COVER: Boreus reductus Carpenter (Mecoptera: Boreidae)

Boreids (order Mecoptera) are interesting for two traits, they are often found wandering

the surface of snow from late November until May, and they can jump. A feat they achieve

using metatibial extension and resilin to store energy in the thorax before a leap. Their

common name of "snow fleas" is both apt and prescient, given that recent molecular stud-

ies have placed them as close relatives of the true fleas. Their biology is poorly known and

most texts state that both adults and larvae "live in moss". This example is a female of the

species Boreus reductus (Carpenter 1933). The image was taken a few hundred metres

from the junction of the Coquihalla Highway and Highway | at the west end of Kamloops

in March 2007. An Olympus E-1 DSLR with a Zuiko ZD 50mm macro lens coupled

to two 25mm_ extension tubes was used. The magnification on

the sensor (18mmx1l4mm) was X1 .5 . This image is a crop of the original image, the

length of the insect from frons to tip of ovipositor is approximately 4 mm. Lighting was

with a 1980's vintage Vivitar 283 flash with a homemade 10cm by 15cm cloth diffuser.

The most important part of the field equipment was a set of chest waders.

The Journal of the Entomological Society of British Columbia is

published annually in December by the Society

Designed and typeset by Hugh Barclay and Jen Perry.

Printed by Vernon Graphics, Vernon, B.C.

Printed on Recycled Paper.

Pug

J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008 1

Journal of the

Entomological Society of British Columbia

Volume 105 Issued December 2008 ISSN #0071-0733

Directors of the Entomological Society of British Columbia, 2008-2009 .0.0.... ec ceessseeeeeeeees 2

G.G.E. Scudder. New provincial and state records for Heteroptera (Hemiptera) in Canada and the

NO IGC CES TALC cate ale ass tame neater Reta cas ana aa punsu een oaNerocaanataas aah ubnkeneedeas meedemeanenetce: 3

Jeremy R. Dewaard, B. Christian Schmidt, Gary G. Anweiler and Leland M. Humble. First Ca-

nadian records of Lampropteryx suffumata ([Denis & Schiffermitller], 1775) (Geometridae:

PEATE TCIM LAG) Seasee otess esieecares tee oh exe tna se gure tans om eat re tron aac eetee cau eam oneoteeueeneeee eee 19

D. Thomas Lowery, Michael G. Bernardy, Robyn M. Deyoung and Chris J. French. Identifi-

cation of new aphid vector species of Blueberry SCOrCHh Virus .......ccccccseeccneeecnneceeneseens 27

John K. Mackenzie, Peter J. Landolt and Richard S. Zack. Sex attraction in Polistes dominulus

(Christ) demonstrated using olfactometers and morphological source extracts .............:006 2)

Gary J.R. Judd and Mark G.T. Gardiner. Efficacy of Isomate-CM/LR for management of leafrol-

lers by mating disruption in organic apple orchards of western Canada .............cccsssceceeeees 45

D.A. Raworth. Climate change and potential selection for non-diapausing two-spotted spider

mites on strawberry in southwestern British Columbia ..............ccccccccccccceeeeesssseceeeeeeeeeeesenes 61

Michael D. Doerr, Jay F. Brunner and Timothy J. Smith. Biology and management of bark

beetles (Coleoptera: Curculionidae) in Washington cherry orchards .............ccccssseeeees 69

Kevin Durden, John J. Brown and Maciej A. Pszczolkowski. Extracts of Ginkgo biloba or Ar-

temisia species reduce feeding by neonates of codling moth, Cydia pomonella (Lepidoptera:

Tortricidae), on apple in a laboratory bi0assay ............cccccssccessccesseeeeseecesseecsseeecssecesseeeesaees 83

S.D. Cockfield and E.H. Beers. Management of dandelion to supplement control of western

flower thrips (Thysanoptera: Thripidae) in apple orchards ..................cccseeecceceeeeseeseeeeees 89

Luis Martinez-Rocha, Elizabeth H. Beers and John E. Dunley. Effect of pesticides on inte-

grated mite management in Washington State ............ccccccccceeeccccuesccseuscceunseseueeseees oF

SCIENTIFIC NOTES

R.D. Kenner and K.M. Needham. Additional records for semiaquatic Hemiptera

musOutha western British: Colt Bialik cae css ssetacen saat stu deti sit Rvahanttectits ttortenc a ettesece Oe 109

Virgiliu M. Aurelian, Mario Lanthier and Gary J.R. Judd. Podosesia syringae (Lepidoptera: Se-

siidae): a new clearwing moth record for British Columbia ..........ccccccccccsesesssesseeeesseseees i!

Willem G. Van Herk, J. Todd Kabaluk, Viola W.M. Lam and Robert S. Vernon. Survival of male

click beetles, Agriotes obscurus L., (Coleoptera: Elateridae) during and after storage at differ-

CTE TE IMP CLAUS Seeks. t aaasseenle ne aocomseewestanecen ecotecelst nae aaSaieats weoaacna rman ner ntesnsinesanitenteeetaaeineret 113

W.G. Van Herk and R.S. Vernon. Effect of handling and morbidity induction on weight,

recovery, and survival of the Pacific Coast wireworm, Limonius canus (Coleoptera: Elateri-

LAS es cae eee rere et en Sac tare te aen ae Pee AME REN We RP ca. ca.s eede cena eee 115

ABSTRACTS

Symposium Abstracts: Biodiversity in Stanley Park. Entomological Society of British Columbia

Annual General Meeting, West End Community Centre Auditorium, Vancouver, BC, October

RU ZOO SOR eee ee CRON denon sitet ost cioaacsaseeaat it sven sie mh atmasvwiieoeaee eaatzattae mtn aR 117

NOTICE TO CONTRIBUTORS .0.0.....cccccccccccccsccssescseeseceesscseesesscessessesnens Inside Back Cover

J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

DIRECTORS OF THE ENTOMOLOGICAL SOCIETY

OF BRITISH COLUMBIA FOR 2008-9

President:

Sheila Fitzpatrick

Agriculture and Agri-Food Canada

President-Elect:

Tom Lowery

Agriculture and Agri-Food Canada

Past-President:

John McLean

University of British Columbia

Secretary/Treasurer:

Lorraine Maclauchlan

B.C. Ministry of Forests and Range

Directors, first term:

Dezene Huber, Alida Janmaat, Leo Rankin

Director, second term:

Jim Corrigan, Rob McGregor, Melanie Hart

Regional Director of National Society:

Bill Riel

Canadian Forest Service, Victoria

Editor, Boreus:

Jennifer Heron

Jennifer. Heron@gov.bce.ca

Editorial Committee, Journal:

Editor-in-Chief: Subject Editors:

Hugh Barclay Sheila Fitzpatrick (Agriculture)

Pacific Forestry Centre Lorraine MacLauchlan (Forestry)

HBarclay@pfc.cfs.nrcan.gc.ca Geoff Scudder (Systematics/Morphology)

Technical Editor: Jen Perry Editor Emeritus: Peter Belton

Editor of Web Site:

Bill Riel

briel@pfc.forestry.ca

Honorary Auditor:

John McLean

University of British Columbia

Web Page: http://www.sfu.ca/biology/esbe/

J. ENTOMOL. SOc. BRIT. COLUMBIA 105, DECEMBER 2008

New provincial and state records for Heteroptera

(Hemiptera) in Canada and the United States

G.G.E. SCUDDER!

ABSTRACT

New provincial records are provided for 52 species of Heteroptera in Canada. Dichaeto-

coris piceicola (Knight) is also reported from Alaska, and Pagasa nigripes Harris from

Washington State.

INTRODUCTION

Since the publication of the checklist of

the Hemiptera of Canada and Alaska (Maw

et al. 2000), new Heteroptera from Canada,

and new provincial records have been pub-

lished by Barnes et al. (2000), Henry et al.

(2008), Jansson (2002), Kenner and

Needham (2004), Paiero et al. (2003), Roch

(2007), Schuh (2000a, 2000b, 2001, 2004a,

2004b), Schuh and Schwartz (2004, 2005),

Schwartz and Scudder (2001, 2003),

Schwartz and Stonedahl (2004), Scudder

(2000, 2004, 2007), Scudder and Foottit

(2006), Scudder and Schwartz (2001),

Wheeler and Hoebeke (2004), and Wheeler

et al. (2006). Wright (1989) and Kerzhner

(1993) also published records not included

in Maw et al. (2000).

Recent research has revealed additional

new provincial records for 52 species. Di-

chaetocoris piceicola (Knight) has been

found in Alaska, and Pagasa nigripes Har-

ris in Washington State.

These are reported below, with Museum

abbreviations as follows:

AAFCL: Agriculture and Agri-Food

Canada, Lethbridge, AB (J.R. Byers).

CNC: Canadian National Collection of

Insects, Agriculture and Agri-Food Canada,

Ottawa, ON (R.G. Foottit).

LM: Lyman Entomological Museum,

Macdonald College, McGill University,

Ste.-Anne-de-Bellevue, QC (T. Wheeler).

MU: Memorial University. St. John’s ,

NF (D. Larson).

NSM: Nova Scotia Museum of Natural

History, Halifax, NS (A. Hebda and C. Ma-

jka).

RBCM: Royal British Columbia Mu-

seum, Victoria, BC (R.A. Cannings).

SM: Saskatchewan Provincial Museum,

Saskatoon, SK (R. Hooper).

UBC: Spencer Entomological Museum,

Department of Zoology, University of Brit-

ish Columbia, Vancouver, BC (K.M.

Needham).

UG: Department of Environmental Biol-

ogy, University of Guelph, Guelph, ON

(S.A. Marshall).

UM: J.B. Wallis Collection, University

of Manitoba, Winnipeg, MB (R.E. Rough-

ley).

UPEI: Department of Biology, Univer-

sity of Prince Edward Island, Charlotte-

town, PE (K.A. Campbell and D. Giber-

son).

NEW PROVINCIAL RECORDS

The systematic order of families and

higher taxa listed below, follows Maw et al.

(2000).

Infraorder NEPOMORPHA

Family CORIXIDAE

' Department of Zoology, University of British Columbia, 6270 University Boulevard, Vancouver, BC

VOT 1Z4. (604) 822-3682. scudder@zoology.ubc.ca

4 J. ENTOMOL. SOc. BRIT. COLUMBIA 105, DECEMBER 2008

Arctocorisa chanceae Hungerford

A subarctic species known from Alaska

and across northern Canada from Yukon to

Newfoundland and Labrador (Scudder

1997; Maw et al. 2000). The species can be

keyed using Hungerford (1948) and Brooks

and Kelton (1967).

New record. BC: 14, Atlin, 10 mi S,

18.viil.1980 (R.J. Cannings) [UBC].

Sigara compressoidea (Hungerford)

An eastern Nearctic species, in Canada

reported from Ontario east to Newfound-

land (Maw et al. 2000), but not previously

reported from Prince Edward Island. Sigara

compressoidea 1s keyed in Hungerford

(1948) and Tinerella and Gundersen (2005),

and a dorsal colour photograph is given in

the latter reference, showing the typically

effaced membrane patterning.

New record. PE: 14 19, Deroche Natu-

ral Protected Area, 46.42114°N 62.94082°

W, kicknet, small wetland, 27.1x.2007 (K.

Alexander Campbell) [UPEI].

Family NOTONECTIDAE

Notonecta spinosa Hungerford

A Cordilleran species, so far in Canada,

recorded only from British Columbia

(Scudder 1977; Maw et al. 2000). The spe-

cies can be keyed using Hungerford (1933).

New records: AB: 1 specimen, Waterton

Lakes Nat. Park, grassland pond with mud-

stone, substrate, near Buffalo Paddock, 49°

07'44"N_ 113°S1'11"W, 9.vii.2005 (R.E.

Roughley & R.D. Kenner) [UBC]; 1 speci-

men, Waterton Lakes Nat. Park, spring-fed

Carex marsh, near West Entrance on Hwy.

5, 49°07'12"N 113°50'S3"W, 9.v11.2005

(R.E. Roughley & R.D. Kenner) [UBC].

Family PLEIDAE

Neoplea striola (Fieber)

This pygmy backswimmer, so far in

Canada is recorded only from Manitoba,

Ontario and Quebec (Maw ef al. 2000). The

species is keyed in Brooks and Kelton

(1967).

