Mix & Max - SETACsesss05.setac.eu/embed/sesss05/7_Felix_Wackers... · environment and society as a...

53
Mix & Max Optimizing agro-ecosystem composition to maximize pollination and biocontrol services

Transcript of Mix & Max - SETACsesss05.setac.eu/embed/sesss05/7_Felix_Wackers... · environment and society as a...

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Mix & Max

Optimizing agro-ecosystem composition

to maximize pollination and biocontrol services

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Agriculture and the Environment

On the landscape level: • Larger fields • Loss of non-crop elements

On the field level: • Fewer crop varieties • Use of agrochemicals

Loss of Biodiversity

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Agriculture Environment

Focus on conservation

Agri Environment Schemes

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Agriculture Environment

Focus on ecosystem services

Agri Environment Schemes

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• Pest control • Disease control • Pollination • Climate regulation

• Erosion regulation • Nutrient retention • Water regulation & purification • …..

Regulating ecosystem services

Human well-being

Cultural ecosystem services, e.g: • Recreational and aesthetic values • Cultural heritage • Education and inspiration • Spiritual and religious values • …..

Cultural ecosystem services

Provisioning ecosystem services, e.g: • Food, fiber, fuel production • Clean water • Biochemicals, pharmaceuticals • Genetic resources • …..

Provisioning ecosystem services

Functional AgroBiodiversity

Ecosystem services to agriculture Positive spin-off to the regional and global

environment and society as a whole €320 billion/year €90 billion/year

(Constanza 1997)

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How to optimize services? Altieri, M.A. 1994 Biodiversity and Pest Management in Agroecosystems. Haworth Press, Inc., NY.

“Biodiversity in agroecosystems performs a variety of ecological services, including regulation of insect pests, pathogens and weeds.”

Traditional paradigm: Enhance diversity (diversity = services)

Biodiversity Ecosystem Services

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Andow D.A. (Ann. Rev. Entomol. 36: 561-586) Vegetational diversity and arthropod population response

• 52% of published studies on agroecosystem diversification

report reduced pest populations • 33% had no effect or variable effects • 15% resulted in increased pest populations.

Effects variable and unpredictable

Biodiversity Ecosystem Services

Does it work?

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Meteorus autographae

Does biodiversity composition matter?

Do bird conservation margins

benefit biological pest control?

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Results

overall sugar levels in Meteorus autographae

0

5

10

15

20

25

30

35

40

control bobwhite cahaba vetch

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Conclusions

High diversity field margins for bobwhite quail conservation failed to provide food to a biological control agent and did not enhance biological pest control in the adjacent crop. Parasitoids did clearly benefit from pure stands of cahaba white vetch. Ecosystem services delivery is not a function of biodiversity per se, but rather of plant species composition (resource availability)

Olson and Wackers, 2007 J. Appl. Ecol. :13-21

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Resource Bottleneck: Lack of nectar and pollen in many cropping systems

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0

5

10

15

20

25

30

5 10 15 20 25 30 35 40

age wasp [days]

aver

age

num

ber p

aras

itize

d ca

terp

illars Control 1 (N=6)

Control 2 (N=5)

Nectar 1 (N=6)

Nectar 2 (N=6)

Winkler et al., (2006)

Consumer benefits The impact of nectar sources on biocontrol efficacy

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Biological control agents depending on nectar/pollen feeding.

Type Plant-

feeding

stage

Arthropod examples can be found within: Type of plant

food utilised

Reference

Life-history

omnivory

adult Neuroptera:

Diptera:

Hymenoptera:

Coleoptera:

Chrysopidae (green lacewings)

Syrphidae (hoverflies)

Cecidomyiidae (gall midges)

Tachinidea (parasitoid flies)

Ichneumonidae, Braconidae,

a.o. (parasitoid wasps)

Vespidae (social wasps)

Formicidae (ants)

Meloidae (blister beetles)

nectar, pollen

nectar, pollen

nectar

nectar

nectar

nectar

nectar, fruit

nectar

nectar, pollen

(Stelzl 1991)

(Hickman et al. 1995)

(Opit 1997)

(Gilbert and Jervis 1998)

(Jervis 1998)

(Lewis 1998)

(Cuautle and Rico-Gray 2003)

(Beattie 1985)

juvenile Heteroptera: Pentatomidae (stink bugs) plant-juice

Temporal

omnivory

adult Hymenoptera:

Coleoptera:

Ichneumonidae, Braconidae,

a.o. (host feeding parasitoids)

Cicindelidae (tiger beetles)

nectar

seeds

(Jervis 1998)

(Wackers 2003)

(Zerm and Adis 2001)

juvenile Araneae: Araneidae (orb web spiders) pollen (Smith and Mommsen 1984)

Permanent

omnivory

adult &

juvenile

Acari:Mesostigmat

Heteroptera:

Neuroptera:

Thysanoptera:

Coleoptera:

Phytoseiidae

(predatory mites)

Pentatomidae (stink bugs)

Miridae (mirid bugs)

Geocorinae (big-eyed bugs)

Anthocoridae (flower bugs)

Chrysopa, Hemerobiidae

(brown lacewings)

Aeolothripidae, Phlaeothripidae

Coccinellidae (ladybirds)

Carabidae (ground beetles)

nectar

pollen

plant juice

plant juice

plant juice

pollen

nectar, pollen

leaves, pollen

nectar

pollen

seeds

(van Rijn and Tanigoshi 1999a)

(van Rijn and Tanigoshi 1999b)

(Ruberson et al. 1986)

(Gillespie and McGregor 2000)

Eubanks & Styrsky, this vol.

Eubanks & Styrsky, this vol.

(Stelzl 1991)

(McEwen et al. 1993)

(Kirk 1997) (Pemberton and Vandenberg

1993)

(Cottrell and Yeargan 1998)

(Goldschmidt and Toft 1997)

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How to optimize services?

