frequency 78-98 Change in the frequency of that plant species in UK Countryside Survey plots between 1978 and 1998 CS 7 Continuous

* Data sources: B&P (Barrow & Pickard, 1985); CS (Countryside Survey data); ECPE (Hodgson et al., 1995); Ecoflora (Fitter & Peat, 1994); Kleyer (Kleyer, 1995); P-J (Prys-Jones, 1982); Stace (Stace, 1997); W (Williams, 1989a).

23 Appendices Restoration and management of bumblebee habitat on arable farmland: literature review BD 1617 Table 3. Bumblebee visits and adjusted preference indices for forage plant species visited at two or more sites.

tongued tongued tongued* - tongued * - - - index index index ong number visits L Short Short All bumblebees Long Number of sites Mean number visits Mean number visits Adjusted preference Adjusted preference Adjusted preference

Forage Species Common name All bumblebees Mean Echium vulgare Viper's bugloss 3 54.67 4.07 33.33 3.47 21.33 3.02 Ballota nigra Black horehound 2 123.50 3.85 122.50 3.56 1.00 0.89 Centaurea nigra Black knapweed 7 115.57 3.82 22.29 2.66 93.29 3.32 Epilobium hirsutum Great willowherb 4 42.25 3.75 30.25 3.14 12.00 2.65 Rubus fruticosus agg. Bramble 6 56.00 3.47 22.17 2.45 33.83 3.29 Cirsium palustre Marsh thistle 3 59.33 3.36 45.00 2.79 14.33 2.19 Melilotus altissimus Tall melilot 2 29.50 3.31 19.50 2.57 10.00 2.31 Trifolium pratense Red clover 8 65.63 3.31 58.50 3.26 7.13 0.91 Carduus acanthoides Welted thistle 3 32.33 3.02 20.00 2.51 12.33 1.93 Odontites vernus Red 5 27.80 2.92 12.40 2.43 15.40 1.70 Lotus corniculatus Bird's-foot trefoil 9 125.22 2.89 11.67 2.23 113.56 1.86 Ononis repens Common restharrow 2 18.50 2.87 4.50 1.87 14.00 1.87 Rhinanthus minor Yellow rattle 7 20.71 2.85 12.43 2.39 8.29 1.65 Cirsium arvense Creeping thistle 8 19.13 2.85 9.00 1.69 10.13 2.47 Cirsium vulgare Spear thistle 8 23.25 2.82 16.25 2.24 7.00 1.90 Mentha aquatica Water mint 2 12.00 2.80 10.50 2.26 1.50 1.09 Leucanthemum vulgare Oxeye daisy 3 29.00 2.79 1.67 0.97 27.33 2.28 Anthyllis vulneraria Kidney vetch 3 23.00 2.73 19.00 2.48 4.00 1.39 Rubus caesius Dewberry 4 20.75 2.68 7.75 2.08 13.00 2.00 Rosa canina Dog rose 2 16.00 2.67 8.00 1.02 8.00 2.42 Vicia cracca Tufted vetch 6 26.83 2.65 15.17 2.44 11.67 0.53 Convolvulus arvensis Field bindweed 3 76.33 2.65 5.33 0.85 71.00 2.76 Chamerion angustifolium Rosebay willowherb 7 22.57 2.50 9.29 1.45 13.29 2.27 Lathyrus pratensis Meadow vetchling 5 46.40 2.38 11.40 2.21 35.00 0.75 Trifolium repens White clover 10 25.70 2.36 10.90 1.64 14.80 1.79 Senecio jacobaea Common ragwort 5 19.00 2.29 6.20 0.70 12.80 2.34 Ajuga reptans Bugle 2 15.00 2.17 13.00 2.08 2.00 0.89 Dipsacus fullonum Wild teasel 3 11.33 2.00 9.33 1.71 2.00 0.95 Prunella vulgaris Selfheal 7 10.71 2.00 9.14 1.80 1.57 0.57 Hypochaeris radicata Cat's-ear 2 10.50 1.99 1.00 0.97 9.50 1.30 Stachys sylvatica Hedge woundwort 3 6.00 1.89 6.00 1.81 0.00 0.19 Field scabious 2 7.50 1.81 1.00 0.47 6.50 1.25 Melilotus officinalis Ribbed melilot 2 11.50 1.73 2.50 1.16 9.00 1.38 Glechoma hederacea Ground-ivy 2 5.50 1.70 5.00 1.33 0.50 0.74 Lychnis flos-cuculi Ragged-Robin 2 6.50 1.62 5.00 1.64 1.50 0.19 ulmaria Meadowsweet 3 5.00 1.55 0.00 -0.05 5.00 1.74 Arctium minus Lesser burdock 2 5.50 1.49 5.50 1.35 0.00 0.14 Iris pseudacorus Yellow iris 2 2.00 1.36 2.00 1.05 0.00 0.40 Clinopodium vulgare Wild basil 2 2.00 1.27 2.00 1.30 0.00 0.00 Lamiastrum galeobdolon Yellow archangel 2 3.50 1.25 0.00 -0.20 3.50 1.53 Trifolium dubium Lesser trefoil 2 4.50 1.19 0.50 0.76 4.00 0.50 Solanum dulcamara Bittersweet 2 1.50 1.09 1.50 0.95 0.00 0.28 Bryonia dioica White bryony 3 1.33 1.03 1.33 0.74 0.00 0.40 Meadow buttercup 2 2.00 0.78 0.50 0.76 1.50 0.01 * Long-tongued species = B. hortorum, B. pascuorum, B. humilis, B. ruderarius, B. sylvarum, B. muscorum. Short-tongued species = B. terrestris, B. pratorum, B. lapidaries, B. lucorum, B. jonellus