New record. BC: 1 specimen, Edge-

wood, F11, edge forest by beaver pond,

25.vill.-5.1x.1988 (H. Knight) [CNC].

Infraorder GERROMORPHA

Family VELIIDAE

Microvelia pulchella Westwood

This cosmopolitan species is recorded

from Alaska, and in Canada from most

provinces from British Columbia to New-

foundland (Maw et al. 2000). The species is

keyed in Brooks and Kelton (1967).

New record. SK: 1¢ 29, Cowan Dam at

Hwy. 55, 54°11'49"N_ 107°27'0"W, Typha

pond, 22.v11.2003 (R.E. Roughley) [CNC].

Family GERRIDAE

Gerris incognitus Drake & Hottes

A species reported across Canada from

British Columbia to Labrador (Maw et al.

2000), but not previously recorded from

Prince Edward Island. The species is keyed

by Drake and Harris (1934).

New record. PE: 1¢ 19, Millburn,

8.v1.1970 (Ray Wenn) [UPE]].

G. pingreensis Drake & Hottes

This species is known from Alaska and

across Canada from Yukon to Labrador

(Maw et al. 2000), but to date there are no

published records for New Brunswick,

Prince Edward Island, Nova Scotia and

Newfoundland. The species is keyed by

Drake and Harris (1934) and Brooks and

Kelton (1967).

New record. NF: 2¢ 19, Plum Point,

ponds, 11.1x.1999 (T. Huxley) [CNC].

Limnoporus notabilis (Drake & Hottes)

This western water strider in Canada 1s

known from British Columbia and Alberta

(Maw et al. 2000; Andersen and Spence

1992). The species is keyed by Andersen

and Spence (1992).

New record. SK: 14, Cypress Hills,

15.vu1.1977 (R. Hopper) [SM].

Family SALDIDAE

Micracanthia humilis (Say)

A Nearctic-Neotropical species, widely

distributed in North America, and in Can-

ada reported from British Columbia and

Northwest Territories east to Newfound-

land, but not previously recorded from

Nova Scotia (Maw et al. 2000). The species

J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

is keyed by Schuh (1967) and Polhemus

and Chapman (1979).

New records. NS: 29, Halifax Co.,

Lawrencetown Beach, viii.1979 (B. Wright)

[NSM]; 19 Halifax Co., Petpeswick Har-

bour, wet sandy upper shore, 29.vu.1971

(Barry Wright) [NSM].

Salda lugubris (Say)

A widely distributed Nearctic species

that also occurs in Mexico, and in Canada is

reported from Yukon to Newfoundland, but

not previously from New Brunswick and

Nova Scotia (Maw et al. 2000). The species

is keyed by Brooks and Kelton (1967) and

Schuh (1967).

New records. NB: 13, French Lake,

2.vii.1928 (W.S. Brown) [CNC]. NS: 19,

CBI, Chiticamp, vi.-vii.1917 (F. Johansen)

[CNC]; 29°, Halifax Co., Port Wallis,

17.vi1.1952 (D.C. Ferguson) [NSM].

Saldula ablusa Drake & Hottes

This is an eastern Nearctic species,

keyed and reported from Ontario by Schuh

(1967). Polhemus (1988) noted that refer-

ences to Saldula xanthochila (Fieber) from

the northeastern United States almost cer-

tainly refer to S. ablusa or S. pallipes

(Fabricius). Wright (1989) reported S. xan-

thochila from Sable Is., Nova Scotia, and

examination of 24 12 specimens in the

Nova Scotia Museum of Natural History,

labeled “NS. Sable Is., brackish ponds east

of station, 26.v1i.1976 (Barry Wright)”

shows these to be S. ablusa. As pointed out

by Polhemus (1985), S. ablusa lacks a dis-

tinct dark distal streak on the ventral side of

the hind femora, and as noted by Schuh

(1967) typically has pale lateral margins to

the pronotum, whereas these are never pre-

sent in the Saldula “pallipes — palustris”

group.

New record. PE: 73 19, Can. Nat.

Park, Dalvay House, 19.vii.1940 (GS.

Walley) [CNC].

S. bouchervillei (Provancher)

This species was previously reported

from Nova Scotia as Salda bouchervillei

(Provancher) by Wright (1989).

Material examined: NS: 29, Sable Is-

land, brackish ponds east of station,

26.vii.1976 (Barry Wright) [NSM].

S. laticollis (Reuter)

Lindskog (1981) clarified the identity of

this species, distinguishing it from the

closely related S. pallipes (Fabricius) and S.

palustris (Douglas) by the presence of long,

curved, semi-recumbent or suberect setae

on the head dorsally, and noted that S. fer-

naldi Drake is a synonym. In Old World

populations of both S. pallipes and S. palus-

tris, the pubescence on the head and dor-

sum generally is uniformly short and re-

cumbent. However, as observed by Schuh

(1967), the Saldula “pallipes — palustris”

species complex has confused systematic

heteropterists for some time, because of the

extreme variability of the “species”. Never-

theless, coastal populations from western

North America and Newfoundland, previ-

ously identified as S. palustris have been

shown to be S. Jaticollis (Lindskog 1981;

Polhemus 1988), the intertidal biology of

which has been described by Stock and

Lattin (1976) under S. palustris.

Wright (1989) reported S. palustris from

Sable Is., Nova Scotia, but examination of

specimens in the Nova Scotia Museum of

Natural History shows these to be S. laticol-

lis. This species is also now known from

New Brunswick and Prince Edward Island.

New records. NB: 14 19, Kouchi-

bouguac Nat. Park, 14.vi.1977 (S.J. Miller)

[CNC]. NS: 2¢ 39, Digby Co., Sandy

Cove, 4.vii.1971 (Barry Wright) [NSM];

22, Guysborough Co., Liscombe, marshy

area above beach, 13.vi.1971 (Barry

Wright) [CNC]; 23 19, Sable Island, West

Light, 23.vii.1976 (Barry Wright) [NSM];

1¢ 19, Sable Island, freshwater pond at

West Light, 13.vi.1977 (Barry Wright)

[NSM]. PE: 16, Brackley Beach,

5.vii.1966 (L.A. Kelton) [CNC]; 34, Green

Gables, Cavendish Beach, 22.vi1.1967

(J.E.H. Martin) [CNC].

Infraorder CIMICOMORPHA

Family ANTHOCORIDAE

6 J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

Anthocoris tomentosus Péricart

This western Nearctic, and Beringian

species occurs in the western United States

south to Arizona (Lewis et al. 2005), and

Alaska to Manitoba, but has not previously

been reported from Saskatchewan (Maw et

al. 2000). The species is keyed in Kelton

(1978) under the name A. melanocerus

Reuter, and characteristically has the he-

melytra entirely shiny, and the pronotum

and antennae completely black. Lewis et al.

(2005) reported A. tomentosus being regu-

larly collected from Alnus, Populus, Pyrus

and Salix growing in and near fruit-growing

regions in Washington State, as well as on

Rumex and psyllid-infested Sheperdia ar-

gentea (Pursh) Nutt.

New record. SK: 192, Fort Qu’Appelle,

17.vi.1967 (R. Hooper) [SM]; 19, Regina,

on elm, 6.x.1986 (K. Roney) [SM].

Dufouriellus ater (Dufour)

This species was described from

Europe, and was first reported from Amer-

ica north of Mexico by Van Duzee (1916).

It is now known to be widely distributed in

North America, with published records in

the USA for New York (Van Duzee 1917),

North Carolina (Blatchley 1926), California

and Kentucky (Blatchley 1928), Idaho

(Harris and Shull 1944), Oregon (Lattin

2004) and Hawaii (Lattin 2005, 2007a),

whereas in Canada it has previously been

reported from British Columbia (Anderson

1962) and Ontario (Kelton 1978).

Dufouriellus ater is usually collected

under the bark of trees, but also is often

associated with stored products (Awadallah

et al. 1984; Arbogast 1984; Lattin 1999).

As a result, it is a useful predator of some

economic importance (Lattin 2000).

The species is keyed by Kelton (1978),

who noted that in Canada it is rare in col-

lections, and probably introduced into Brit-

ish Columbia and Ontario. Although D.

ater was not included as an alien in Canada

by Scudder and Foottit (2006), Lattin

(2004, 2007b) considered the species to be

non-indigenous to America north of Mex-

ico.

New record. NS: 14, Halifax, Grain

Elevators, from dust and debris samples,

17.vu1.1991 (J. Hulton) [NSM].

Family NABIDAE

Nabis inscriptus (Kirby)

This Holarctic species, with somewhat

abbreviated wings is difficult to distinguish

from shorter winged specimens of N.

americoferus Carayon. Reliable separation

is based on the shape of the male parameres

and structure of the copulatory pouch of the

female, as illustrated in Vinokurov (1988)

and discussed by Kerzhner (1963). Nabis

inscriptus 1s recorded from Alaska and

across northern Canada (Maw ef al. 2000),

but as noted by Henry and Lattin (1988),

reported distribution records need to be

verified.

New records. NS: 19, Antigonish Co.,

Pomquet Beach, 31.v.1978 (B. Wright)

[NSM]; 16), Halifax, found on corpse, Path.

Lab., 1.xi.1979 (B. Wright) [NSM].

N. roseipennis Reuter

This species is recorded from British

Columbia to Nova Scotia in Canada (Maw

et al. 2000), but has not previously been

reported from Prince Edward Island. The

species is keyed in Blatchley (1926) and

Harris (1928) and characteristically has

black spots on the hind tibiae.

New record: PE: 63 89, Charlottetown,

UPEI, nr. East edge, 46°15'25"N 63°

O8'08"W, sweeping, 20.1x.2004 (G.G.E.

Scudder) [CNC, UPEI].

Pagasa nigripes Harris

Kerzhner (1993) raised Pagasa fusca

var. nigripes Harris to specific rank, and

recorded the species in Canada from Al-

berta, Quebec and Saskatchewan. He also

reported P. nigripes from Alaska, Colorado,

Massachusetts, New Hampshire, New Mex-

ico, New York, Pennsylvania, Vermont and

Wyoming. He distinguished it from Pagasa

fusca (Stein) by differences in the male and

female genitalia, and noted that the legs

tend to be brown or brownish yellow in P.

nigripes, whereas they are yellow in P.

fusca, with the femora often orange or red-

dish. However, the legs in P. fusca can

J. ENTOMOL. SOc. BRIT. COLUMBIA 105, DECEMBER 2008

sometimes be partly brownish or entirely

black. Hence, the genitalic characters are

the most reliable. The parameres in P. fusca

are relatively large with the outer margin

rounded, whereas in P. nigripes they are

slightly smaller than in P. fusca and dis-

tinctly angulate on the outer margin.

Pagasa nigripes is here recorded for the

first time from British Columbia, Northwest

Territory and Yukon, where the species was

previously reported as P. fusca in Maw et

al. (2000): P. fusca does not occur in

Alaska, Northwest Territory and Yukon.

However, in British Columbia, P. fusca co-

occurs with P. nigripes, the species having

been collected together at Merritt (23 km E,

Hamilton Commonage), Osoyoos (Mt. Ko-

bau in Montane Spruce habitat), Vaseux

Creek (Kennedy bench), and the Winder-

mere Valley.

New records. BC: 14 , Canal Flats, 10.8

km S., 31.viii.1998 (G.G.E. Scudder)

[CNC]; 13, Fairview, White L., BGxhl,

SWm, pan trap WL/P-1, 4.vit.-11.vii.1995

(J. Jarrett) [UBC]; 32, Merritt, 23 km E,

Hamilton Commonage, Upper Fescue

grassland, early seral, 1250 m, 16.ix.2000

(G.G.E. Scudder) [CNC, UBC]; 19, Mer-

ritt, 35 km S, 14.viii.1988 (G.G.E. Scudder)

[CNC]; 164, Nicola, 24.vii.1932 (GJ.