1. Identify resource requirement of target organisms providing services 2. Identify plant species that are effective in providing these resources

Different organisms have different requirements

Targeted landscape management

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Informed selection of non-crop plants as a multifunctional tool

to optimize ecosystem services

- Select plants that optimize biological pest control or pollination

- Select plants that avoid stimulation of pests - Select plants that generate multiple ecosystem services

Targeted landscape management

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Selecting plants that optimize biological pest control

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Aegopodium podagraria

Daucus carota

Origanum vulgare

Galium mollugo Leucanthemum vulgare

Erigeron annuus

Trifolium repens Medicago lupulina Trifolium pratense

Achillea millefolium

Vicia sativa

Accessible

nectar

Inaccessible

nectar

Att

ract

ive

Non-a

ttra

ctiv

e

Select to optimize BC benefits

(Wäckers, 2004)

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Flower exploitation by predator group

Choice Longevity (AFLI)

family species

Floral Nectar

depth

Hoverfly

E. balteatus

Hoverfly

E. balteatus

Lacewing

C. carnea Parasitoids

References parasitoids

(species)

Apiaceae Ammi majus 0 + + + - Geneau et al., unpubl. (Microplitis mediator)

Apiaceae Coriandrum sativum 0 + + +/- Vattala et al., 2006 (Microtonus hyperodae)

Apiaceae Daucus carota 0 +/- + ++ + Winkler et al., 2009 (Cotesia glomerata)

Apiaceae Foeniculum vulgare 0 + + + Winkler et al., 2009 (Cotesia glomerata)

Apiaceae Heracleum spondylium 0 +/- + +/- Winkler et al., 2009 (Cotesia glomerata)

Apiaceae Pastinaca sativa 0 + + ++ +/- Foster & Ruessink, 1984 (Meteorus rubens)

Polygonaceae Fagopyrum esculentum 0 + + + + Winkler et al., 2009 (Cotesia glomerata)

Boraginaceae Borago officinalis 0 - + ++ - Nilsson et al., unpubl. (Trybliographa rapae)

Ranunculaceae Ranunculus acris 0 + + - Kehrli & Bacher, 2008 (Minotetrastichus frontalis)

Caryophyllaceae Gypsophila elegans 1 +/- + ++ Asteraceae Matricaria chamomilla 1 +/- + + - Nilsson et al., unpubl. (Trybliographa rapae)

Asteraceae Achillea millefolium 1 +/- + +/- - Wäckers 2004 (Cotesia glomerata)

Asteraceae L Cichorium intybus 1 - - + Asteraceae Chrysanthemum segetum 2 +/- + + Asteraceae Anthemis tinctoria 2 - +/- +/- Asteraceae Leucanthemum vulgare 2 +/- + - Wäckers 2004 (Cotesia glomerata)

Asteraceae Tanacetum vulgare 2 - - +/- Asteraceae Calendula officinalis 3 - - Rahat et al., 2005 (Trissolcus basalis)

Asteraceae Centaurea cyanus (+EFN) 3 +/- + ++ +/- Winkler et al., 2009 (Cotesia glomerata)

Asteraceae Helianthus annuus (+EFN) 3 + + Asteraceae Cosmos bipinnatus 4 - - +/- + Rahat et al., 2005 (Trissolcus basalis)

Malvaceae Malva sylvestris 4 - - - Boraginaceae Phacelia tanacetifolia 4 +/- +/- +/- - Irvin et al., 2007 (Gonatocerus spp.)

Fabaceae Medicago sativa 4 - - - Kehrli & Bacher, 2008 (Minotetrastichus frontalis)

Fabaceae Vicia sativa (+EFN) 4 - + ++ Geneau et al., unpubl. (Microplitis mediator)

Fabaceae Lotus corniculatus 4 - -

(Wäckers and van Rijn, 2012)

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Taking it to the field

Large scale biodiversity project in the Hoekse Waard working with conventional growers. Addition of annual and perennial field margins to existing landscape features (polders, dikes, creeks, canal borders).

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Aphid natural enemies in the adjacent brassica crop

Impact of flower strips on

in aardappel, 15 juni - 4 juli

0.00 0.01 0.02 0.03 0.04

Gaasvlieg

Zweefvlieg

Roofwants

Lieveheersbeest

Galmug

Sluipwesp

(aantal/blad)

met bloemstrokenzonder bloemstroken

flower strips control

Lacewings

Hoverflies

Predatory bugs

Lady bugs

Gall midges

Parasitic wasps

Number per leaf

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Melige Koolluis(aantal/plant)

0

20

40

60

80

TuinboonBloemen

Veld nabij akkerrand met:Cabbage aphids

Control Vetch Mixed flowers

24 26 28 30 32 34 36

week

Control

Broad bean

Mixed flowers

Brussels Sprouts

Cabbage aphids

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Potato Aphid peak: mid July

Wheat mid June

bladluizen

0

2

4

6

8

2005 2006 2007

aant

al/h

alm A

BCD

bedrijf

rover/luis

0.0

0.1

0.2

0.3

0.4

2005 2006 2007

verh

oudi

ng

01234567

2005 2006 2007

no./l

eaf

ABCD

farm

aphids

predator/prey

0.0

0.2

0.4

0.6

0.8

1.0

2005 2006 2007

ratio

Three year trends in aphid and natural

enemy populations at conventional farms

aphids aphids

Predator/prey

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Selecting plants that avoid stimulation of pests

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0

2

4

6

8

10

12

14

16

18

20

Contro

l (wate

r)

Centau

rea ja

cae

Origan

um vu

lgare

Ornitho

pus s

ativu

s

Medica

go sp

.

Trifolium

prate

nse

Centau

rea cy

anus

Daucu

s caro

ta

Anethu

m grav

eolen

s

Fagop

yrum es

culen

tum

surv

ival

(day

s)

Pieris rapaeCotesia glomerata

Pest

Biological control

agent

(Winkler et al., 2009)

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Selecting plants that generate multiple ecosystem services

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Pol

linat

or m

ix

Bio

cont

rol m

ix

Com

bina

tion

(Campbell, Biesmeijer Varma & Wäckers, 2012)

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Conclusions

• Insects providing ecosystem services differ distinctly with regard to their flower associations

• Flower mixes targeting particular insect groups are not necessarily suitable for other beneficials • Visitation by pollinators and biocontrol agents can be preserved when plants are combined Informed choice of flowering vegetation allows us to - optimize specific ecosystem services - avoid benefits to pests - successfully stack ecosystem services

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Optimizing Ecosystem Services in Terms of Agronomy and Conservation

(ECOSTAC.CO.UK)

Growing Wild

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Seed selection

- Database of >100 plants - Categorized by 16 criteria

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Flower species mix

Tailor made - to match grower´s requirements - to maximize ecosystem services

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Acknowledgements

US Robert Bugg George Heimpel Joe Lewis Dawn Olson Switzerland Claire Bonifay Agata Filipini Stephanie Horn Christoph Isenegger Nora Lawo Lena Obrist Ursula Ott Jörg Romeis Ted Turlings Rene Wunderlin Doris Zuber

Germany Teja Tscharntke Wolfgang Völkl

Ines Vollhardt Anke Steppuhn Netherlands Martijn Bezemer Joop van Lenteren Suzanne Lommen Paul van Rijn Jeroen van Schelt Susan Urbanus Louise Vet Matthijs Vos Roel Wagenaar Karin Winkler

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Thanks

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More beneficials Fewer pesticides

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Potatoes

Conservation Biological Control

Does it work at larger scales?