24 Appendices Restoration and management of bumblebee habitat on arable farmland: literature review BD 1617 APPENDIX 1. Table 4. Ecological characteristics and flower attributes of bumblebee forage pla nts visited at two or more sites. (see Table 2 for key to abbreviations)

98

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Forage Species Family name Life history Flower type ACT/ ZYG Flower colour Flower arrangement & orientation Abundance in arable habitats Period of flowering Earliest Flowering time Latest Flowering time Length flowering period Floristic diversity Height (cm) Log for analysis Corolla length mm Change in frequency 78 Ajuga reptans Per ZYG Blue 5 -- SPRIL 5 7 3 4 30 8.60 -14 Anthyllis vulneraria Per ZYG Yellow 2 -- ESUML 6 9 4 4 60 9.70 -1 Arctium minus Asteraceae Bien ACT Purple 3 . SUMAU 7 9 3 3 200 3.50 -1 Ballota nigra Lamiaceae Per ZYG Purple 2 SUMAU 6 10 5 100 8.70 9 Bryonia dioica Cucurbitaceae Per ACT White 7 ESUML 5 9 5 300 -6 Carduus acanthoides Asteraceae Bien ACT Purple 3 - SUMAU 6 8 3 3 120 3.14 Centaurea nigra Asteraceae Per ACT Purple 3 -- ESUML 6 9 4 4 60 3.00 -62 Chamerion angustifolium Onagraceae Per ZYG Pink 5 -- SUMAU 7 9 3 2 120 2.40 -19 Cirsium arvense Asteraceae Per ACT Purple 3 + SUMAU 7 9 3 3 90 1.20 -8 Cirsium palustre Asteraceae Bien ACT Purple 3 -- SUMAU 7 9 3 4 150 1.30 -14 Cirsium vulgare Asteraceae Bien ACT Purple 3 . SUMAU 7 10 4 4 150 5.60 -65 Clinopodium vulgare Lamiaceae Per ZYG Pink 5 -- SUMAU 7 9 3 4 80 1 Convolvulus arvensis Convolvulaceae Per ACT White 8 ++ ESUML 6 9 4 3 75 5.90 9 Dipsacus fullonum Dipsacaceae Bien ZYG Purple 5 ESUMS 7 8 2 150 9.60 -2 Echium vulgare Boraginaceae Bien ZYG Blue 5 ESUML 6 9 4 90 6.43 1 Epilobium hirsutum Onagraceae Per ACT Pink 4 -- ESUMS 7 8 2 2 150 3.30 -9 Filipendula ulmaria Per ACT White 2 -- ESUML 6 9 4 2 120 -26 Glechoma hederacea Lamiaceae Per ZYG Blue 5 -- SPRIS 3 5 3 2 30 11.40 -8 Hypochaeris radicata Asteraceae Per ACT Yellow 3 - ESUML 6 9 4 3 60 6.20 Iris pseudacorus Iridaceae Per ZYG Yellow 5 -- ESUML 5 11 7 