Spencer) [UBC]; 14', Osoyoos, East Bench,

Artemisia/Purshia assoc., pitfall trap,

1S.vii. 17.vili. 1990 (G.G.E. Scudder)

[CNC]; 14, Osoyoos IRI, Inkaneep,

BGxhl, AN, pitfall trap T1-1, 6.vii.-

9.viii.1995 (G.G.E. Scudder) [CNC]; 16,

id., 9.viit.-9.ix.1995 [CNC]; 19, id., 9.ix.-

6.x.1995 [CNC]; 12 id, T2-1, 9.viii.-

9K 199s NENG lia 9d.. 12-5. 4 viz

7.1x.1994 [CNC]; 14, id., T4-1, 9.viii.-

9.1x.1995 [CNC]; 19, id., T4-3, 6.vii.-

9.vii.1995 [CNC]; 19, id., T4-5, 4.viii.-

7.1x.1994 [CNC]; 19, id. T5-2, 9.viii.-

9.1x.1995 [CNC]; 14, Osoyoos, Mt. Kobau,

MSxh, VK, Pitfall trap K4A-5, 10.viii.-

8.1x.1995 (J. Jarrett) [UBC]; 19, id., K4A-

3, 18.viii.-28.1x.1997 [UBC]; 14, Osoyoos,

Mt. Kobau Rd., IDFdk1, pitfall trap K3B-5,

18.viii.-28.ix.1996 (J. Tome) (UBC); 16;

id., IDFxhl, pitfall trap K2B-4, 18.viii-

28.ix.1996 [UBC]; 13, id., PPxhl, pitfall

trap K1A-2, 28.vii.-18.viii.1997 [UBC]; 12

12, Tatlayoka L., 16.vii.1978 (G.G.E.

Scudder) [CNC]; io Vaseux Cr., CWS

bench, BGxhl, AN, pitfall trap Y1-4, 6.ix.-

4.x.1995 (G.G.E. Scudder) [CNC]; 19, id.,

Y2-5 [CNC]; 16, id. Y1-4, 8.vii-

3.viii.1994 [CNC]; 19, ee Y2-5, 12.viii.-

64x. 1995 [ONC]: (12, ids. > Y¥2-5,) -61x.-

4.x.1995 [CNC]; 19, id. Y3-3, 5.vii.-

12.viii.1995 [CNC]; 23, id., Y3-3, 6.ix.-

4.x.1995 [CNC]; 16, id., Y3-5, 5.vii.-

12.viii.1995 [CNC]; 14, id, Y4-l,

12.viii.6.ix.1995 [CNC]; 13, id, Y4-5,

3.viii.-6.1x.1994 [CNC]; 164, id., Y4-5,

5.vii.-12.viii.1995 [CNC]; 14, Vaseux Cr.,

‘Kennedy bench’, 49°16'N 119°30'W,

BGxhl, AN, pitfall trap Z2-4, 3.vi-

8.vii.1994 (G.G.E. Scudder) [CNC]; 19,

id., Z2-5, 12.viii.-6.ix.1995 [CNC]; 10,

Vaseux Cr., ‘Kennedy flats’, 49°15'N 119°

31'W, BGxhl, AN, pitfall trap X2-1, 6.ix.-

4.x.1995 (G.G.E. Scudder) [CNC]; 19, id.,

X3-4 [CNC]; 14, X3-5, 12.vii.-6.ix.1995

[CNC]; 19, id., X4-1 [CNC]; 14. Vaseux

L., Wildlife Res., BGxhl, AN:F, pitfall trap

VLI-2, 27.vi.-17.vi.1997 (J. Jarrett)

[UBC]; 14, Westwick L., Cariboo,

28.vi.1961 (J. Scudder) [UBC]; 19, id

17.viii.1962 (G.G.E. Scudder) [CNC]; 33

49, id., 18.viii.1962 [CNC, RBCM, UBC];

12, White L., BGxhl, SWm, pitfall trap

WL2-3, 17.viii.-28.ix.1996 (J. Jarrett)

[UBC]; 145 29, Windermere Valley, pitfall

trap No. 5, 26.vii.-17.1x.2000 (R. Sargent)

[CNC, UBC]. NT: 19, Fort Smith,

27.v.1950 (W.G. Helps) [CNC]; 19, Fort

Smith, 6.viii.1950 (J.B. Wallis) [CNC]. YT:

2°, Alaska Hwy. km 1768, Duke R.,

9.vii.1983 (G.G.E. Scudder) [CNC]; 16,

Alaska Hwy. mi 1054, Kluane L.,

16.vii.1962 (G.G.E. Scudder) [CNC];1¢,

id., 7.vii.1983 [CNC]; 24, Canyon, Aishi-

hik R., 9.vu.1983 (G.G.E. Scudder) [CNC];

Bet 39, Carcross, 8.vii.1983 (G.G.E. Scud-

der) [CNC, UBC]; 14, Lapie R., 1 km E on

Campbell Hwy., 28.vii.1981 (C.S. Guppy)

[UBC]; 192, Pelly Crossing, 17.vii.1983

(G.G.E. Scudder) [CNC]; 19, Tatchun Cr.,

62°17'N 136°17'W, 17.vu.1983 (G.G.E.

Scudder) [CNC]; 19, Whitehorse,

17.vii.1959 (R. Madge) [CNC]; 19, White-

8 J. ENTOMOL. SOc. BRIT. COLUMBIA 105, DECEMBER 2008

horse, 31.vii.1981 (C.S. Guppy) [UBC].

I have also collected P. nigripes in

Washington State, USA, as follows: WA:

19, Oroville, E. Osoyoos L., 48°53'N 119°

25'W, Purshia assoc., AN BGxhl, pitfall

trap O2-1, 10.1x.-4.x.1995 (G.G.E. Scud-

der) [CNC].

Family MIRIDAE

Ceratocapsus modestus (Uhler)

A widely distributed eastern Nearctic

species, previously reported from Sas-

katchewan east to Quebec in Canada (Maw

et al. 2000). Recorded hosts in West Vir-

ginia are Quercus alba L. and Vitis sp.

(Wheeler et a/. 1983). The species is keyed

by Knight (1941), Henry (1979), Kelton

(1980) and Larochelle (1984).

New records. NS: 19, Grand Pre, on

Picea, 10.vii.1966 (L.A. Kelton) [CNC];

19, Grand Pre, Pinus _ sylvestris,

10. vil. 1966 (L.A. Kelton) [CNC].

Conostethus americanus Knight

To date this species in Canada has been

reported only from Alberta, Northwest Ter-

ritories and Saskatchewan (Maw ef al.

2000). On the prairies C. americanus oc-

curs on grasses (Kelton 1980). It is keyed

and illustrated in Kelton (1980).

New record. YT: 43° 79, Whitehorse,

Dillabough’s graze lease, 8V 6754911

490889, 12.vii.2005 (G.E. Hutchings)

[RBCM].

Cyrtorhinus caricis (Fallén)

A Holarctic species, reported to occur

on sedge (Carex spp.) across Canada

(Kelton 1980), and also recorded from

Alaska and Minnesota, with an apparent

relict population occurring in Colorado

(Wheeler and Henry 1992). The species is

keyed and illustrated by Kelton (1980).

New record. NS: 14, Lake Egmont,

18.vii.1991 (B. Wright) [NSM].

Deraeocoris quercicola Knight

An eastern Nearctic species, widely

distributed and in Canada reported to date

from Saskatchewan east to Quebec (Maw ef

al. 2000). Recorded hosts include Carya

sp., Quercus alba, Q. ilicifolia Wangenh

and Tilia americana L. (Wheeler et al.

1983), as well as Quercus macrocarpa

Michx. where it preys on aphids (Kelton

1980). The species is keyed by Knight

(1921), Kelton (1980) and _ Larochelle

(1984).

New record. NB: 19, St. Johns, Rock-

wood Pk., 5.viii.1954 (J.F. Brimley)

[CNC].

D. triannulipes Knight

This Nearctic species in Canada has

previously been reported from British Co-

lumbia east to Quebec (Maw ef al. 2000).

Deraeocoris triannulipes 1s reported to feed

on aphids on Populus tremuloides Michx.

and A/nus spp. (Kelton 1980). It is keyed by

Knight (1921), Kelton (1980) and La-

rochelle (1984).

New records. NS: 14, Exfern, on apple,

4.vii.1950 (F.T. Low) [CNC]; 19, Halifax,

Pyrus, 22.vii.1976 (L.A. Kelton) [CNC];

2°, Kentville, on apple, 10-14.vii.1976

(L.A. Kelton) [CNC]; 14, Kentville, Tilia

cordata, 15-17.vii.1976 (L.A. Kelton)

[CNC].

Dichaetocoris piceicola (Knight)

A western Nearctic species, known from

Colorado in the USA, and in Canada re-

corded from Alberta, British Columbia and

Yukon (Maw ef al. 2000). The genus is

keyed by Knight (1968). The species D.

piceicola is distinguished from the only

other northern species (D._ gillespiei

Schwartz and Scudder) by Schwartz and

Scudder (2003). It 1s recorded from Picea

engelmanni Parry (Polhemus 1994), and in

British Columbia has been collected on

Abies lasiocarpa (Hook.) Nutt., Picea sp.,

Pinus contorta Dougl. and Tsuga hetero-

phylla (Raf.) Sarg. (Scudder, unpublished).

New record. AK: 12, Mosquito L., 59°

27'N 136°02'W, 6.vii.1983 (G.G.E. Scud-

der) [CNC].

Labops verae Knight

A western Nearctic and Beringian spe-

cies distributed from Alaska to Manitoba

and south to Washington State (Henry and

J. ENTOMOL. Soc. BRIT. COLUMBIA 105, DECEMBER 2008

Wheeler 1988; Maw ef al. 2000), but not

previously reported from Saskatchewan.

The species is keyed in Slater (1954) and

Kelton (1980). The host plants are unknown

(Kelton 1980).

New record. SK: 1, Stony Rapids,

30.vi.1975 (R. Hooper) [SK].

Lygidea salicis Knight

This Nearctic species in Canada is pre-

viously reported from Alberta east to New-

foundland (Maw ef al. 2000), and in the

United States from New York to Minne-

sota, Colorado, and California (Henry and

Wheeler 1988). It has not previously been

recorded from British Columbia. Lygidea

salicis Knight is a small species with the

average length in the male of 5.8 mm, and

in the female 6.2 mm. The species is keyed

by Kelton (1980), who notes that the pilos-

ity on the second antennal segment is

shorter than the thickness of this segment.

Lygidea salicis is usually collected on Salix

spp. (Kelton 1980; Wheeler et a/. 1983).

New records. BC: 24 39, Fernie, gold-

enrod, 23.vui.1959 (L.A. Kelton) [CNC];

2¢ 29, Mt. Revelstoke Nat. Pk., Salix,

17.vii.1970 (L.A. Kelton) [CNC].

Megalopsallus femoralis Kelton

This species so far has been reported

from Alberta, Manitoba, Saskatchewan,

Colorado, South Dakota and Wyoming

(Schuh 2000b). Megalopsallus femoralis

has been collected on Salicornia rubra

Nels. (Kelton 1980). It is keyed by Kelton

(1980) and Schuh (2000b), and is illustrated

in colour in the latter reference.

New record. BC: 3¢ 29, Kamloops,

Ironmask L., 10U 6804 56152, saline flats,

Salicornia/Plantago, 730 m., 14.vi.1995

(S.G. Cannings) [RBCM].

Orthotylus alni Knight

This Nearctic species is distributed from

Yukon to Newfoundland, and south to New

York and Minnesota in the eastern United

States (Henry and Wheeler 1988; Maw ef

al. 2000). However, it has not previously

been recorded from Nova Scotia. Orthoty-

lus alni is keyed by Kelton (1980), and has

been collected on Alnus rugosa (DuRoi)

Spreng. (Kelton 1980), as well as A. tenui-

folia Nutt., Betula glandulosa Michx., Lu-

pinus sp. and Salix sp. (Scudder 1997).

New records. NS: 14, Chester,

10.vii.1969 (B. Wright) [NSM]; 19, Ches-

ter, 16.vii.1969 (B. Wright) [NSM].

O. nyctalis Knight

Described originally from Minnesota

(Knight 1927), this species has been re-

ported in the USA also from Iowa, Illinois,

New York and Wisconsin (Henry and

Wheeler 1988). It has not previously been

recorded from Canada under this name.

According to Knight (1927), O. nyctalis

can be recognized chiefly on the structure

of the male genital claspers. The left clasper

is slender with two short dorsal prongs, and

the right clasper decurved on the apex and

devoid of spines, but the dorsal margin has

a prominent spine at the basal third, and

two other spines just before the decurved

apex.