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125 m 125 m

135 m

250 m

5 m

20 m

40 m

5 m

20 m

40 m

Annual flower margin

Fallow

Brussels Sprouts

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Diamond back moth larvae

0

0.2

0.4

0.6

0.8

1

1.2

1.4

1.6

1.8

2

13/6/

06

23/6/

063/7

/06

13/7/

06

23/7/

062/8

/06

12/8/

06

22/8/

06

aver

age

larv

ae /

plan

t

flowers 5mflowers 20mflowers 40mcontrol 5mcontrol 20mcontrol 40m

Brussels Sprouts

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Functional plant species (perrenials for Autumn sowing with non-competitive grass mix) Native (UK)1 Growth Height Resources Beneficaries Other attributes Soil6

Hardiness (1 (artic) - 10 (tropical), where most of the UK = 7 (8 in Cornwall, 6 in the North, 4 up a mountain) Climate and competetion6 Typical flowering season6 Typical seed/fruit ripening6

Common name Anticipated main beneficeries Family name Latin bi-nomial Habit (in cm) Life cycle Nectar2 Pollen2 EFN Alternate prey/Associated phytophages3 Bird food4 Bees Hoverflies Parasitoids

Other beneficials (pest natural enemies) Pests of significance Birds Over-wintering species Weed status Availability5

Cost per 100g unless stated (£) Seed per g Competitive dominance Establishment liklihood Other Light (sand) Medium (loam) Heavy (clay) Acid Neutral Alkaline To usda Zone Drought exposure Waterlogging exposure Maritime exposure Frost exposure Competition J F M A M J J A S O N D J F M A M J J A S O N D

European elderberry Bi., Be., NE., Ph. Adoxacaeae Sambucus nigra Native Woody shrub 60-150 Perennial 0 2 YES ######## 4 (Berries) Yes (potential nesting site - Miro et al 2001) - -

Predatory Phytosid mites (Dabrowski et al 2008)

No. Relatively low diversity and abundance of phytophagous insects, year-round host for Mirid potato/beet/fruit pest L. pabulinus (Atkinson & Atkinson 2002 RT)

Yes (Atkinson & Atkinson 2002 RT; Boatman et al 2008) - Moderate eSeeds/manual collection 2.99 per sachet - - Good

Nitrophilous. Should tolerate mowing. Y Y Y Y Y Y Hardy to Z5 Tolerant - Tolerant Tolerant Shade tolerant

Fennel Be., Ho., Pa., NE., Ph. Apiaceae Foeniculum vulgare Naturalised Forb 90-150 Perennial 4 2 ######## - Yes (selected species only - de Toledo et al 2003)

Yes (Olkowski et al 1996; Colley & Luna 2000; Dufour 2000 RT; Luna et al 2000; Langoya & van Rijn 2008)

Yes (Olkowski et al 1996; Al-Doghairi & Cranshaw 1999; Dufour 2000 RT; Witting-Bissinger et al 2008)

Coccinelids (Al-Doghairi & Cranshaw 1999; Dufour 2000 RT), predatory mites (Olkowski et al 1996; Dufour 2000 RT), Araneae, Tachinids (Dufour 2000 RT)

Maybe - CARROT: Psila rosae (Degan 1998, Degan et al 1999). Low risk of supporting pests (van Rijn et al 2008). High acceptability to Psila rosae (Degan 1998, Degan et al 1999). - - None CNS/Herbi 3.65 (CNS) / 24.00 (Herbi) 175 (CNS) - Good - Y Y Y Y Y Y Hardy to Z5 Tolerant Intolerant Intolerant Tolerant Shade intolerant

Greater burnet saxifrage Ph., ?? Apiaceae Pimpinella major Native Forb 30-90 Perennial 2 1

AGROMYZIDAE: Melanagromyza sativae, Phytomyza adjuncta, Phytomyza pauliloewi, Phytomyza pimpinellae. APHIDIDAE: Anuraphis catonii, Semiaphis pimpinellae, Semiaphis pimpinellae. CECIDOMYIIDAE: Contarinia umbellatarum, Diodaulus traili, Kiefferia pericarpiicola, Lasioptera carophila, Macrolabis heraclei. EPERMENIIDAE: Cataplectica profugella, Phaulernis dentella. GEOMETRIDAE: Eupithecia absinthiata, Eupithecia centaureata, Eupithecia pimpinellata, Eupithecia trisignaria. OECOPHORIDAE: Depressaria pimpinellae. PAPILIONIDAE: Papilio machaon. - - Yes (van Rijn per com) -

Crysopids (van Rijn per comm)

No. Unsuitable for Psila rose larval development (Degan et al 1999) - - None Em. 35.00 (Em.) 410 (Em.) - Good - Y Y Y Y Y Y Hardy to Z5 Intolerant - - Tolerant Shade tolerant

Yarrow Be., Ho., Pa., NE, Ph. Asteraceae Achillea millefolium Native Forb 30-90 Perennial 1 2 ######## 1 (Seeds)

Yes (Valenzuela 1994), low visitation rate (Carvell et al 2007), Asteraceae generally good for bees (Meek et al 2002 RT)

Yes (Al-Doghairi & Cranshaw 1999; Colley & Luna 2000; Dufour 2000 RT; Luna et al 2000; Langoya & van Rijn 2008), good visitation rate (Powell et al 2004)

Yes (Valenzuela 1994; Olkowski et al 1996; Dufour 2000 RT). No - repellent (Wackers 2004)

Coccinelids, Tachinids (Valenzuela 1994; Al-Doghairi & Cranshaw 1999), Chrysopids (Al-Doghairi & Cranshaw 1999), Orius spp (Dufour 2000 RT; Alomar et al 2006), Nabids (Olkowski et al 1996; Dufour 2000 RT), coccinelids (Dufour 2000 RT)

No. Used by thrips including T. Tabaci (BRC).

Yes - Asteraceae used a food source by farmalnd birds per se (Wilson et al 1999) Yes (parasitic wasps - ref) None Em. 9.50 (Em.) 6000 (Em.) - Good - Y Y Y Y Y Y Hardy to Z2 Tolerant Intolerant Tolerant Tolerant Shade tolerant

Perrenial cornflower Bi., Be., ?? Asteraceae Centaurea montana Introduced Forb 30-60 Perennial 3 2 YES

Trama troglody tes (root

aphid) (ref).