1 150 6.60 1 Knautia arvensis Dipsacaceae Per ZYG Blue 2 - SUMAU 7 9 3 4 100 5.00 Lamiastrum galeobdolon Lamiaceae Per ZYG Yellow 5 -- ESUMS 4 5 2 1 60 7.20 -1 Lathyrus pratens is Fabaceae Per ZYG Yellow 5 - ESUMS 5 8 4 4 120 4.80 -45 Leucanthemum vulgare Asteraceae Per ACT White 3 . ESUML 6 8 3 4 100 0.00 -13 Lotus corniculatus Fabaceae Per ZYG Yellow 5 -- ESUML 5 10 6 4 35 5.09 -36 Lychnis flos-cuculi Caryophyllaceae Per ACT Pink 8 -- ESUMS 5 6 2 4 75 5.86 -2 Melilotus altissimus Fabaceae Bien ZYG Yellow 5 ESUML 6 8 3 150 3.64 Melilotus officinalis Fabaceae Bien ZYG Yellow 5 ESUMS 7 9 3 150 3.64 Mentha aquatica Lamiaceae Per ZYG Purple 2 -- SUMAU 7 10 4 2 90 4.10 -10 Odontites vernus Ann ZYG Pink 5 + ESUML 6 8 3 4 50 10 Ononis repens Fabaceae Per ZYG Pink 8 -- ESUMS 6 9 4 4 60 6.33 -2 Prunella vulgaris Lamiaceae Per ZYG Purple 5 . ESUMS 6 9 4 5 30 7.50 -33 Ranunculus acris Per ACT Yellow 8 . SPRIL 5 10 6 4 60 0.00 -14 Rhinanthus minor Scrophulariaceae Ann ZYG Yellow 5 -- ESUML 5 8 4 5 50 6.30 6 Rosa canina Rosaceae Per ACT White 8 ESUMS 6 7 2 300 0.00 Rubus caesius Rosaceae Per ACT White 8 -- ESUMS 6 9 4 2 50 1.40 Rubus fruticosus agg. Rosaceae Per ACT White 8 - ESUMS 5 9 5 1 200 1.40 -3 Senecio jacobaea Asteraceae Per ACT Yellow 8 - ESUMS 6 10 5 4 150 2.91 6 Solanum dulcamara Solanaceae Per ACT Purple 8 -- ESUMS 6 9 4 1 200 3 Stachys sylvatica Lamiaceae Per ZYG Purple 8 . SUMAU 7 8 2 2 100 9.40 1 Trifolium dubium Fabaceae Ann ZYG Yellow 2 + ESUML 5 10 6 5 25 2.80 -8 Trifolium pratense Fabaceae Per ZYG Pink 2 ++ ESUMS 5 9 5 4 100 9.20 -60 Trifolium repens Fabaceae Per ZYG White 2 + ESUML 6 9 4 3 50 2.80 -69 Vicia cracca Fabaceae Per ZYG Purple 5 . ESUMS 6 8 3 2 200 5.70 5

25 Appendices Restoration and management of bumblebee habitat on arable farmland: literature review BD 1617 Table 5. Results of ANOVA tests on differences in adjusted bumblebee preference indices between discontinuous plant ecological characteristics. Values represent the mean number of visits to each category. * p<0.05. ** p<0.01. ***p<0.001