The record of O. candidatus Van Duzee

from Saskatchewan (Kelton 1980; Maw ef

al. 2000; Roch 2007) is evidently referable

to O. nyctalis, as is the record of O. candi-

datus from Ontario (Maw et al. 2000; Roch

2007). The recorded occurrence of O. can-

didatus in Quebec (Henry and Wheeler

1988; Roch 2007) may also refer to O.

nyctalis. Some specimens from Dawson

and Moose Creek in the Yukon, listed as

Orthotylus sp. in Scudder (1997) are actu-

ally O. nyctalis (see below), but it may be

noted that O. candidatus also occurs at both

these localities. Orthotylus nyctalis evi-

dently occurs on Populus tremuloides

(Kelton 1980).

New records. AB: 16, Stettler,

3.vili.1957 (A. & J. Brooks) [CNC]: 19,

Vermilion Provincial Park, Beaverdam

Loop Trail, Populus tremuloides Michx.,

22.viil.1993 (M.D. Schwartz) [CNC]. MB:

14, Falcon L., S.viii.-10.viii.1978 (L.A.

Kelton) [CNC]; 19, id., 6.vii.1978 (L.A.

Kelton) [CNC]; 19, Rennie, 16.viii.1961

(F.I.S.) [CNC]. ON: 13, One Sided Lake,

Salix sp., 1.viii.1960 (Kelton and Whitney)

[CNC]; 19, Tillsonburg, 18.vii.1962

10 J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

(Kelton and Thorpe) [CNC]. SK: 19, Cy-

press Hills Prov. Park., Sheperdia canaden-

sis, 19.1x.1951 (L.A. Konotopetz) [CNC].

YT: 14, Dawson, 14 mi E, Populus sp.,

29.vii.1962 (R.E. Leech) [CNC]; 19, Daw-

son, Salix sp., 23.vi1.1983 (L.A. Kelton)

[CNC]; 12, Moose Creek, Salix sp.,

28.vil.1983 (L.A. Kelton) [CNC].

Phytocoris buenoi Knight

An eastern Nearctic species, in Canada

previously reported from Ontario and Que-

bec (Maw et al. 2000). The species is keyed

by Blatchley (1926). Knight (1920) re-

ported it to occur on Norway spruce (Picea

abies (L.) Karst.) in the eastern United

States, and Wheeler et al. (1983) added

Picea glauca (Moench) Voss and P. rubens

Sarg.

New records. NS: 16, Chester,

29.vill.1968 (B. Wright) [NSM]; 19, Sandy

Cove, 4.viii.1971 (B. Wright) [NSM]; 18,

Baddeck, 28.vili.1972 (B. Wright) [NSM];

14, Kemptville, 24.viii.1982 (Agriculture

Canada) [NSM].

P. procteri Knight

This species, which is a member of the

P. junceus Knight group, was described

from Maine (Knight 1974), and is reported

from Quebec (Roch 2007). The frons has

definite transverse red lines, the pronotum

is pallid with a basal submarginal strong

black band, and the propleura are brownish

black. The clavus is more or less fuscous

external to the claval vein, with insect

length of 8.0 mm. The first antennal seg-

ment is pallid, but clothed with black, re-

cumbent setae, and without distinct dark

spots. The first antennal segment is longer

than the width of the vertex, but does not

exceed the width of the pronotum. The sec-

ond antennal segment is without annuli or

coloured bands. There are no records of a

host for this species.

New records. NS: 1 specimen (abdomen

missing), Lake Kejimkujik, 13.vu.1961

(D.C. Ferguson) [NSM]; 1, Chester,

24.v11.1968 (B. Wright) [NSM].

Pilophorus neoclavatus Schuh &

Schwartz

This eastern Nearctic species to date in

Canada has been reported from Alberta east

to Quebec (Maw et al. 2000), and is keyed

by Schuh and Schwartz (1988). Hosts in-

clude Alnus rugosa, Quercus ilicifolia, Q.

palustris, Q. stellata Wangenh and Salix

longifolia Muhl. (Schuh and Schwartz

1988).

New record. NS: 19, Kentville,

8.viii.1952 (C.R. McL.) [LM].

Rhinocapsus rubricans (Provancher)

An eastern Nearctic species, in Canada

to date reported from Saskatchewan east to

Quebec (Maw eft al. 2000). The species is

keyed and illustrated in Kelton (1980), who

reports it collected on Kalmia polifolia

Wang.

New record. NS: 164, Chester,

4.vii.1969 (B. Wright) [NSM].

Sixeonotus deflatus Knight

An eastern Nearctic species, in Canada

previously only reported from Quebec

(Larochelle 1984; Maw et al. 2000). The

species 1s keyed by Larochelle (1984) as S.

insignis Reuter, who records the host as

Symplocarpus foetidus (L.).

New records. NB: 14, St. John,

9.viii.1954 (J. Brimley) [CNC]. ON: 83

59, St. Catherines, 22.vi.1961 (Kelton and

Brampton) [CNC].

Family TINGIDAE

Acalypta lillianis Torre-Bueno

This Nearctic tingid is widely distrib-

uted in North America and Beringia (Drake

and Lattin 1963; Scudder 1997). It occurs

in Alaska, and from Yukon to Newfound-

land, but has not previously been reported

from Manitoba (Maw et al. 2000). It is

keyed by Drake and Lattin (1963), who

note that host records are mosses.

New record. MB: 16, Bird Cove, 4 km

NE, Churchill North Studies Centre, 58°

46'14"N 93°50'33"W, pit trap in tundra

zone, 18.viil.2006 (Boreal & Arctic Ento-

mol.) [UM].

J. ENTOMOL. Soc. BRIT. COLUMBIA 105, DECEMBER 2008

Family ARADIDAE

Aradus uniannulatus Parshley

A Nearctic and Beringian species, dis-

tributed from Yukon to Quebec, and in the

United States, south to New York and

Colorado (Scudder 1997). The species is

keyed by Matsuda (1977) who notes that it

is reported to be associated with Pinus con-

torta murrayana (Balf.) Critchfield else-

where.

New records. BC: 19, Fernie,

22.viii.1934 (Hugh Leech) [CNC]; 19,

Lorna, Picea engelmanni, 17203 Lot 1,

29.vi.1929 (R. Hopping) [CNC]; 13, id.,

17203 Lot 2, 10.vii.1924 [CNC]; 19, id.,

Pinus contorta, 17203 Lot 8, 12.vu1.1924

[CNC]; 13, id., Picea engelmanni, 17203

Lot 14, 12.vii.1924 [CNC]; 3, id., 17203

Lot 25, 25.vii.1924 [CNC]; 16, id., 17203

Lot 26, 27.vii.1924 [CNC] 19, Midday

Valley, Merritt, Pinus ponderosa, Exp.

17501 Lot 1035, 30.v.1923 (R. Hopping)

[CNC]; 19, Pine Pass, Picea, 11.vii.1972

(D.E. Bright) [CNC].

Infraorder PENTATOMOMORPHA

Family COREIDAE

Leptoglossus occidentalis Heidemann

This leaf-footed bug, commonly called

the western conifer seed bug, feeds on nu-

merous conifer species (Koeber 1963;

Krugman 1969; Hedlin et al. 1981; Schae-

fer and Mitchell 1983; Gall 1992; Mitchell

2000) although the apparently strong reli-

ance on Pinaceae as a food source is not

absolute (Mitchell 2000). However, it is of

considerable economic importance because

it can cause significant losses in conifer

seed orchards (Koeber 1963; Schowalter

and Sexton 1990; Blatt and Borden 1996;

Mitchell 2000; Strong et a/. 2001; Bates et

al. 2002; Strong 2006). The species is

keyed in Allen (1969) and McPherson et al.

(1990), and illustrated by Koeber (1963)

and Ruth et al. (1982).

Originally considered a western Nearc-

tic species, L. occidentalis has naturally

expanded its range eastwards in the past

few decades (Schaffner 1967; McPherson ef

al. 1990; Marshall 1991; Gall 1992; Ridge-

O’Connor 2001), and has invaded Europe

(Taylor et al. 2001; Gogala 2003; Tescari

2004; Hilpold 2005; Rabitsch and Heiss

2005; Ribes and Oleguer 2005; Foldessy

2006; Moulet 2006). It is now known to

occur in Nova Scotia.

New records. NS: 19, Kings Co., Mid-

dleton, in house, 20.ix.2006 (J. Parks)

[NSM]; 19°, Halifax Co., Halifax, on house,

3.x.2006 (B. Fay) [NSM]; 29, Kings Co.,

Lakeville, 1 of 4, 12.111.2007 (J. Morton)

[NSM 36185]; 1°, Halifax Co., Halifax, in

dwelling, 15.x.2007 (John Sherwood)

[NSM Cat. 36219].

Family RHOPALIDAE

Liorhyssus hyalinus (Fabricius)

This cosmopolitan species is widely

distributed in North America, and in Can-

ada has so far been reported from British

Columbia, Manitoba and Ontario (Maw ef

al. 2000). It is keyed by Blatchley (1926),

Slater and Baranowski (1978), and Hoe-

beke and Wheeler (1982), and illustrated in

Slater and Baranowski (1978). Schaefer and

Chopra (1982) report Abutilon, Euphorbia,

Lactuca and Sonchus as host plant genera.

New record. QC: 19, Terrebonne Co.,

Lac Carre, Lot 31, Range 8, 19-23.viii.1968

(W. Boyle and R. La Conde) [LM].

Family LYGAEIDAE

Kleidocerys ovalis Barber

This widely distributed Nearctic species

has so far in Canada only been recorded

from British Columbia and Ontario (Maw

et al. 2000), as well as Quebec (Roch

2007). It is keyed in Barber (1953) and

Scudder (1962). In British Columbia, K.

ovalis has been collected on Abies lasio-

carpa, Alnus sp., Betula occidentalis Hook.,

B. papyrifera Marsh., Malus sp., and Pinus

ponderosa Dougl. (Scudder, unpublished).

New records. AB: 50 29, Drumheller,

18.vi.1957 (Brooks, MacNay) [CNC]; 72

69, id., 11.vili.1957 (A.R. & J.E. Brooks)

[CNC]; 23 39, Empress, 7.vi.1957

(Brooks, MacNay) [CNC]; 19, Lethbridge,

3.vii.1929 (J.H. Pepper) [CNC]; 13, Lund-

breck, 7.vii1.1930 (J.H. Pepper) [CNC].

MB: 19°, Aweme, 5.v1i.1920 (H.A. Robert-

son) [CNC]; 19, id., 8.vii.1920 (P.N.

12 J. ENTOMOL. Soc. BRIT. COLUMBIA 105, DECEMBER 2008

Vroom) [CNC]; 39, id., Betula, 30.vi.1922

(N. Criddle) [CNC]; 19, id., 3.v.1923 (N.

Criddle) [CNC]; 14 19. Carberry, 9.v.1953

(Brooks, Kelton) [CNC]; 14 39, Ninette,

Betula glandulosa, 21.vi.1958 (J.F. McAI-

pine) [CNC]; 13, id., 14.vii.1958 (R.B.

Madge) [CNC]; 16, id., 15.vii.1958 (R.L.

Hurley) [CNC]; 1¢, Onah, 10.v.1923 (R.M.

White) [CNC]; 84 19, id., Betula papyrif-

era, 10.ix.1930 (R.M. White) [CNC]; 19,

Turtle Mtn., 22.vii.1953 (Brooks, Kelton)

[CNC]. SK: 19, Punnichy, 21.v.1965 (R.

Hooper) [SM].

K. resedae (Panzer)

This Holarctic species is widely distrib-

uted in North America, and occurs in

Alaska and from Yukon to Newfoundland

and Labrador, but has not previously been

recorded from Prince Edward Island. It is

keyed in Barber (1953) and Scudder (1962).

Kleidocerys resedae usually occurs on A/-

nus spp. and Betula spp. (Scudder 1997).

New records. PE: 33 19, Blooming

Point, 46°24'33"N 62°58'07"W, sweeping,

20.x.2004 (G.G.E. Scudder) [CNC, UPE]];

2¢ 49, Charlottetown, UPEI, nr. NE point,

46°15'39"N 63°08'19"W, sweeping,

20.x.2004 (G.G.E. Scudder) [CNC, UPEI].