AGROMYZIDAE:

Liriomyza centaureae,

Phy tomyza autumnalis,

Phy tomyza horticola,

Phy tomyza spinaciae.

CHYSOMELIDAE:

Psy lliodes chalcomera,

Cassida vibex. 2 (Seeds)

Yes (Goulson 2008). Asteraceae generally good for bees (Meek et al 2002 RT) - - - No.

Yes - genus is food source for farmland birds (Wilson et al 1999) - None CNS 110 (CNS) 80 (CNS) - Good - Y Y Y Y Y Y Hardy to Z3 Tolerant Intolerant - Tolerant Shade intolerant

Oxeye daisy Ho., Ph. Asteraceae Leucanthemum vulgare Native Forb 30-60 Perennial 2 1 ######## 1 (Seeds)

Yes (ref), Asteraceae generally good for bees (Meek et al 2002 RT). No - low visitation rate (Backman & Tiainen 2002; Carvell et al 2007),

Yes (Dicks et al 2002; van Rijn per comm). No - not visited (Powell et al 2004) - - No.

Yes - Asteraceae used a food source by farmalnd birds per se (Wilson et al 1999) - None Em. 9.50 (Em.) 2000 (Em.) - Good - Y Y Y N Y Y Hardy to Z3 - Intolerant Intolerant Tolerant Shade intolerant

Bird's foot trefoil Be., Ph. Fabaceae Lotus corniculatus Native Forb 5--40 Perennial 3 1 ? (Mckey 1989) ######## 1 (Veg)

Yes (Fussel & Corbet 1992; Meek et al 2002; Pywell et al 2006;Carvell et al 2007; Kleijn & Raemakers 2008; Goulson 2008)

Fabaceae are poor syrpid plants (Weiss & Stettmer, 1991) - -

Yes - PEAS: Sitona lineatus; Acrythosiphon pisum (BRC), BRASSICA: Meligethes spp (BRC). Lygus bugs (including L. rugulipennis) (BRC)

Yes - Fabaceae used a food source by farmalnd birds per se (Wilson et al 1999) - None Em. 21.00/3.00 (Em.) 500 (Em.) - Good Legume Y Y Y Y Y Y Hardy to Z5 - Intolerant Tolerant - Shade intolerant

Red clover Bi., Be., Ph. Fabaceae Trifolium pratense Native Forb 15-50 Perennial 3 3 ######## 3 (Veg)

Yes (Fussel & Corbet 1992; Comba et al 1999; Goulson & Darvill 2004; Goulson et al 2005; Pywell et al 2005; Carvell et al 2006; 2007; Pywell et al 2006; Diekotter et al 2006; Kleijn & Raemakers 2008; Goulson 2008)

Fabaceae are poor syrpid plants (Weiss & Stettmer, 1991)

Yes (Tscharntke et al 2002). Clovers generally good for Brachonids (Olkowski et al 1996). No - not exploited (Winkler et al 2003); repellent (Wackers 2004)

Orius spp (Alomar et al 2006)

Yes - MULTIPLE: Autographa gamma (BRC),. PEAS: Acrythosiphon pisum. Not exploited by P. rapae and P. xylostella

(immigrant) (Winkler et al 2003), not visited by Pieris

spp (Comba et al 1999)

Yes (Boatman et al 2008). Genus is food source for farmland birds (Wilson et al 1999) - None Em. 9.50 (Em.) 750 (Em.) - Good Legume Y Y Y Y Y Y Hardy to Z6 Intolerant Intolerant - Tolerant Shade intolerant

White clover Bi., Be., Ph. Fabaceae Trifolium repens Native Forb 5--25 Perennial 4 3 ######## 3 (Veg)

Yes (Fussel & Corbet 1992; Comba et al 1999; Diekotter et al 2006; Pywell et al 2006; Kleijn & Raemakers 2008; Goulson 2008)

Fabaceae are poor syrpid plants (Weiss & Stettmer, 1991)

Yes (Dufour 2000 RT), clovers generally good for Brachonids (Olkowski et al 1996). Non-attractive (Wackers 2004)

Carabidae (in orchards - Dufour 2000 RT)

Yes - BRASSICA: Meligethes spp (BRC), MULTIPLE: Autographa gamma (BRC).

Yes (Boatman et al 2008). Genus is food source for farmland birds (Wilson et al 1999) - None Em. 3.00 (Em.) 877 - Good Legume Y Y Y Y Y Y Hardy to Z4 Intolerant Intolerant - Tolerant Shade intolerant

Bush vetch Bi., Be., Ph. Fabaceae Vicia sepium Native Forb 30-100 Perennial 3 1 YES ######## 3 (Veg) Yes (Fussel & Corbet 1992; Goulson 2008)

Fabaceae are poor syrpid plants (Weiss & Stettmer, 1991)

No - repellent (Wackers 2004) -

No. Acyrthosiphon pisum; Aphis fabae; Megoura viciae;

Yes (Boatman et al 2008). Genus is food source for farmland birds (Wilson et al 1999) - None Scotia

1.71 per seed tray packet (Naturescape) 60 - Good Legume Y Y Y Y Y Y Hardy to Z6 Intolerant Intolerant - - Shade tolerant

Oregano / wild majoram Be., Pa., Ph. Lamiaceae Origanum vulgare Native Forb 30-60 Perennial 3 2 ######## - Yes (Comba et al 1999) Yes (ref) Yes (Wackers 2004; Dorn et al 2006) - ######## - - None Em. 20.00 (Em.) 11000 (Em.) - Good

Provides good ground cover for preventing weed establishment. Very high seed/g - may need to be included at low % rate. Y Y Y Y Y Y Hardy to Z5 - Intolerant Intolerant Tolerant Shade tolerant

Common sorrel Bi., Ph. Polygonaceae Rumex acetosa Native Forb 50-100 Perennial 0 2 ######## 4 (Seeds) Yes (pollen only - ref) Poor visitation rate (Powell et al 2004) - - No.

Yes (Boatman et al 2008). Key food source for farmland birds (Wilson et al 1999). Polygonaceae generally produce large quantities of seed (Wilson et al 1999) - None Em. 16.00 (Em.) 2000 (Em.) - Good - Y Y Y Y Y Y Hardy to Z3 - Intolerant - Tolerant Shade tolerant

Guelder rose Adoxiacaeae Viburnum opulus Native Woody shrub 150-300 Perennial 1 1 YES ######## 3 (Berries) Yes (ref) - - -

Yes - BRASSICA: Myzus persicae (BRC). Aphis fabae (BRC).