Ecological Long- Short- characteristic n All bees SE tongued SE tongued SE Family name Asteraceae 9 34.85 11.30 14.10 4.64 20.75 9.38 Boraginaceae 1 54.67 33.33 21.33 Caryophyllaceae 1 6.50 5.00 1.50 Convolvulaceae 1 76.33 5.33 71.00 Cucurbitaceae 1 1.33 1.33 0.00 Dipsacaceae 2 9.42 1.91 5.16 4.17 4.25 2.25 Fabaceae 10 37.68 11.20 15.36 5.21 22.32 10.50 Iridaceae 1 2.00 2.00 0.00 Lamiaceae 8 22.28 14.50 21.02 14.60 1.26 0.41 Onagraceae 2 32.41 9.84 19.77 10.50 12.65 0.65 Ranunculaceae 1 2.00 0.50 1.50 Rosaceae 4 24.44 11.00 9.48 4.62 14.96 6.50 Scrophulariaceae 2 24.26 3.55 12.42 0.02 11.85 3.56 Solanaceae 1 1.50 1.50 0.00

F 13,30 2.14 1.04 4.27 Significance * ns *** Life history Annual 3 17.67 6.90 8.44 3.97 9.23 3.32 Biennial 8 28.43 7.06 18.93 5.07 9.50 2.41 Perennial 33 29.16 6.10 13.14 3.95 16.03 4.66

F 2,41 0.72 1.50 0.43 Significance ns ns ns Flower type Actinomorphic 19 28.49 6.90 10.99 2.78 17.50 5.67 Zygomorphic 25 28.06 6.64 16.06 5.06 12.00 4.54

F 1,42 0.00 1.67 4.11 Significance 0.97 ns * Flower colour Blue 4 20.67 11.50 13.08 7.20 7.58 4.76 Pink 7 26.46 8.25 17.42 7.72 9.05 2.36 Purple 13 34.38 11.30 22.48 8.87 11.91 6.94 White 8 28.76 9.02 7.14 2.53 21.62 8.10 Yellow 12 24.82 9.89 7.22 2.11 17.60 9.11

F 4,39 0.73 2.35 1.20 Significance ns ns ns Flower arrangement Upwards (U), clustered 8 33.35 14.70 27.86 15.10 5.49 1.53 & orientation Upwards, Composite 8 36.83 12.60 15.09 5.14 21.74 10.60 Upwards, single flower 1 42.25 30.25 12.00 Horizontal (H) 16 25.95 7.64 10.51 2.03 15.44 6.95 H, U or downw ards 1 1.33 1.33 10.00 H or U 10 22.26 7.80 6.70 1.88 15.56 6.96

F 5,38 1.60 1.11 1.33 Significance ns ns ns Arable habitats Infrequent 6 28.62 8.05 10.30 3.77 18.33 5.17 Largely absent 18 27.83 8.69 11.18 2.77 16.65 7.59 Widespread 7 14.76 4.26 7.75 2.32 7.01 3.76 Common 4 19.28 5.26 8.20 2.66 11.08 2.64 V Common, cht 2 70.98 5.35 31.92 26.60 39.07 31.90

F 4,32 0.59 0.15 1.22 Significance ns ns ns Flowering period ESUML 15 36.72 10.20 10.22 2.57 26.50 9.28 ESUMS 15 23.76 5.10 12.76 3.86 11.00 2.74 SPRIL 2 8.50 6.50 6.75 6.25 1.75 0.25 SPRIS 1 5.50 5.00 0.50 SUMAU 11 28.46 10.70 22.46 10.70 6.01 1.74

F 4,39 0.87 0.26 1.25 Significance ns ns ns

26 Appendices Restoration and management of bumblebee habitat on arable farmland: literature review BD 1617

a) Short-tongued bumblebees

30 Centaurea nigra Actinomorphic

x r = -0.89 ** Zygomorphic e Rubus fruticosus agg. d n i Echium vulgare e

c 20 n e

r Convolvulus arvensis

e Epilobium hirsutum f e

r Cirsium arvense p

Melilotus altissimus

d 10 Cirsium palustre e

t Rubus caesius Lotus corniculatus s Trifolium repens

u Rhinanthus minor j Melilotus officinalis Anthyllis vulneraria d Knautia arvensis Ballota nigra