Melacoryphus lateralis (Dallas)

A widely distributed Nearctic species,

so far only recorded from British Columbia

and Saskatchewan in Canada (Maw eft al.

2000). The species is keyed by Slater

(1988). Specimens of M. lateralis collected

at light in Wyoming contained cardenolides

in the body (Scudder and Duffey 1972), and

thus showed evidence of feeding on Ascle-

piadaceous host plants.

New record. ON: 16, Guelph,

3.vill.1977 (W.A. Attwater) [UG].

Family RHYPAROCHROMIDAE

Antillocoris minutus (Bergroth)

An eastern Nearctic species, in Canada

previously recorded from Ontario east to

Newfoundland (Maw et al. 2000). The ge-

nus is keyed in Blatchley (1926), Slater and

Baranowski (1978), and Larochelle (1984),

with key to species given by Barber (1952)

and Larochelle (1984). The biology of A.

minutus 1n New England has been de-

scribed by Sweet (1964), who notes the

species typically occurs on the ground and

usually is found in forest litter, most fre-

quently found beneath gray birch (Betula

populifera Marsh) and white birch (B. pa-

pyrifera), but also occurs under hemlock

(Tsuga canadensis (L.) Carr.), and in

sphagnum bogs.

New record. MB: 19, Winnipeg, St.

Charles Rifle Rge., Block B Refuge, Pitfall

trap, 6-13.x.1999 (D.A. Pollock, J.K. Diehls

and R.E. Roughley) [UM].

Drymus unus (Say)

An eastern Nearctic species, in Canada

so far recorded from Saskatchewan east to

Nova Scotia (Maw et al. 2000). The species

is keyed by Blatchley (1926) and La-

rochelle (1984), and illustrated by Slater

and Baranowski (1978). Sweet (1964) de-

scribed the biology D. unus in New Eng-

land, and noted that this is a ground-

dwelling species, most abundant in subcli-

max forests, particularly where black birch

(Betula lenta L.) and red maple (Acer ru-

brum L.) are associated with oak (Quercus

spp.) and hickory (Carya spp.).

New record. NF: 1, St. John’s, Long

Pond, ix.-x.2001 (Biology 4150) [MU].

Perigenes constrictus (Say)

This eastern Nearctic species is distrib-

uted throughout the northern and central

United States, and in Canada so far reported

from Nova Scotia, Ontario and Quebec

(Maw et al. 2000). The species is keyed by

Blatchley (1926) and Larochelle (1984),

and illustrated by Blatchley (1926) and Sla-

ter and Baranowski (1978). Sweet (1964)

described the biology of P. constrictus in

New England and noted that it typically

occurs in temporary habitats, such as vacant

lots, roadsides and newly fallow fields.

New record. SK: 14, Big Beaver,

9.vil.1974 (R. Hooper) [SM].

Plinthisus americanus Van Duzee

In Canada previously reported from

Alberta east to New Brunswick (Maw et al.

J. ENTOMOL. Soc. BRIT. COLUMBIA 105, DECEMBER 2008

2000). Plinthisus americanus is distin-

guished by the hemelytra of the female be-

ing densely pilose as noted by Sweet

(1964), whereas in the closely related P.

compactus (Uhler) the hemelytra of the

female are glabrous. Sweet (1964) noted

that P. americanus is a forest species in

New England, most abundant in Tsuga lit-

ter.

New record. BC: 192, Attachie, 32 km

W of Charlie L., 5.vili.1982 (R.A. Can-

nings) [RBCM].

Trapezonotus arenarius (Linnaeus)

A Holarctic species with a wide distri-

bution in both the Nearctic and Palearctic,

in Canada recorded from British Columbia

and Yukon east to Quebec (Maw ef al.

2000), and in the United States evidently

restricted to the highlands of New England

and northern New York (Sweet 1964). The

species is keyed by Blatchley (1926) and

Larochelle (1984), and illustrated by Slater

and Baranowski (1978). Sweet (1964) re-

ported that in New England T. arenarius is

a species of open upland habitats, particu-

larly well-drained and rather dry sites.

New record. NS: 1, Lunenberg,

7.vili.1991 (B. Wright) [NSM].

Family CYDNIDAE

Amnestus pusillus Uhler

A widely distributed species in North

America, with recorded occurrence also in

Mexico and Guatemala (Froeschner 1960).

In Canada, so far reported only from On-

tario and Quebec (Maw et al. 2000). The

species is keyed in Froeschner (1960),

McPherson (1982), and Larochelle (1984).

McPherson (1982) reported that elsewhere

A. pusillus has been collected from vegeta-

tion along streams and margins of roadsides

and cultivated fields, as well as beneath

rubbish in sandy places.

New record. NB: 192, Woodstock,

22.v.1966 (L.A. Kelton) [CNC].

Family PENTATOMIDAE

Acrosternum hilare (Say)

A widely distributed Nearctic species, in

Canada so far recorded only from British

Columbia, Ontario and Quebec (Maw et al.

2000). This species has been collected from

numerous plants (McPherson 1982) and can

damage some crops (Panizzi et al. 2000).

The species is keyed in Blatchley (1926),

McPherson (1982), Rolston (1983), and

Larochelle (1984).

New record. NS: 19, Debert, 1.ix.1952

(R.L. Horsburgh) [LM].

Cosmopepla intergressus (Uhler)

A widely western Nearctic species, in

Canada so far recorded only from British

Columbia (Maw ef al. 2000). Cosmopepla

intergressus 1s keyed by McDonald (1986),

who reported records of the species on

“currants”, Rubus parviflorus Nutt. and

Ribes sp.

New record. AB: 3¢ 39, Lethbridge,

black current, 21.1x.2005 (J.R. Byers)

[AAFCL].

Euschistus euschistoides

(Vollenhoven)

A Nearctic species widely distributed in

North America, and in Canada recorded

from British Columbia to Nova Scotia, but

not previously reported from New Bruns-

wick (Maw ef al. 2000). It has been re-

corded from numerous host plants

(McPherson 1982), and the species has

caused yield and quality losses to several

crops (Panizzi et al. 2000). The species is

keyed in McPherson (1982) and Larochelle

(1984).

New records. NB: 19, Jonah Mt.,

3.vi.1976 (P. Kevan) [LM]; 192, Whittier

Ridge, 30.v.1976 (P. Kevan) [LM].

sServus

Menecles insertus (Say)

Widely distributed in North America,

and in Canada previously recorded from

Nova Scotia, Ontario and Quebec. Mene-

cles insertus is nocturnal, arboreal, and

phytophagous, and has been collected on

deciduous trees (McPherson 1982). The

species is keyed in Blatchley (1926), Rol-

ston (1973), McPherson (1982) and La-

rochelle (1984).

New record. SK: 19, Buffalo Pound

Park, 5.v.1975 [SM].

14 J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

Zicrona caerulea (Linnaeus)

This Holarctic species is widely distrib-

uted in North America, Europe and Asia,

and occurs in the Oriental region (De

Clercq 2000). In Canada it is reported from

British Columbia east to New Brunswick

(Maw eft al. 2000). This predaceous species

attacks only small prey (De Clercq 2000),

and is keyed by McPherson (1982), La-

rochelle (1984), and Thomas (1992).

New record. NF: 1, Red Indian L.,

Winddrift Lot 5, 25.vi.1980 (Brennan and

Larson) [MU].

Family THYREOCORIDAE

Corimelaena pulicaria (Germar)

A widely distributed species in North

America, also reported from Mexico and

Guatemala. In Canada recorded from Brit-

ish Columbia to Nova Scotia, but not pre-

viously reported from New Brunswick

(Maw et al. 2000). The species has been

collected on many plants (McPherson

1982), and is keyed in Blatchley (1926),

McPherson (1982), and Larochelle (1984).

New record. NB: 19, Fredericton,

French Lake, 10.vi.1931 (C.W. Maxwell)

[LM].

ACKNOWLEDGEMENTS

The research for this paper was sup-

ported by grants from the Natural Sciences

and Engineering Research Council of Can-

ada. I thank the curators of the various

collections for permission to examine the

material in their care and/or loan of speci-

mens. Dr. M.D. Schwartz helped me in my

study of the Canadian National Collection

of Arthropods in Agriculture and Agri-

Foods Canada in Ottawa, kindly identified

or confirmed the identity of most of the

Miridae listed, and allowed me to include

the records of Orthotylus nyctalis. Dr. T.J.

Henry (United States Department of Agri-

culture, Washington, DC) provided useful

comments on the manuscript. I thank

Launi Lucas for the final preparation of the

paper.

REFERENCES

Allen, R.C. 1969. A revision of the genus Leptoglossus Guerin (Hemiptera: Coreidae). Entomologica

Americana (New Series) 45:35-140.

Andersen, N.M. and J.R. Spence. 1992. Classification and phylogeny of the Holarctic water strider genus

Limnoporus Stal (Hemiptera, Gerridae). Canadian Journal of Zoology 70:753-785.

Anderson, N.H. 1962. Anthocoridae of the Pacific Northwest with notes on distribution, life histories, and

habits (Heteroptera). Canadian Entomologist 94:1325-1334.

Arbogast, R.T. 1984. Demography of the predaceous bug Dufouriellus ater (Hemiptera: Anthocoridae).

Environmental Entomology 13:990-994.

Awadallah, K.T., F.S. Tawfik, N. Abou-Zeid and El-Husseini. 1984. The life-history of Dufouriellus ater

(Duf.) (Hemiptera: Anthocoridae). Bulletin de la Societé Entomologique d'Egypt 63:191-197.

Barber, H.G. 1952. The genus Anftillocoris Kirk. in the United States (Hemiptera-Lygaeidae). Bulletin of the

Brooklyn Entomological Society 47:85-87.

Barber, H.G. 1953. A revision of the genus Kleidocerys Stephens in the United States (Hemiptera: Lygaei-

dae). Proceedings of the Entomological Society of Washington 55:273-283.

Barnes, D.I., H.E.L. Maw and G.G.E. Scudder. 2000. Early records of alien species of Heteroptera

(Hemiptera: Prosorrhyncha) in Canada. Journal of the Entomological Society of British Columbia 97:95-

102.

Bates, S.L., C.G. Lait, J.H. Borden and A.R. Kermode. 2002. Measuring the impact of Leptoglossus occi-

dentalis (Hemiptera: Coreidae) on seed production in lodgepole pine using an antibody-based assay.

Journal of Economic Entomology. 95:770-777.

Blatchley, W.S. 1926. Heteroptera or True Bugs of Eastern North America, with especial reference to the

Faunas of Indiana and Florida. Nature Publishing Company, Indiana. 1116 pp.

Blatchley, W.S. 1928. Notes on the Heteroptera of eastern North America with descriptions of new species.

1. Journal of the New York Entomological Society 36:1-23.

Blatt, S.E. and J.H. Borden. 1996. Distribution and impact of Leptoglossus occidentalis Heidemann

(Hemiptera: Coreidae) in seed orchards in British Columbia. The Canadian Entomologist 128:1065-1076.

J. ENTOMOL. Soc. BRIT. COLUMBIA 105, DECEMBER 2008 15

Brooks, A.R. and L.A. Kelton. 1967. Aquatic and semiaquatic Heteroptera of Alberta, Saskatchewan, and

Manitoba (Hemiptera). Memoirs of the Entomological Society of Canada 51:1-92.

De Clercq, P. 2000. Predaceous stinkbugs (Pentatomidae: Asopinae). Pp. 737-789. Jn Schaefer, C.W. and

A.R. Panizzi (eds.). Heteroptera of Economic Importance. CRC Press, Boca Raton, FL.

Drake, C.J. and H.M. Harris 1934. The Gerrinae of the Western Hemisphere (Hemiptera). Annals of the

Carnegie Museum 23:179-240.

Drake, C.J. and J.D. Lattin. 1963. American species of the lacebug genus Aca/ypta (Hemiptera: Tingidae).

Proceedings of the United States National Museum 115:331-345.

Foldessy, M. 2006. The presence of Leptoglossus occidentalis Heidemann, 1910 (Heteroptera: Coreidae) in

North-east Hungary. Folia Historico Naturalia Musei Matraensis 30:203-204.