Yes - fruits and seeds taken by several bird species, although not necessarily preferred over other food sources (Englund 1993) - None - - - - Good - Y Y Y Y Y Y Hardy to Z3 Intolerant - - Tolerant Shade tolerant

Common knapweed Asteraceae Centaurea nigra/jacea Native Forb 15-120 Perennial 3 2 ######## 3 (Seeds)

Yes (Fussell & Corbet 1992; Croxton et al 2002; Goulson & Darvill 2004; Carvell et al 2006; 2007; Pywell et al 2006; Goulson 2008), Asteraceae generally good for bees (Meek et al 2002 RT)

Good visitation rate (Powell et al 2004) - -

Maybe - BRASSICA: Pieris spp (Corbet 2000). Heavily used by Leps, visited by Pieris spp (Corbet 2000).

Yes (Boatman et al 2008) genus is food source for farmland birds (Wilson et al 1999) - None Em. 16.00 (Em.) 400 - Good - Y Y Y Y Y Y Hardy to Z5 Tolerant Intolerant - Tolerant Shade intolerant

Greater knapweed Asteraceae Centaurea scabiosa Native Forb 30-120 Perennial - - ######## - None - Good - Y Y Y Y Y Y Hardy to Z4 Tolerant Intolerant - - Shade intolerant

Chickory Asteraceae Cichorium intybus Naturalised Forb 15-120 Perennial 3 3 ######## 1 (Seeds)

Asteraceae generally good for bees (Meek et al 2002 RT) - - -

Yes - CARROT: Psila rosae (BRC).

Yes - Asteraceae used a food source by farmalnd birds per se (Wilson et al 1999) - None CNS 12.70 (CNS) 500 - Good - Y Y Y Y Y Y 3 to 9 Intolerant Intolerant - Tolerant Shade intolerant

Tansy Asteraceae Tanacetum vulgare Native Forb 60-120 Perennial 2 2 ######## 1 (Seeds)

Asteraceae generally good for bees (Meek et al 2002 RT)

Yes (Olkowski et al 1996; Dufour 2000 RT)

Yes (Olkowski et al 1996; Dufour 2000 RT)

Coccinelids (Dufour 2000 RT), Orius (Olkowski et al 1996; Dufour 2000 RT), Chrysopids (Dufour 2000 RT)

No. Repels a range of insects (ref)

Yes - Asteraceae used a food source by farmalnd birds per se (Wilson et al 1999) - None CNS 70.5 (CNS) 2000 - Good - Y Y Y Y Y Y Hardy to Z4 - Intolerant Intolerant Tolerant Shade intolerant

Sweet Alyssum Brassicaceae Lobularia maritima Introduced Forb 10--20 Annual/perennial 2 1

Thrips - not F. occidentalis (Alomar et al 2006). PIERIDAE: Pieris brassicae. 1 (Veg) Yes (ref)

Yes (Valenzuela 1994; Al-Doghairi & Cranshaw 1999; Dufour 2000 RT; Alomar et al 2006; 2008)

Yes (Valenzuela 1994; Chaney 1998; Johanowicz & Mitchell 2000; Begum et al 2006; Berndt et al 2006; Fiedler et al 2008 RT)

Tachinids (Valenzuela 1994; Olkowski et al 1996; Dufour 2000 RT), Orius sp, predatory mites (Alomar et al 2006; 2008), predatory mites (Dufour 2000 RT)

Yes - BRASSICA: Pieris brassicae (BRC), Plutella xylostella (Winkler et al 2003). Flower use by P.

xylostella (immigrant) (Winkler et al 2006), potential dead-end trap crop for P. xylostella (de Groot et al 2005).

Yes - Brassicaceae used a food source by farmalnd birds per se (Wilson et al 1999) - None CNS Variable (CNS) 2600 - Good

Better for selected parasitoid than borage, coriander and buckwheat (Begum et al 2006) Y Y Y Y Y Y Hardy to Z7 - Intolerant Tolerant - Shade intolerant

Field scabious Dipsacaceae Knautia arvensis Native Forb 15-60 Perennial 1 1 ######## 3 (Seeds)

Yes (Dramstad & Fry 1995; Comba et al 1999; Backman & Tiainen 2002; Carvell et al 2007 RT; Franzen & Nilsson 2008; Goulson 2008)

Good visitation rate (Powell et al 2004) No (Kehrli & Bacher, 2008) -

No. Visited by Pieris spp (Comba et al 1999), though not reported as associated. Yes (Boatman et al 2008) - None Em. 23.00 (Em.) 150 - Good - Y Y Y Y Y Y Hardy to Z6 - Intolerant - - Shade tolerant

Lucerne Fabaceae Medicago sativa Introduced Forb 30-80 Perennial 3 1 ######## 1 (Veg) Yes (ref)

Fabaceae are poor syrpid plants (Weiss & Stettmer, 1991), poor nectar plant for syrphids (van Rijn, unpublished) No (Kehrli & Bacher 2008)

Orius spp (Alomar et al 2006), Coccinelids (Taffetani et al 2003)

Yes - BRASSICA: Pieris brassicae (BRC), Plutella xylostella (Winkler et al 2003). CEREAL: Helicoverpa armigera (immigrant). MULTIPLE: Lygus rugulipennis. Not exploited by P. rapae, exploited by P. xylostella

(immigrant) (Winkler et al 2003).

Yes - Fabaceae used a food source by farmalnd birds per se (Wilson et al 1999) - None Em. - - - Good Legume Y Y Y Y Y Y Hardy to Z5 Tolerant Intolerant - - Shade intolerant

Musk mallow

Malvaceae

Malva moschata Native Forb 30-70 Perennial 3 1

APIONIDAE: Apion aeneum, CHRYSOMELIDAE: Podagrica fuscicornis, Podagrica fuscipes. ERIOPHYIDAE: Eriophyes gymnoproctus, Aceria geranii. -

Yes (Comba et al 1999), pollen unacceptable to honeybees (Comba et al 1999 RT) Yes (Comba et al 1999) - -

No. Not visited by Pieris spp (Comba et al 1999) - - None Em. 20.00 (Em.) 500 - Good - Y Y Y Y Y Y Hardy to Z3 Intolerant Intolerant Intolerant Tolerant Shade tolerant

Common verbena Verbenaceae Verbena officinalis Naturalised Forb 30-75 Perennial 2 1 AGROMYZIDAE: Amauromyza labiatarum. - Yes (ref) - - - No. - - None CNS 78.00 (CNS) 3400 - Unknown - - - - - - - - - - - - -