A Trifolium dubium Glechoma hederacea Ranunculus acris 0 Stachys sylvatica 0 5 10 Flower corolla length (mm)

b) Long-tongued bumblebees

Ballota nigra 35 Actinomorphic Echium vulgare

x Zygomorphic

e r = 0.58 ns

d 30 n

i Trifolium pratense

e 25 Epilobium hirsutum c n e

r 20 e

f Cirsium palustre e

r Centaurea nigra

p 15

Vicia cracca Anthyllis vulneraria

d Rubus fruticosus agg. Rhinanthus minor

e Lathyrus pratensis t 10 Rubus caesius Ajuga reptans s Ononis repens Stachys sylvatica u

j Cirsium arvense Prunella vulgaris

d Arctium minus Glechoma hederacea Le5ucanthemum vulgare

A Convolvulus arvensis Senecio jacobaea Lamiastrum galeobdolon 0 0 5 10 Flower corolla length

APPENDIX 1. Figure 1. Relationships between flower corolla length and adjusted forage plant preference index for a) short-tongued and b) long-tongued bumblebees. Flower types (actinomorphic or zygomorphic) are also shown.

27 Appendices Restoration and management of bumblebee habitat on arable farmland: literature review BD 1617 APPENDIX 2. Table 1. Bumblebee forage plants showing a significant change in frequency from Countryside Survey data between 1978 and 1998. Values represent percentage change in recorded frequency from CS plots. Results of Mc Nemar chi-square test: * p<0.05. ** p<0.01. *** p<0.001

Bumblebee distribution region a

Universal Forage plant species GB p Northern p N. local p Southern p S. local p Ajuga reptans -53.85 * Calluna vulgaris -53.85 *** -54.39 *** -76.19 ** Calystegia sepium 2000.00 *** 1600.00 *** 900.00 * Centaurea nigra -59.05 *** -56.14 *** -62.50 *** -64.86 ** Cirsium palustre 250.00 * -67.74 ** -72.41 ** Cirsium vulgare -37.57 *** -34.94 * -40.00 ** -48.21 ** -80.00 * -100.00 * Digitalis purpurea 153.33 ** Erica tetralix -52.11 *** -55.07 *** -70.59 *** Filipendula ulmaria -40.63 * -72.22 * Heracleum sphondylium -49.06 *** -58.21 *** -42.39 ** -43.64 * Hypochoeris spp./Leontodon spp. -21.95 * Lamium purpureum -54.76 ** -60.00 * Lathyrus pratensis -57.69 *** -53.19 ** -64.52 ** Leucanthemum vulgare -68.42 * -90.00 * Lotus corniculatus -41.38 ** -54.29 * -53.57 * Odontites vernus 200.00 * 7.00 * Papaver rhoeas -90.00 * Potentilla reptans -41.67 * Prunella vulgaris -30.00 * -33.78 * Ranunculus repens -25.64 ** -24.03 * -27.62 * Rhinanthus minor 6.00 * 6.00 * Rubus spp. 96.30 *** 46.15 * 53.19 * Taraxacum agg. -30.77 * Trifolium pratense -55.56 *** -61.67 *** -47.92 * -52.78 * Trifolium repens -31.94 *** -47.37 *** -40.68 * Ulex spp. -71.43 *** -68.75 * -73.68 ** -77.78 ** Vicia sepium -50.91 ** -51.43 * Viola tricolor -90.00 * -100.00 * a See Appendix 3 for description of bumblebee distribution regions

28 Appendices Restoration and management of bumblebee habitat on arable farmland: literature review BD 1617 APPENDIX 2. Table 2. Significant changes in CS plot cover represented by bumblebee forage plants, grouped by aggregate vegetation class and plot type. Results of modified paired t-test: * p<0.05. ** p<0.01. *** p<0.001

Bumblebee Mean% change in distribution Aggregate vegetation Total no. of proportion of region class a CS Plot type plots forage cover Significance N.local All All 236 -3.85 * Southern All All 219 -2.73 * Universal GB All All 1572 -2.21 *** Northern All Fields/unenclosed 382 -2.96 ** Universal GB All Fields/unenclosed 865 -2.48 *** Southern All Hedges 41 -9.68 * N.local All Roadsides 25 -8.12 * Northern Heath/Bog All 104 -9.58 *** Universal GB Heath/Bog All 221 -7.92 *** N.local Heath/Bog All 113 -6.02 * Northern Heath/Bog Fields/unenclosed 95 -9.95 *** Universal GB Heath/Bog Fields/unenclosed 195 -8.00 *** Universal GB Infertile grassland Fields/unenclosed 192 -3.03 * S.local Lowland wooded All 59 -10.47 ** Universal GB Lowland wooded All 119 -8.64 ** S.local Lowland wooded Hedges 40 -11.18 * Universal GB Lowland wooded Hedges 79 -8.00 *

a see Bunce et al.(1999a).