Froeschner, R.C. 1960. Cydnidae of the Western Hemisphere. Proceedings of the United States National

Museum 111:337-680.

Gall, W. 1992. Further eastern range extensions of Leptoglossus occidentalis. Great Lakes Entomologist

25:159-171.

Gogala, A. 2003. Listonozka (Leptoglossus occidentalis) ze v Sloveniji (Heteroptera: Coreidae). Acta Ento-

mologica Slovenica 11:189-190.

Harris, H.M. 1928. A monographic study of the hemipterous family Nabidae as it occurs in North America.

Entomologica Americana 9:1-97.

Harris, H.M. and W.E. Shull. 1944. A preliminary list of Hemiptera of Idaho. Iowa State College Journal of

Science 18:199-208.

Hedlin, A.F., H.O. Yates, D.C. Tovar, B.H. Ebel, T.W. Kroeber and E.P. Merkel. 1981. Cone and Seed

Insects of North American Conifers. Canadian Forestry Services, USDA Forest Service and Secretaria de

Agricultura y Recursos Hidraulicos, Mexico. 122 pp.

Henry, T.J. 1979. Review of the Ceratocapsus lutescens group, with descriptions of seven new species from

the eastern United States (Hemiptera: Miridae). Proceedings of the Entomological Society of Washington

81:401-423.

Henry, T.J. and J.D. Lattin 1988. Family Nabidae Costa, 1853. The Damsel Bugs. Pp. 508-520. Jn T.J.

Henry and R.C. Froeschner (eds.). Catalog of the Heteroptera, or True Bugs, of Canada and the Conti-

nental United States. E.J. Brill, Leiden.

Henry, T.J. and A.G. Wheeler, Jr. 1988. Family Miridae Hahn, 1833 (= Capsidae Burmeister, 1835). The

Plant Bugs. Pp. 251-507. Jn T.J. Henry and R.C. Froeschner (eds.). Catalog of the Heteroptera, or True

Bugs, of Canada and the Continental United States. E.J. Brill, Leiden.

Henry, T.J., A.G. Wheeler, Jr. and W.E. Steiner, Jr. 2008. First North American records of Amphiareus

obscuriceps (Poppius) (Hemiptera: Heteroptera: Anthocoridae), with a discussion of dead-leaf microhabi-

tats. Proceedings of the Entomological Society of Washington 110:402-416.

Hilpold, A. 2005. Neu fuer Suedtirol: Leptoglossus occidentalis Heidemann, 1910 (Heteroptera, Coreidae).

Gredleriana 5:358.

Hoebeke, E.R. and A.G. Wheeler, Jr., 1982. Rhopalus (Brachycarenus) tigrinus, recently established in

North America, with a key to the genera and species of Rhopalidae in eastern North America (Hemiptera:

Heteroptera). Proceedings of the Entomological Society of Washington 84:213-224.

Hungerford, H.B. 1933. The genus Notonecta of the world. University of Kansas Science Bulletin 21:5-195.

Hungerford, H.B. 1948. The Corixidae of the Western Hemisphere (Hemiptera). University of Kansas Sci-

ence Bulletin 32:5-827.

Jansson, A. 2002. New records of Corixidae (Heteroptera) from northeastern USA and eastern Canada, with

one new synonymy. Entomologica Fennica 13:85-88.

Kelton, L.A. 1978. The Anthocoridae of Canada and Alaska. Heteroptera: Anthocoridae. The Insects and

Arachnids of Canada. Part 4. Agriculture Canada Research Branch Publication 1639: 101 pp.

Kelton, L.A. 1980. The plant bugs of the Prairie Provinces of Canada. The Insects and Arachnids of Can-

ada. Part 8. Agriculture Canada Research Branch Publication 1703: 408 pp.

Kenner, R.D. and K.M. Needham. 2004. New waterboatmen records for Western Canada (Hemiptera:

Corixidae). Journal of the Entomological Society of British Columbia 101:147-148.

Kerzhner, I.M. 1963. Beitrag zur Kenntnis der Unterfamilie Nabinae (Heteroptera: Nabidae). Acta Ento-

mologica Musei Nationalis Prague 35:5-61.

Kerzhner, I.M. 1993. New and little-known Nabidae from North America (Heteroptera). Zoosystematica

Rossica | (1992):37-45.

Knight, H.H. 1920. New and little-known species of Phytocoris from the eastern United States

(Heteroptera-Miridae). Bulletin of the Brooklyn Entomological Society 15:49-66.

Knight, H.H. 1921. Monograph of the North American species of Deraeocoris (Heteroptera: Miridae). Uni-

versity of Minnesota Agricultural Experimental Station Technical Bulletin 1:75-210.

Knight, H.H. 1927. Descriptions of seven new species of the genus Orthotylus Fieber (Hemiptera, Miridae).

16 J. ENTOMOL. Soc. BRIT. COLUMBIA 105, DECEMBER 2008

The Canadian Entomologist 59:176-181.

Knight, H.H. 1941. The plant bugs, or Miridae, of Illinois. Illinois Natural History Survey Bulletin 22:1-

234.

Knight, H.H. 1968. Taxonomic review: Miridae of the Nevada test site and the western United States. Brig-

ham Young University Science Bulletin (Biology Series) 9(3):1-282.

Knight, H.H. 1974. A key to species of Phytocoris Fallen belonging to the Phytocoris junceus Kngt. group

of species (Hemiptera, Miridae). Iowa State Journal of Research 49:123-135.

Koeber, T.W. 1963. Leptoglossus occidentalis (Hemiptera, Coreidae), a newly discovered pest of conifer-

ous seed. Annals of the Entomological Society of America. 56:229-234.

Krugman, S.L. 1969. Effect of cone feeding by Leptoglossus occidentalis on ponderosa pine seed develop-

ment. Forest Science 15:104-111.

Larochelle, A. 1984. Les Punaises Terrestres (Hemipteres: Géocorises) du Quebec. Fabreries Supplement 3:

513 pp.

Lattin, J.D. 1999. Bionomics of the Anthocoridae. Annual Review of Entomology 44:207-231.

Lattin, J.D. 2000. Minute pirate bugs (Anthocoridae). Pp. 607-637. In C.W. Schaefer and A.R. Panizzi

(eds.). Heteroptera of Economic Importance. CRC Press, Boca Raton, FL.

Lattin, J.D. 2004. Solenonotus angustatus Poppius, a synonym of Dufouriellus ater Fieber (Hemiptera:

Heteroptera: Anthocoridae). Proceedings of the Entomological Society of Washington 106:722-724.

Lattin, J.D. 2005. Dufouriellus ater (Puton), Macrotrachelia nigronitens (Stal) and Xylocoris (Arrostelus)

flavipes (Reuter) (Hemiptera: Heteroptera: Cimicoidea: Anthocoridae): first records from the Hawaiian

Islands. Proceedings of Entomological Society of Washington 107:466-468.

Lattin, J.D. 2007a. The Lasiochilidae, Lyctocoridae, and Anthocoirdae (Hemiptera: Heteroptera) of the

Hawaiian Islands: native or introduced? Proceedings of Entomological Society of Washington 109:75-80.

Lattin, J.D. 2007b. The non-indigenous Lyctocoridae and Anthocoridae (Hemiptera: Heteroptera; Cimi-

coidea) of America north of Mexico. Proceedings of Entomological Society of Washington 109:366-376.

Lewis, T.M., D.R. Horton and D.A. Broers. 2005. New state and United States records for Anthocoridae

(Hemiptera: Heteroptera). Pan-Pacific Entomologist 81:59-67.

Lindskog, P. 1981. On the identity of Saldula laticollis (Reuter) (Heteroptera: Saldidae). Pan-Pacific Ento-

mologist 57:321-326.

Marshall, S.A. 1991. A new Ontario record of a seed eating bug (Hemiptera: Coreidae) and other examples

of the role of regional insect collections in tracking changes in Ontario’s fauna. Proceedings of the Ento-

mological Society of Ontario 122:109-111.

Matsuda, R. 1977. The Aradidae of Canada. Hemiptera: Aradidae. The Insects and Arachnids of Canada.

Part 3. Research Branch Canada Department of Agriculture Publication. 1634: 116 pp.

Maw, H.E.L, R.G. Foottit, K.G.A. Hamilton and G.G.E. Scudder. 2000. Checklist of the Hemiptera of Can-

ada and Alaska. NRC Research Press, Ottawa. 220 pp.

McDonald, F.J.D. 1986. Revision of Cosmopepla Stal (Hemiptera: Pentatomidae). Journal of the New York

Entomological Society 94:1-15.

McPherson, J.E. 1982. The Pentatomoidea (Hemiptera) of northeastern North America, with emphasis on

the fauna of Illinois. Southern IIlinois University Press, Carbondale and Edwardsville. 240 pp.

McPherson, J.E., R.J. Packauskas, S.F. Taylor and M.F. O’Brien. 1990. Eastern range extension of Lep-

toglossus occidentalis, with a key to Leptoglossus species of America north of Mexico (Heteroptera:

Coreidae). Great Lakes Entomologist 23:99-104.

Mitchell, P.L. 2000. Leaf-footed bugs (Coreidae). Pp 337-403. In C.W. Schaefer and A.R. Panizzi (eds.).

Heteroptera of Economic Importance. CRC Press, Boca Raton, FL.

Moulet, P. 2006. Un nouveau Coreide en France: Leptoglossus occidentalis Heidemann, 1910 (Heteroptera

Coreidae). Entomologiste, Paris 62:183-184.

Paiero, S.M., S.A. Marshall and K.G.A. Hamilton. 2003. New records of Hemiptera from Canada and On-

tario. Journal of the Entomological Society of Ontario 134:115-129.

Panizzi, A.R., J.E. McPherson, D.G. James, M. Javahery and McPherson, R.M. 2000. Stink Bugs

(Pentatomidae). Pp. 421-474. In Schaefer, C.W. and A.R. Panizzi (eds.). Heteroptera of Economic Impor-

tance. CRC Press, Boca Raton, FL.

Polhemus, D.A. 1994. An annotated checklist of the plant bugs of Colorado (Heteroptera: Miridae). Pan-

Pacific Entomologist 70:122-147.

Polhemus, J.T. 1985. Shore Bugs (Heteroptera, Hemiptera; Saldidae). A world overview and taxonomy of

Middle American forms. The Different Drummer, Englewood, Co. 252 pp.

Polhemus, J.T. 1988. Family Saldidae Amyot and Serville, 1843. The Shore Bugs. Pp. 665-681. Jn T.J.

Henry and R.C. Froeschner (eds.). Catalog of the Heteroptera, or True Bugs, of Canada and the Conti-

nental United States. E.J. Brill, Leiden.

Polhemus, J.T. and H.C. Chapman. 1979. Family Saldidae / Shore Bugs. Pp. 16-33. Jn A.S. Menke (ed.).

J. ENTOMOL. SOc. BRIT. COLUMBIA 105, DECEMBER 2008 17

The Semiaquatic and Aquatic Hemiptera of California (Heteroptera: Hemiptera). Bulletin of the Califor-

nia Insect Survey. Volume 21.

Rabitsch, W. and E. Heiss. 2005. Leptoglossus occidentalis Heidemann, 1910, eine amerikanische Adventi-

vart auch in Oesterreich aufgefunden (Heteroptera: Coreidae). Berichte des Naturwissenschaftlich Mediz-

inischen Vereins in Innsbruck 92:131-135.

Ribes, J. and E.-B. Oleguer. 2005. Leptoglossus occidentalis Heidemann, 1910, hemipter nearctic trobat a

Catalunya (Hemiptera: Heteroptera: Coreidae). Sessio Conjunta d’ Entomologia ICHN SCL 13 (2003):47-

50.

Ridge-O’Connor, G.E. 2001. Distribution of the western conifer seed bug, Leptoglossus occidentalis Heide-

mann (Heteroptera: Coreidae) in Connecticut and parasitism by a tachinid fly, Trichopoda pennipes (F.)

(Diptera: Tachinidae). Proceedings of the Entomological Society of Washington 103:364-366.

Roch, J.-F. 2007. Liste des punaises du Québec et des Régions adjacentes (Hemiptera: Heteroptera). Ento-

mofauna du Québec Document Faunique 27: 42 pp.

Rolston, L.H. 1973. The genus Menecles Stal (Hemiptera: Pentatomidae). Journal of the New York Ento-

mological Society 80:234-237.