Red campion Caryophyllaceae Silene dioica Forb 30-90 Perennial/biennial

Ragged robin Caryophyllaceae Lychnis flos-cuculi Forb 30-90 Perennial

Functional plant species (annuals and biennials for spring sowing without grass seed) Native (UK)1 Growth Height Resources Beneficaries Other attributes Soil6

Hardiness (1 (artic) - 10 (tropical), where most of the UK = 7 (8 in Cornwall, 6 in the North, 4 up a mountain) Climate and competetion6 Typical flowering season6 Typical seed/fruit ripening6

Common name Anticipated main beneficeries Family name Latin bi-nomial Habit (in cm) Life cycle Nectar2 Pollen2 EFN Alternate prey/Associated phytophages3 Bird food4 Bees Hoverflies Parasitoids

Other beneficials (pest natural enemies) Pests of significance Birds Over-wintering species Weed status Availability5 Cost per 100g unless stated Seed per g Competitive dominance Establishment liklihood Other Light (sand) Medium (loam) Heavy (clay) Acid Neutral Alkaline To usda Zone Drought exposure Waterlogging exposure Maritime exposure Frost exposure Competition J F M A M J J A S O N D J F M A M J J A S O N D

Cornflower Bi., Be., Ho., Pa., NE. Asteraceae Centaurea cyanus Naturalised Forb 30-60 Annual 3 2 YES

AGROMYZIDAE:

Liriomyza strigata.

APHIDIDAE: Dactynotus

jaceae, Trama troglody tes

(root aphid).

CHRYSOMELIDAE:

Psy lliodes chalcomera.

TEPHRITIDAE:

Chaetostomella cy lindrica. 2 (Seeds)

Yes (Weiss & Stettmer 1991; Comba et al 1999; Carreck et al 1999; Meek et al 2002; Pywell et al 2006; Witting et al., 2006; Goulson 2008), Asteraceae generally good for bees (Meek et al 2002 RT)

Yes (Weiss & Stettmer 1991; Solomon et al 1999; Alomar et al 2006; Witting et al., 2006; Langoya & van Rijn 2008). Good visitation rate (Powell et al 2004)

Yes (Weiss & Stettmer 1991; Winkler 2005; Witting et al 2006)

Coccinelids, Chrysopids (Weiss & Stettmer 1991), Anthocorids (Solomon et al 1999;Fitzgerald & Solomon 2004), natural enemies per se (van Rijn et al 2006)

Yes - BRASSICA: Plutella xylostella (Winkler et al 2003). Not exploited by P.

rapae, exploited by P.

xylostella (immigarnt) (Winkler et al 2003), low risk of supporting pests (van Rijn et al 2008)

Yes - genus is food source for farmland birds (Wilson et al 1999) - None Em. 9.00 (Em.) 200 (Em.) - Good - Y Y Y Y Y Y 3 to 10 Tolerant Intolerant - Tolerant Shade intolerant

Sunflower Bi., Be., Ho., Pa., NE. Asteraceae Helianthus annuus Introduced Forb 100-400 Annual 3 3 YES

Trama troglodytes (root aphid) (ref). AGROMYZIDAE: Liriomyza strigata. AHIDIDAE: Aulacorthum solani. CHYSOMELIDAE: Psylliodes chalcomera. CICADELLIDAE: Eupteryx atropunctata. 4 (Seeds)

Yes (Jones & Gillett 2005; Backman & Tiainen 2002; Goulson 2008), Asteraceae generally good for bees (Meek et al 2002 RT)

Yes (Olkowski et al 1996; Dufour 2000 RT)

Yes (Dufour 2000 RT; Jones & Gillett 2005)

Numerous (Jones & Gillett 2005), Nabids, Chysopids (Olkowski et al 1996; Dufour 2000 RT), coccinelids (Dufour 2000 RT) No.

Yes - genus is food source for farmland birds (Wilson et al 1999). Exploited by a range of farmland birds (Henderson et al 2004) - None CNS Variable (CNS) 15-40 (CNS) - Good - Y Y Y Y Y Y Hardy to Z7 Tolerant Intolerant - Intolerant Shade tolerant

Borage Bi., Be, Ho., Pa., NE., Ph. Boraginaceae Borago officinalis Introduced Forb 20-60 Annual 4 2 ######## 3 (Seeds)

Yes (Weiss & Stettmer 1991; Carreck et al 1999; Carvell et al 2006; Goulson 2008)

Yes (Weiss & Stettmer 1991; Alomar et al 2006; Langoya & van Rijn 2008)

Yes (Baggen & Gurr 1997; Baggen et al 1999). No (Campbell 2009)

Coccinelids, Chrysopids (Weiss & Stettmer 1991), natural enemies per se (van Rijn et al 2006)

No. Repels tomato worm (ref), not exploited by potato moth (Baggen & Gurr 1997; Baggen et al 1999), low risk of supporting pests (van Rijn et al 2008)

Yes - but only of importance to a small number of species (Stoate et al 2004) - None CNS 8.80 (CNS) 60 (CNS) - Good

Use at low seed rate as high coverage from a single plant (van Rijn per comm) Y Y Y Y Y Y Hardy to Z0 Tolerant Intolerant - Tolerant Shade tolerant

Scorpion weed Bi., Be., Ho., Pa. Boraginaceae Phacelia tanacetifolia Introduced Forb 20-80 Annual 4 3 - 4 (Seeds)

Yes (Carreck & Williams 1997; Kells et al 2001; Pontin et al 2000; 2006; Petanidou 2003 RT; Ambrosino et al 2006; Goulson 2008) ########

Yes (Holland & Thomas 1996; Fiedler et al 2008 RT) Tachinids (Dufour 2000 RT)

No. Supports abundant mirids (Holland & Thomas 1996), minimally effected by slug damage (Sievwright et al 2006). No impact on mites and springtails in broccoli (Wyland et al 1996)

Yes - high in gamebird chick food (mirid bugs) (Holland & Thomas 1996), exploited by a range of farmland birds (Henderson et al 2004) - None CNS 13.00 (500g) (CNS) 525 (CNS) - Good

Aethestically popular with general public (Sievwright et al 2006). More attractive than buckwheat to bumblebess (Pontin et al 2006 RT) Y Y Y Y Y Y Hardy to Z8 Intolerant Intolerant - Tolerant Shade intolerant

Common vetch Bi., Be., Ph. Fabaceae Vicia sativa (subsp. seqetalis) Naturalised Forb 10-100 Annual 3 1 YES ######## 3 (Veg) Yes (Fussel & Corbet 1992; Comba et al 1999)

Yes (Langoya & van Rijn 2008). Fabaceae are poor syrpid plants (Weiss & Stettmer, 1991) Yes (Wackers et al 2006)

Orius spp (Alomar et al 2006)

Yes - PEAS: Sitona lineatus; Acyrtosiphon pisum (BRC), BRASSICA: Pieris rapae (Comba et al 1999). Aphis fabae; Megoura viciae; Low risk of supporting nectar feeding pests (van Rijn et al 2008), visited by P. rapae (Comba et al 1999).