29 Appendices Restoration and management of bumblebee habitat on arable farmland: literature review BD 1617

100% 90% 80% 70% 60% 50% 40% 30% 20% 10% % changes in forage plant frequency 0% Universal GB Southern n=62 Southern local Northern n=57 Northern local n=69 n=55 n=27

Bumblebee distribution region

Increased plants Decreased plants

APPENDIX 2. Figure 1. Total number of changes in forage plant frequency between 1978 and 1998, expressed as the percentage of increased and decreased plants detected across the bumblebee distribution regions.

30 Appendices Restoration and management of bumblebee habitat on arable farmland: literature review BD 1617

a) Short-tongued bumblebees

30 Centaurea nigra Rubus fruticosus agg. x e d n i Echium vulgare r = -0.04 ns e 20 c

n Convolvulus arvensis e r Epilobium hirsutum e f

e Cirsium arvense r Senecio jacobaea p Leucanthemum vulgare

d 10 Cirsium palustre

e Cirsium vulgare Ononis repens t Trifolium repens Lotus corniculatus s Filipendula ulmaria Odontites vernus

u Anthyllis vulneraria j Mentha aquatica

d Ballota nigra Trifolium pratense Prunella vulgaris Vicia cracca

A Stachys sylvatica Ranunculus acris Lathyrus pratensis 0 -70 -60 -50 -40 -30 -20 -10 0 10 Change in frequency 1978-1998

b) Long-tongued bumblebees

Ballota nigra 35 Echium vulgare

x r = -0.03 ns e 30

d Trifolium pratense n i 25 Epilobium hirsutum e c n

e 20 r

e Cirsium palustre f Centaurea nigra e

r 15

p Rubus fruticosus agg. Rhinanthus minor d Cirsium vulgare Lotus corniculatus Mentha aquatica

e 10 Lathyrus pratensis Ajuga reptans t Stachys sylvatica s Prunella vulgaris Trifolium repens Lychnis flos-cuculi u j 5 Glechoma hederacea d Trifolium duSbeiunmecio jacobaea

A Filipendula ulmaria 0 -70 -60 -50 -40 -30 -20 -10 0 10 Change in frequency 1978-1998

APPENDIX 2. Figure 2. Relationships between change in forage plant frequency from the Countryside Survey 1978 – 1998 and adjusted preference index for a) short-tongued and b) long-tongued bumblebee species. (r = linear regression coefficient)

31 Appendices Restoration and management of bumblebee habitat on arable farmland: literature review BD 1617

APPENDIX 3.

Bumblebee distribution regions used for analyses of changes in forage plant abundance.

These range descriptions are based on updates of the Bumblebee Distribution Maps Scheme (Alford,1980) and patterns identified by Williams (1982). Some have been compiled by the Bees, Wasps and Ants Recording Society and published in recent Newsletters, and others are based on current knowledge of the International Bee Research Association, Edwards (pers. comm.) and Macdonald (pers. comm.). Most represent the known post-1970 to present distribution of each bumblebee (Bombus) species.

The species fall into six groups as detailed below:

Universal (occurs throughout the British Isles) B. lucorum B. terrestris B. pratorum B. lapidarius B. pascuorum B. hortorum

Universal, local (occurs throughout British Isles but is restricted in abundance, even within main area of distribution) B. jonellus B. muscorum This group was not distinguished from the universal group for the analysis of changes in plant species abundance.

Northern (restricted to, or with a strong bias to the north of the Humber-Mersey line) B. monticola (not included in current dataset)

Northern, local (restricted to the extreme north of Scotland; Western Isles, Coll, Tiree, Caithness, Sutherland and Orkney) B. distinguendus

Southern (restricted to, or with a strong bias to the south of the Humber-Mersey line) B. ruderarius

Southern, local (restricted to the southern coastal band running south from Norfolk in the east and up to the Isle of Anglesey in the west, having seriously declined since 1970’s) B. humilis B. sylvarum

32 Appendices Restoration and management of bumblebee habitat on arable farmland: literature review BD 1617 APPENDIX 4. Table 1. Consideration of bumblebee forage plants for inclusion in seed mixtures.