Rolston, L.H. 1983. A revision of the genus Acrosternum Fieber, subgenus Chinavia Orian, in the Western

Hemisphere (Hemiptera: Pentatomidae). Journal of the New York Entomological Society 91:97-176.

Ruth, D.S., G.E. Miller and J.R. Sutherland. 1982. A guide to common insect pests and diseases in spruce

seed orchards in British Columbia. Canadian Forestry Service, Environment Canada. BC-X-231.

Schaefer, C.W. and N.P. Chopra. 1982. Cladistic analysis of the Rhopalidae, with a list of food plants. An-

nals of the Entomological Society of America 75:224-233.

Schaefer, C.W. and P.L. Mitchell. 1983. Food plants of the Coreidae (Hemiptera: Heteroptera). Annals of

the Entomological Society of America 76:591-615.

Schaffner, J.C. 1967. The occurrence of Theognis occidentalis in the midwestern United States

(Heteroptera: Coreidae). Journal of the Kansas Entomological Society 40:141-142.

Schowalter, T.D. and J.M. Sexton. 1990. Effect of Leptoglossus occidentalis (Heteroptera: Coreidae) on

seed development of Douglas fir at different times during the growing season in western Oregon. Journal

of Economic Entomology 83:1485-1486.

Schuh, R.T. 2000a. Revision of Oligotylus Van Duzee with descriptions of ten new species from western

North America and comments on Lepidargyrus in the Nearctic (Heteroptera: Miridae: Phylinae: Phylini).

American Museum Novitates 3300: 44 pp.

Schuh, R.T. 2000b. Revision of the North American plant bug genus Megalopsallus Knight, with descrip-

tion of eight new species from the west (Heteroptera: Miridae: Phylinae). American Museum Novitates

3305: 69 pp.

Schuh. R.T. 2001. Revision of New World Plagiognathus Fieber, with comments on the Palearctic fauna

and the description of a new genus (Heteroptera: Miridae: Phylinae). Bulletin of the American Museum

of Natural History 266: 267 pp.

Schuh, R.T. 2004a. Revision of Tuxedo Schuh (Hemiptera: Miridae: Phylinae). American Museum Novi-

tates 3435: 26 pp.

Schuh, R.T. 2004b. Revision of Europiella Reuter in North America, with the description of a new genus

(Heteroptera; Miridae: Phylinae). American Museum Novitates 3463: 58 pp.

Schuh. R.T. and M.D. Schwartz. 1988. Revision of the New World Pilophorini (Heteroptera: Miridae:

Phylinae). Bulletin of the American Museum of Natural History 187:101-201.

Schuh, R.T. and M.D. Schwartz. 2004. New genera, new species, new synonyms, and new combinations in

North American and Caribbean Phylinae (Heteroptera: Miridae). American Museum Novitates 3436: 36

Pp.

Schuh, R.T. and M.D. Schwartz. 2005. Review of North American Chlamydatus Curtis species, with new

synonymy and the description of two new species (Heteroptera: Miridae: Phylinae). American Museum

Novitates 3471: 55 pp.

Schuh, T. 1967. The shore bugs (Hemiptera: Saldidae) of the Great Lakes region. Contributions of the

American Entomological Institute 2(2): 35 pp.

Schwartz, M.D. and G.G.E. Scudder 2001. Miridae (Heteroptera) new to Canada, with some taxonomic

changes. Journal of the New York Entomological Society 108 (2000):248-267.

Schwartz, M.D. and G.G.E. Scudder. 2003. Seven new species of Miridae (Heteroptera) from British Co-

lumbia and Alaska and synonymy of Adelphocoris superbus (Uhler). Journal of the New York Entomo-

logical Society 111:65-95.

Schwartz, M.D. and G.M. Stonedahl. 2004. Revision of Phoenicocoris Reuter with descriptions of three

new species from North America and a new genus from Japan (Heteroptera: Miridae: Phylinae). Ameri-

can Museum Novitates 3464: 55 pp.

Scudder, G.G.E. 1962. The Ischnorhynchinae of the world (Hemiptera: Lygacidae). Transactions of the

18 J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

Royal Entomological Society of London 114:163-194.

Scudder, G.G.E. 1977. An annotated checklist of the aquatic and semiaquatic Hemiptera (Insecta) of British

Columbia. Syesis 10:31-38.

Scudder, G.G.E. 1997. True Bugs (Heteroptera) of the Yukon. Pp. 241-336. Jn H.V. Danks and J.A. Dow-

nes (eds.). Insects of the Yukon. Biological Survey of Canada (Terrestrial Arthropods), Ottawa.

Scudder, G.G.E. 2000. Heteroptera (Hemiptera: Prosorrhyncha) new to Canada. Part. 1. Journal of the Ento-

mological Society of British Columbia 97:51-56.

Scudder, G.G.E. 2004. Heteroptera (Hemiptera: Prosorrhyncha) new to Canada. Part 2. Journal of the Ento-

mological Society of British Columbia 101:125-129.

Scudder, G.G.E. 2007. Two alien Heteroptera (Hemiptera) new to Canada. Journal of the Entomological

Society of British Columbia 104:81-84.

Scudder, G.G.E. and S.S. Duffey. 1972. Cardiac glycosides in the Lygaeinae (Hemiptera: Lygaeidae). Ca-

nadian Journal of Zoology 50:35-42.

Scudder, G.G.E. and R.G. Foottit 2006. Alien true bugs (Hemiptera: Heteroptera) in Canada: composition

and adaptations. The Canadian Entomologist 138:24-51.

Scudder, G.G.E. and M.D. Schwartz 2001. The genus Leptopterna Fieber (Heteroptera: Miridae:

Stenodemini) in North America. Proceedings of the Entomological Society of Washington 103(4):797-

806.

Slater, A. 1988. A new genus and two new species of Lygaeinae (Heteroptera: Lygaeidae). Journal of the

Kansas Entomological Society 61:308-316.

Slater, J.A. 1954. Notes on the genus Labops Burmeister in North America, with the description of three

new species (Hemiptera: Miridae). Bulletin of the Brooklyn Entomological Society 49:57-65, 89-94.

Slater, J.A. and R.M. Baranowski 1978. How to know the true bugs (Hemiptera-Heteroptera). Wm. C.

Brown Company Publishers, Dubuque, Iowa. 256 pp.

Stock, M.W. and J.D. Lattin. 1976. Biology of intertidal Sa/dula palustris (Douglas) on the Oregon coast

(Heteroptera: Saldidae). Journal of the Kansas Entomological Society 49:313-326.

Strong, W.B. 2006. Seasonal changes in seed reduction in Lodgepole pine cones caused by feeding of Lep-

toglossus occidentalis (Hemiptera: Coreidae). The Canadian Entomologist 138:888-896.

Strong, W.B., S.L. Bates and M.U. Stoehr. 2001. Feeding by Leptoglossus occidentalis Heidemann

(Hemiptera: Coreidae) reduces seed set in lodgepole pine. The Canadian Entomologist 133:857-865.

Sweet, M.H. 1964. The biology and ecology of the Rhyparochrominae of New England (Heteroptera: Ly-

gaeidae). Part II. Entomologica Americana (new series) 44:1-201.

Taylor, S.J., G. Tescari and M. Villa. 2001. A nearctic pest of Pinaceae accidentally introduced into Europe:

Leptoglossus occidentalis (Heteroptera: Coreidae) in northern Italy. Entomological News 112:101-103.

Tescari, G. 2004. First record of Leptoglossus occidentalis (Heteroptera: Coreidae) in Croatia. Entomologia

Croatica 8:73-75.

Thomas, D.B. 1992. Taxonomic synopsis of the Asopine Pentatomidae (Heteroptera) of the Western Hemi-

sphere. The Thomas Say Foundation Volume 16: 156 pp.

Tinerella, P.P. and R.W. Gundersen 2005. The water boatmen (Insecta: Heteroptera: Corixidae) of Minne-

sota. Schafer-Post Series: North Dakota Insects Publication 23: 119 pp.

Van Duzee, E.P. 1916. Checklist of the Heteroptera (excepting the Aphidiae, Aleurodidae and Coccidae) of

America, north of Mexico. New York Entomological Society, New York. 111 pp.

Van Duzee, E.P. 1917. Catalogue of the Hemiptera of America north of Mexico, excepting the Aphididae,

Coccidae and Aleurodidae. University of California Publications, Technical Bulletins, Entomology 2:

902 pp.

Vinokurov, N.N. 1988. Heteroptera of Yakutia. Oxonian Press Pvt. Ltd., New Delhi, Calcutta. 328 pp.

Wheeler, Jr., A.G., and T.J. Henry 1992. A Synthesis of the Holarctic Miridae (Heteroptera): Distribution,

Biology, and Origin, with Emphasis on North America. The Thomas Say Foundation Volume 15: 282 pp.

Wheeler, Jr., A.G., T.J. Henry and E.R. Hoebeke 2006. Palearctic plant bugs (Hemiptera, Miridae) in New-

foundland, Canada: First North American records for Phytocoris longipennis Flor and Pilophorus cin-

namopterus (Kirschbaum), new records of eight other species, and review of previous reported species.

Denisia 19 (2006):997-1014.

Wheeler, Jr., A.G., T.J. Henry and T.L. Mason, Jr. 1983. An annotated list of the Miridae of West Virginia

(Hemiptera-Heteroptera). Transactions of the American Entomological Society 109:127-159.

Wheeler, Jr., A.G. and E.R. Hoebeke. 2004. New records of Palearctic Hemiptera (Sternorrhyncha, Cicado-

morpha, Heteroptera) in the Canadian maritime provinces. Proceedings of the Entomological Society of

Washington. 106:298-304.

Wright, B. 1989. The fauna of Sable Island. Nova Scotia Museum Curatorial Report 68: 93 pp.

J. ENTOMOL. Soc. BRIT. COLUMBIA 105, DECEMBER 2008

First Canadian records of Lampropteryx suffumata

({[Denis & Schiffermiiller], 1775) (Geometridae: Larentiinae)

JEREMY R. DEWAARD””, B. CHRISTIAN SCHMIDT?,

GARY G. ANWEILER* and LELAND M. HUMBLE'”

ABSTRACT

The first Canadian records of the Holarctic species Lampropteryx suffumata ({Denis &

Schiffermiiller], 1775) are documented, based on collections from Alberta and British

Columbia. Widespread and common throughout much of Eurasia, the larvae feed on

Galium species (Rubiaceae). Diagnostic descriptions and images are provided to aid in

future recognition of this species. The specimens were originally detected while con-

structing a DNA barcoding library for western North American Geometridae, and pro-

vide a good example of how genetic methods can enhance the construction of regional

inventories and aid in surveillance for invasive species.

Key Words: Lampropteryx suffumata, black-banded carpet, DNA barcoding, invasive

species

INTRODUCTION

The genus Lampropteryx Stephens in-

cludes ten species, most of which are re-

stricted to Asia, with two species also oc-

curring in Europe (Scoble 1999). The

black-banded carpet Lampropteryx suffu-

mata ({Denis & Schiffermiiller], 1775),

described from Vienna, Austria, occurs

from western Europe and the northern

Mediterranean region to northern Scandina-

via, east through the Tien Shan and Altai

mountain ranges of south-central Asia to

the Kamchatka Peninsula, Russia and Hok-

kaido, Japan (Skou 1986; Beljaev and Vasi-

lenko 2002). Previously known in North

America only from Alaska (Choi 2000), we

report here historical and contemporary

records in British Columbia and Alberta,

flagged by DNA barcoding.