Yes (Boatman et al 2008). Genus is food source for farmland birds (Wilson et al 1999) - None Em. 10.00 (Em.) 50 (Em.) - Good

Legume. High germination rate typical of cultivated Vicia = low seeding rate required (van Rijn per comm) Y Y Y Y Y Y Hardy to Z5 Intolerant Intolerant - Tolerant Shade tolerant

Red dead nettle Bi., Be., Ph. Lamiaceae Lamium purpureum Naturalised Forb 07--30 Annual 2 1 ######## 3 (Seeds?)

Yes (Goulson 2008; Lye te al in press). V.important for queens in spring due to early flowering period (S. O'Conner per.comm.)

Poor visitation rate (Powell et al 2004) - - No.

Yes (Boatman et al 2008). Seeds of no value (Storkey 2006) - Moderate B. 87.00 (B.) 1050 - Good - Y Y Y Y Y Y Hardy to Z4 Intolerant Intolerant - Tolerant Shade tolerant

Yellow rattle Be., Ph. (plus sward) Orobanchaceae Rhinanthus minor Native Forb 20-80 Annual 2 0 ######## -

Yes (Meek et al 2002; Pywell et al 2006; Goulson 2008) - - -

Yes - BRASSICA: Helicoverpa armigera (imigrant) (BRC) - - None Em. 21.00 (Em.) 300 (Em.) - Good

Hemi-parasitic - may restrict grass growth to encourage forb diversity (Pywell et al 2004; Westburry & Dunett 2008) Y Y Y Y Y Y Hardy to Z6 Intolerant - - - Shade intolerant

Buckwheat Bi., Be., Ho., Pa., NE. Polygonaceae Fagopyrum esculentum Introduced Forb 15-60 Annual 4 3 ? (Zim. 1932)

Exploited by a large number of phytophages (Thapa 2006). CHYSOMELIDAE: Chaetocnema concinna. NOCTUIDAE: Euxoa tritici. 4 (Seeds) Yes (Thapa 2006)

Yes (Valenzuela 1994; Dufour 2000 RT; Luna et al 2000; Pontin et al 2006; Thapa 2006; van Rijn et al 2006; Langoya & van Rijn 2008)

Yes (Valenzuela 1994; Dufour 2000 RT; Wratten et al 2003; Lee et al 2004; Begum et al 2006 RT; Winkler et al 2006; Irvin et al 2006; Berndt et al 2006; Thapa 2006; Fiedler et al 2008 RT; Witting-Bissinger et al 2008)

Chrysopids, Orius, Tachinids, Coccinelids (Valenzuela 1994), Tachinids (Olkowski et al 1996; Dufour 2000 RT), Chrysopids, coccinelids, orius (Dufour 2000 RT), coccinelids (Thapa 2006)

Yes - BRASSICA: Exploited by Pieris rapae and Plutella

xylostella (immigrant) (Winkler et al 2003), Pieris rapae (Thapa 2006). Low risk of supporting pests (van Rijn et al 2008). Exploited by potato moth (Baggen & Gurr 1997) and Lygus bugs (ref).

Yes (exploited by a range of farmland birds - Henderson et al 2004). Polygonaceae generally produce large quantities of seed (Wilson et al 1999) - None Suffolk Herbs

1.50 per 150g (Suffolk Herbs) 45 - Good

Soil improver (Valenzuela 1994). Better for selected parasitoids than fennel (Witting-Bissinger et al 2008) Y Y Y Y Y Y Hardy to Z0 Intolerant Intolerant - Intolerant Shade intolerant

Viper's bugloss Be. , Ph. Boraginaceae Echium vulgare Native Forb 30-100 Biennial 3 2 ######## - Yes (Goulson et al 2005; Goulson 2008) Yes (ref) - -

Yes - BRASSICA: Meligethes spp (BRC). - - None Em. 16.00 (Em.) 350 (Em.) - Good - Y Y Y Y Y Y Hardy to Z0 - Intolerant Tolerant Tolerant Shade intolerant

Teasel Bi., Be., Ph. Dipsacaceae Dipsacus fullonum Native(?) Forb 90-200 Biennial 3 2 ######## 4 (Seeds)

Yes (Comba et al 1999; Kells et al 2001; Goulson 2008) Yes (Comba et al 1999) - -

No. Not visited by Pieris spp (Comba et al 1999)

Yes (Boatman et al 2008). High value seed source for some spp (Stoate et al 2004) Teasel moth (per. obs.) None Em. 9.50 (Em.) 300 (Em.) - Good - Y Y Y Y Y Y Hardy to Z5 Intolerant - - - Shade intolerant

Bishops weed Ho., NE. Apiaceae Ammi majus Naturalised/introduced? Forb 60-100 Annual - -

APHIDIDAE: Dysaphis apiifolia. CECIDOMYIDAE: Lasioptera carophila. ERIOPHYIDAE: Eriophyes peucedani. PAPILIONIDAE: Papilio machaon. - -

Yes (Olkowski et al 1996; Dufour 2000 RT) -

Orius, Tachinids (Olkowski et al 1996), Tachinids, coccinelids, orius (Dufour 2000 RT) No.