Adjusted Preference Performance Soil Commercial Forage Species Common name index index a types b availability Ajuga reptans Bugle 2.17 N Anthyllis vulneraria Kidney vetch 2.73 0.14 C Y Arctium minus Lesser burdock 1.49 C Y Ballota nigra Black horehound 3.85 N Bryonia dioica White bryony 1.03 N Campanula rotundifolia Harebell 0.00 NCW Y Carduus acanthoides Welted thistle 3.02 N Centaurea cyanus Cornflower NC Y Centaurea nigra Black knapweed 3.82 0.34 NCAW Y Greater knapweed -0.03 C Y Chamerion angustifolium Rosebay willowherb 2.50 N Cirsium arvense Creeping thistle 2.85 NCA N Cirsium palustre Marsh thistle 3.36 CW Y Cirsium vulgare Spear thistle 2.82 NC N Clinopodium vulgare Wild basil 1.27 NCA Y Convolvulus arvensis Field bindweed 2.65 N Dipsacus fullonum Wild teasel 2.00 NW Y Echium vulgare Viper's bugloss 4.07 A Y Epilobium hirsutum Great willowherb 3.75 N Filipendula ulmaria Meadowsweet 1.55 0.16 CW Y Geranium pratense Meadow cranesbill NCW Y Glechoma hederacea Ground-ivy 1.70 A N Hypochaeris radicata Cat's-ear 1.99 0.46 CAW Y Iris pseudacorus Yellow iris 1.36 W Y Knautia arvensis Field scabious 1.81 0.09 NC Y Lamiastrum galeobdolon Yellow archangel 1.25 N Lamium album White deadnettle N Lathyrus pratensis Meadow vetchling 2.38 0.22 CW Y Leucanthemum vulgare Oxeye daisy 2.79 0.79 NCAW Y Linaria vulgaris Common toadflax A Y Lotus corniculatus Bird's-foot trefoil 2.89 0.45 NCAW Y Lotus pedunculatus Greater bird's-foot trefoil W Y Lychnis flos-cuculi Ragged-Robin 1.62 0.26 CW Y Malva moschata Musk mallow NCA Y Melilotus altissimus Tall melilot 3.31 N Melilotus officinalis Ribbed melilot 1.73 N Mentha aquatica Water mint 2.80 W Y Odontites vernus Red bartsia 2.92 NC Y Onobrychis viciifolia Sainfoin C Y Ononis repens Common restharrow 2.87 C Y Spiny restharrow C Y Origanum vulgare Wild marjoram C Y Prunella vulgaris Selfheal 2.00 0.53 NCAW Y Ranunculus acris Meadow buttercup 0.78 0.18 NCAW Y Rhinanthus minor Yellow rattle 2.85 0.06 NCAW Y Rosa canina Dog rose 2.67 N Rubus caesius Dewberry 2.68 N Rubus fruticosus agg. Bramble 3.47 N Senecio jacobaea Common ragwort 2.29 N Solanum dulcamara Bittersweet 1.09 N Stachys officinalis Betony 0.07 NCAW Y Stachys palustris Marsh woundwort W Y Stachys sylvatica Hedge woundwort 1.89 NCA Y Succisa pratensis Devil's-bit scabious 0.03 CAW Y Teucrium scorodonia Wood sage CA Y Trifolium dubium Lesser trefoil 1.19 0.25 NCAW Y Trifolium pratense Red clover 3.31 0.32 NCAW Y Trifolium repens White clover 2.36 NCAW Y Vicia cracca Tufted vetch 2.65 0.15 NCW Y Vicia sativa Common vetch NCAW Y a Index of performance in existing restoration schemes (Pywell et al., 1999) b Most suited soil types of commercially available species N, neutral; C, calcareous; A, acidic; W, wet.