Toxic and causes photosensitisation if consumed in large enough quantities (Egyed et al 1974) - None CNS 17.00 (CNS) 1200 - Good Good winter survival Y Y Y Y Y Y Hardy to Z0 Intolerant - - - Shade tolerant

Dill Apiaceae Anethum graveolens Naturalised/introduced? Forb 45-120 Annual 2 2

APHIDIDAE: Dysaphis apiifolia. CECIDOMYIIDAE: Lasioptera carophila. PAPILIONIDAE: Papilio machaon. PSILIDAE: Psila rosae. - Yes (ref)

Yes (Olkowski et al 1996; Valenzuela 1994; Dufour 2000 RT)

Yes (Baggen & Gurr 1997; Al-Doghairi & Cranshaw 1999; Dufour 2000 RT; Winkler 2005; Dorn et al 2006) ########

Yes - CARROT: Psila rosae (BRC), BRASSICA: Plutella xylostella (Winkler et al 2003). Not exploited by P.

rapae, exploited by P.

xylostella (immigrant) (Winkler et al 2003). Low acceptability to Psila rosae (Degan 1998) - - None CNS 3.00-5.00 (CNS) 600 - Good - Y Y N Y Y Y Hardy to Z8 Intolerant Intolerant - - Shade intolerant

Coriander Apiaceae Coriandrum sativum Introduced Forb 30-45 Annual 4 1 PSILIDAE: Psila rosae. - Yes (ref)

Yes (Olkowski et al 1996; Al-Doghairi & Cranshaw 1999; Colley & Luna 2000; Dufour 2000 RT; Ambrosino et al 2006; Pontin et al 2006 RT; Langoya & van Rijn 2008)

Yes (Baggen & Gurr 1997; Al-Doghairi & Cranshaw 1999; Dufour 2000 RT Fiedler et al 2008 RT)

Coccinelids (Al-Doghairi & Cranshaw 1999; Landis et al 2000 RT), Tachinids (Valenzuela 1994; Dufour 2000 RT), Chysopids (Landis et al 2000), Orius (Olkowski et al 1996; Dufour 2000 RT) natural enemies per se (van Rijn et al 2006)

Yes - CARROT: Psila rosae (BRC). Repels aphids (ref), low risk of supporting pests (van Rijn et al 2008), exploited by potato moth (Baggen & Gurr 1997). - - None CNS approx 3.00 (CNS) 100 - Good - Y Y N Y Y Y Hardy to Z5 - Intolerant - Tolerant Shade tolerant

Yellow chamomile Asteraceae Anthemis tinctoria Introduced Forb 30-60 Biennial 2 2

AGROMYZIDAE: Ophiomyia curvipalpis. MIRIDAE: Orthocephalus coriaceus. PYRALIDAE: Euzophera cinerosella, Apion confluens, Apion dispar, Apion laevigatum. 1 (Seeds)

Asteraceae generally good for bees (Meek et al 2002 RT) - - - No.

Yes - Asteraceae used a food source by farmalnd birds per se (Wilson et al 1999) - None CNS 10.50 (CNS) 5000 - Good - Y Y Y N Y Y Hardy to Z6 - Intolerant Tolerant Tolerant Shade intolerant

Chrysanthemum/corn marigold Asteraceae Chrysanthemum spp (segetum?) Native Forb 30-60 Annual - -

Brachycaudus cardui (ref).

CURCULIONIDAE:

Ceutorhynchus triangulum.

DIASPIDIDAE:

Hemiberlesia lataniae.

NOCTUIDAE: Phlogophora

meticulosa. TINGIDAE:

Catoplatus fabricii. 1 (Seeds)

Asteraceae generally good for bees (Meek et al 2002 RT)

Spp segetum - variable effect on longevity (Langoya & van Rijn 2008) -

Anthocirids (Solomon et al 1999). Parasitic nematodes (Dufour 2000 RT) No.

Yes - Asteraceae used a food source by farmalnd birds per se (Wilson et al 1999) - None Em. 8.00 (Em.) 700 - Good - Y Y Y Y Y N Hardy to Z0 Intolerant Intolerant - - Shade tolerant

Chamomile Asteraceae Matricaria chamomilla Native Forb 15-40 Annual - -

Brachycaudus cardui (ref).

TEPHRITIDAE: Trupanea

stellata. 1 (Seeds)

Asteraceae generally good for bees (Meek et al 2002 RT) Yes (ref) - Coccinelids (ref) No.

Yes - Asteraceae used a food source by farmalnd birds per se (Wilson et al 1999) - None Em. (2 wk delay) - - - Good - - - - - - - - - - - - -

White dead nettle Lamiaceae Lamium album Naturalised Forb 30-60 Annual 2 1 ######## 3 (Seeds?)

Yes (Fussel & Corbet 1992; Comba et al 1999; Goulson 2008; Lye et al in press). V.important for queens in spring due to early flowering period (S. O'Conner per.comm.) - -

Predatory Phytosid mites (Dabrowski et al 2008)

Yes - BRASSICA: Meligethes spp Yes (Boatman et al 2008) - Moderate - - - - Good - Y Y Y Y Y Y Hardy to Z4 Intolerant Intolerant - Tolerant Shade tolerant

Red bartsia Orobanchaceae Odontites vernus Native Forb 15-40 Annual 2 1

AGROMYZIDAE: Phytomyza isais. GEOMETRIDAE: Eupithecia plumbeolata, Perizoma bifaciata - Yes (Goulson 2008) - - - No. - - None - - - - Good

Hemi-parasitic - may restrict grass growth to encourage forb diversity (Goulson 2008) - - - - - - - - - - - -

Herb robert Geraniaceae Geranium robertianum 10--60

Dovesfoot cranesbill Geraniaceae Geranium molle 05--40

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Taking it to the field

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parasitoid distribution relative to margin

0

2

4

6

8

10

1 2 3 4 5 15 20

brassica field row

num

ber o

f par

asito

ids

flower margincontrol

20 m

5 m

Annual flower margin

Grass (control)

15 m

2 m 3 m

4 m

1 m

Do parasitoids/predators actually utilize flowering plants?

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Nutritional benefits?

Or migration effects?

X X

X X

X X

X

X X

X

X

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05

1015202530

05

1015202530

05

1015202530

05

1015202530

flower margin

field

control field

row 1 to 5 row 15 to 20

*

*

Fed, high sugar levels

Fed, low sugar levels

Unfed

*

*

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0 25 50 75 100 125 150 175 200 225

fru

ctose

/tota

l su

gars

0.0

0.1

0.2

0.3

0.4

0.5

0.6

0.7

Newly-emerged

Unfed

Fed < 15 days

Fed > 15 days

Separation lines

g total sugars/parasitoidg total sugars/parasitoid

0 25 50 75 100 125 150 175 200 225

fru

ctose

/tota

l su

gars

0.0

0.1

0.2

0.3

0.4

0.5

0.6

0.7

Separation lines

0-2 m

2-4 m

4-6 m

6-8 m

8-10 m

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0 25 50 75 100 125 150 175 200 225

fru

ctose

/tota

l su

gars

0.0

0.1

0.2

0.3

0.4

0.5

0.6

0.7

Newly-emerged

Unfed

Fed < 15 days

Fed > 15 days

Separation lines

g total sugars/parasitoid

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Identifying honeydew sources

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How to select seed mixes that optimize

ecosystem services?

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