33 Appendices Restoration and management of bumblebee habitat on arable farmland: literature review BD 1617

APPENDIX 4. Table 2. Recommended wildflower and grass seed mixtures to benefit bumblebees on field margins or grassland sites of specific soil types. NB. Flowering phenology may vary according to local climate and management.

Flowering Phenology Bumblebees

Spring- Late Early Mid Summer- Short- Long- Summer Summer Autumn tongued tongued NEUTRAL/ LOAMY SOILS Wild a Centaurea nigra + + + + Geranium pratense + + + Knautia arvensis + + + + Leucanthemum vulgare + + + Lotus corniculatus + + + + + Prunella vulgaris + + + Ranunculus acris + + + + Rhinanthus minor + + + + Trifolium pratense + + + + Vicia cracca + + + +

Grasses Agrostis capillaris Anthoxanthum odoratum Cynosurus cristatus Festuca ovina Festuca rubra spp. commutata Trisetum flavescens

CALCAREOUS SOILS Wild flowers Anthyllis vulneraria + + + Centaurea nigra + + + + Centaurea scabiosa + + + + Clinopodium vulgare + + + Knautia arvensis + + + + Leucanthemum vulgare + + + Lotus corniculatus + + + + + Malva moschata + + Ononis repens + + + + Origanum vulgare + + + Prunella vulgaris + + + Ranunculus acris + + + + Rhinanthus minor + + + + Trifolium pratense + + + + Vicia cracca + + + +

Grasses Agrostis capillaris Cynosurus cristatus Festuca ovina Festuca rubra spp. commutata Phleum bertolonii Trisetum flavescens

34 Appendices Restoration and management of bumblebee habitat on arable farmland: literature review BD 1617 APPENDIX 4. Table 2 cont.

Flowering Phenology Bumblebees

Spring- Late Early Mid Summer- Short- Long- Summer Summer Autumn tongued tongued ACIDIC/ SANDY SOILS Wild flowers Centaurea nigra + + + + Echium vulgare + + + + Leucanthemum vulgare + + + Linaria vulgaris + + + Lotus corniculatus + + + + + Malva moschata + + Prunella vulgaris + + + Rhinanthus minor + + + + Stachys officinalis + + + Trifolium pratense + + + + Vicia sativa + + + +

Grasses Agrostis capillaris Anthoxanthum odoratum Cynosurus cristatus Deschampsia flexuosa Festuca ovina Festuca rubra spp. juncea

WET SOILS Wild flowers Centaurea nigra + + + + Filipendula ulmaria + + + Lotus pedunculatus + + + + Lychnis flos-cuculii + + + Prunella vulgaris + + + Ranunculus acris + + + + Stachys officinalis + + + Stachys palustris + + Succisa pratensis + + + + Trifolium pratense + + + +

Grasses Alopecurus pratensis Anthoxanthum odoratum Deschampsia cespitosa Festuca rubra spp. juncea a Sowing rates: Seed merchants generally provide advice on sowing rates which are in the region of 80% grasses to 20% flowers depending on cost factors.

35 Appendices Restoration and management of bumblebee habitat on arable farmland: literature review BD 1617

APPENDIX 4. Table 3. Forage plant species to encourage in hedgerows and farm woodlands, not commercially available.

HEDGEROW/ TREE SPECIES Flowering Phenology Bumblebees Spring- Late Early Mid Summer- Short- Long- Summer Summer Autumn tongued tongued Ajuga reptans + + Ballota nigra + + + Bryonia dioica + + + + Carduus acanthoides + + + + Chamerion angustifolium + + + + Cirsium arvense * + + + Cirsium vulgare * + + + + Convolvulus arvensis + + + Cotoneaster spp. + + + Dipsacus fullonum + + + Epilobium hirsutum + + + Glechoma hederacea + + + Lamiastrum galeobdolon + + + Lamium album + + + Melilotus altissimus + + + Melilotus officinalis + + + Rosa canina + + + Rubus caesius + + + Rubus fruticosus agg. + Salix cinerea + + + Senecio jacobaea * + + + Solanum dulcamara + + + +

* These species are invasive which require control under the Injurious Weeds Act 1959.

36 Appendices Restoration and management of bumblebee habitat on arable farmland: literature review BD 1617