MATERIALS AND METHODS

During the course of documenting the

molecular diversity of western Canadian

geometrid moths from museum and field

collections using standard DNA barcoding

methods (Hajibabaei et al. 2005; deWaard

et al. 2008), it became evident that a num-

ber of specimens variously identified as

Antepirrhoe Warren or Xanthorhoe Hibner

were highly divergent compared to other

congeners. Using the identification engine

of the Barcode of Life Database (BOLD)

(Ratnasingham and Hebert 2007), their cy-

tochrome oxidase I (COI) sequences were a

nearly identical match to those of Lamprop-

teryx suffumata ({[Denis & Schiffermiiller],

1775) specimens from Bavaria, Germany

University of British Columbia, Department of Forest Sciences, Forestry Sciences Centre, Vancouver,

BC, Canada V6T 1Z4

? Royal British Columbia Museum, Entomology, 675 Belleville Street, Victoria, BC, Canada V8W 9W2

> Canadian Food Inspection Agency, Canadian National Collection of Insects, Arachnids and Nematodes,

KW. Neatby Bldg., 960 Carling Ave., Ottawa, ON, Canada K1A 0C6

: E.H. Strickland Entomological Museum, University of Alberta, Edmonton, AB, Canada T6G 2E9

Natural Resources Canada, Canadian Forest Service, Pacific Forestry Centre, 506 West Burnside Road,

Victoria, BC, Canada V8Z 1M5

20 J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

(Figure 1). Although many larentiines are

very similar in habitus and are difficult to

identify when the wing pattern 1s worn,

subsequent genitalic examination of the

suspect Antepirrhoe and Xanthorhoe speci-

mens showed unequivocally that they are in

fact L. suffumata.

To determine the Canadian distribution,

and whether or not the species is likely na-

tive, we examined historical and contempo-

rary Antepirrhoe and Xanthorhoe speci-

mens from various Canadian collections.

We identified specimens in the Royal Brit-

ish Columbia Museum, Victoria, BC

(RBCM), the E.H. Strickland Entomolog-

cal Museum, University of Alberta, Edmon-

ton, AB (UASM) the Canadian National

Collection of Insects, Agriculture and Agri-

Food Canada, Ottawa, ON (CNC), and the

Biodiversity Institute of Ontario, University

of Guelph, Guelph, ON (BIOUG) as L. suf-

fumata. The collections of the Pacific For-

estry Centre, Canadian Forest Service, Vic-

toria, BC (PFCA), the Spencer Entomologi-

cal Museum, University of British Colum-

bia, Vancouver, BC (UBCZ), and the

Northern Forestry Centre, Canadian Forest

Service, Edmonton, AB (NFRC) do not

contain any specimens of L. suffumata.

RESULTS

Specimens examined (all specimens are

single, pinned adults; the BOLD accession

number (italicized) is provided for speci-

mens that have been barcoded).

AB: Hillcrest, 49.568N 114.377W, 20-

vi-1919 (K. Bowman) [UASM,

UASM10792]; West Castle River, W Cas-

tle R. Rd., 15 km SW, 49.294N 114.273W,

23-v-1999 (B.C. Schmidt) [CNC,

CNCLEP00033310, GWNC311-07]; BC:

Elkford, 35 km north, 50.266N 114.921W,

12-Jun-1988 (C.S. Guppy) [RBCM,

ENT991-006550, GWNR470-07]; Glacier

National Park, Abandoned Rails Trail west

of Rogers Pass Centre, 51.2902N

117.516W, 04-Jul-2005 (K. Pickthorn)

[BIOUG, HLC-20568, LBCA568-05]; Gla-

cier National Park, Glacier National Park

Compound at Rogers Pass, 51.3032N

117.519W, 28-Jun-2005 (K. Pickthorn)

[BIOUG, HLC-20320, LBCA320-05]; Gla-

cier National Park, Illecillewaet Camp-

grounds west of Rogers Pass, 51.2648N

117.494W, 24-Jun-2005 (K. Pickthorn)

[BIOUG, HLC-20175, LBCA175-05]; Gla-

cier National Park, Glacier National Park

Compound at Rogers Pass, 51.3032N

117.519W, 16-Jun-2005 (K. Pickthorn)

[BIOUG, HLC-20022, LBCA022-05]; Trin-

ity Valley Field Station, 50.400N

118.917W, 18-May-1961 (W.C. McGuffin)

[CNC, CNCLEP00054030].

Identification. A medium-sized, broad-

winged moth with a wingspan of 2.5—3.2

cm (Figure 2a.). The forewing basal and

median bands are dark, varying from red-

brown to black, being dark brown in most

specimens. The median band has a jagged

proximal and distal margin, with the distal

margin extending towards the base just be-

low the median area, such that the median

band is narrower along the anal third than

on the upper half. The apex is also dark-

ened, divided by a white apical dash. There

is a subterminal line of white spots or

wedges, and the fringe is checkered. It is

very similar to Antepirrhoe semiatrata

(Hulst), but can be readily separated by the

following characters: forewing pale

antemedian band faintly bordered with two

whitish lines both proximally and distally

(only one pale border line in A. semiatrata);

forewing subapical dark patch bordered

towards costal margin by contrasting pale

line (indistinctly so in A. semiatrata). The

dorsal markings on the abdomen are the

most reliable external features for diagnos-

ing L. suffumata, which has a row of black

triangles along the midline (Fig. 2a),

whereas Antepirrhoe species have two

black dots broken at the midline by a pale

line/spot. Some specimens may be melanic

and lack the contrasting white forewing

bands present in most specimens. Xan-

thorhoe species are superficially similar,

but lack the combination of broad, dark

J. ENTOMOL. Soc. BRIT. COLUMBIA 105, DECEMBER 2008

pas

CNCLEP00033427 - Antepirrhoe atrifasciata (Canada: BC)

CNCLEP00033426 - A. fasciata (United States: Washington)

CNCLEP00033425 - A. semiatrata (Canada: NB)

O HLC-20320 (Canada: BC)

O HLC-20022 (Canada: BC)

Lampropteryx

suffumata

O CNCLEP00033310 (Canada: AB)

O HLC-20175 (Canada: BC)

AYK-06-7334 - L. minna (Japan: Chubu)

BC ZSM Lep 13521 - L. jameza (China: Hebei)

Figure 1. Neighbour-joining tree of Lampropteryx suffumata and related species. Tree was

reconstructed with the barcode fragment of the COI gene. Sequences shaded in grey are de-

rived from specimens previously misidentified as Antepirrhoe or Xanthorhoe spp. The 13 se-

quences are publicly available in the Barcode of Life Database and GenBank (accession nos.

FJ37663 1-FJ3 76643).

basal and median bands with a contrast-

ingly bordered subapical dark patch that

extends to the distal wing margin. Genitalic

examination of L. suffumata will easily

segregate this species: the male valve is

simple and lobe-shaped, costa lacking api-

cal process; socii prominent, about half as

long as valve, with bundle of apical setae as

long as socius; aedeagus uncurved, vesica

with two cornuti (Figure 2b,c.). Identifica-

tion through genitalic examination of males

can usually be made by brushing away the

terminal abdominal scales to reveal the api-

cal portion of the valve which lacks the

pointed, dorsally projecting costal process

of A. semiatrata, in addition to the long

tubular socii (stout and triangular without

apical hair pencils in A. semiatrata). Male

genitalic structure of Xanthorhoe species is

very different, with a comparatively mas-

Sive costal process that extends beyond the

valve apex and is variously enlarged, broad-

ened and/or armed with spines.

Distribution and Habitat: Great Brit-

ain and northern Europe east to southern

Siberia, Kamchatka and Japan (Skou 1986;

Beljaev and Vasilenko 2002); in North

America, known from two areas: Alaska

(Choi 2000) and southwestern British Co-

lumbia and adjacent Alberta (Figure 3). It is

likely that this species occurs in intervening

regions of northern British Columbia and

the Yukon, but these areas have not been

adequately surveyed. The single historical

collection from Hillcrest, Alberta, coupled

with the fact that L. suffumata occurs in

relatively remote, mountainous habitats but

has not been recorded near the international

shipping ports of the coastal Pacific North-

west, suggests that L. suffumata is native to

Canada. Furthermore, it likely expanded

over Beringia during the Pleistocene, a

common pattern in the western Canadian

arthropod fauna, as evident by present

ranges and fossil evidence of past ranges

(Danks et al. 1997). Its habitat appears to be

open wooded areas, edges and meadows.

Life History and Notes: There is a sin-

22 J. ENTOMOL. SOc. BRIT. COLUMBIA 105, DECEMBER 2008

0.5mm

Figure 2. Adult male of Lampropteryx suffumata: a) dorsal view b) genital capsule c)

aedeagus.

gle annual brood, with adults in late May to

early July. Adults are nocturnal and come

to light. The only reported larval hosts are

bedstraw species (Galium sp.), particularly

G. aparine Linnaeus (Skou 1986). The

pupa overwinters underground (Skou

1986). Based on the scarcity of specimens

in Canadian collections, we conclude the

species is rarely collected and likely rare.

The COI barcode sequences are publicly

available in the Barcode of Life Database

and GenBank (accession nos. FJ376631—

FJ376643).

DISCUSSION

The late discovery of a relatively large

and conspicuous native macromoth in

Western North America is surprising, but

we believe it can be explained simply by

the paucity of taxonomic expertise and lit-

erature on the group. The Canadian larenti-

ines are notoriously hard to discriminate,

due in part to the lack of a treatment of this

J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

ao ATER i ae aI a iB ehPPEIL L ADI PEE BE OAS BE Rae PA RETR ETE i]

a Banff National Park | fe

; ' Yoho yn ;

Glacier ‘National Lake Louise AK

National) Park i"

i Fath ee NE Banff

i, Golden i -

Mt, Revelstoke: viek 'y

National Le «ll

rie ols Keatcney \ | . es

Park ‘National po, are

va Park ; bo i

Revelstoke | “eR

Invermere’:

Elkford*

" Nakusp

eLumby

4 Coleman

Kimberley °

‘ ‘Fernie \

Cranbrook a .

Waketor Lakes

Natromg!

Park

hyte ee

ne ae

Figure 3. Distribution of Lampropteryx suffumata in North America. Black squares are loca-

tions of records.

subfamily in McGuffin’s ‘Guide to the Ge-

ometridae of Canada’ series (1967, 1972,

1977, 1981, 1987, 1988). While a few lar-

entiine genera have been revised

(Hydriomena Hibner: McDunnough 1954;

Eupithecia Curtis: Bolte 1990; Entephria

Htibner: Troubridge 1997), most are in dire

need of revision, and the Xanthorhoini in

particular contain a number of genera that

need attention, with cryptic and previously

unrecognized species awaiting description

(e.g., Psychophora Kirby, Xanthorhoe and

Zenophleps Hulst: B.C.S. unpublished data;

Antepirrhoe: J.R.D. et al. unpublished

data). It is reasonable to assume that the

few specimens of this rarely collected (and

presumably rare) species could go unno-

ticed due to the lack of reliable guides and

keys for the group.

Although L. suffumata is in all likeli-

hood native, its discovery clearly illustrates

how DNA barcoding can assist in the detec-

tion and surveillance of nonindigenous or-

ganisms (Armstrong and Ball 2005; Chown

et al. 2008). A monitoring program that

incorporates DNA barcoding can flag po-

tential introduced species in one of two

ways. First, as in this study, a barcode

match is made with one or more specimens

collected from the native range. The poten-

tial nonindigenous specimens can then be

verified by morphological examination or

further genetic analysis. At that point, na-

tional and regional collections can be exam-

ined for historical and contemporary speci-

mens in the new range to determine if the

Species is native or introduced. Secondly,

with a barcode library for a regional fauna

complete (e.g., Geometridae of British Co-

lumbia — J.R.D. et al. unpublished data),

any barcoded specimens that do not match

the database are flagged as potentially non-

indigenous and again warrant further ex-

amination. Using genetic methods for this

initial screening has numerous advantages,

most notably the ability to differentiate spe-

cies objectively across all life stages as well

as using damaged specimens. It is also ap-

24 J. ENTOMOL. Soc. BRIT. COLUMBIA 105, DECEMBER 2008

parent that with the current costs of genetic

analysis steadily dropping and new tech-

nologies emerging (Hajibabaei et al. 2007),

genetic screening may soon be more cost-

and time-efficient than current morphologi-

cal methods of biodiversity monitoring.

ACKNOWLEDGEMENTS

This study was facilitated by the funding

provided by the RBCM as part of the

NSERC-RBCM — Systematics Research

Graduate Supplement program; for that, we

are grateful to Rob Cannings, Kelly Sendall

and Grant Hughes. Additional funding was

provided by a Forest Investment Account -

Forest Science Program Student Grant and

NSERC Graduate Scholarship (to JRD).

Molecular analyses were supported by the

Canadian Barcode of Life Network from

Genome Canada through the Ontario Ge-

nomics Institute, NSERC (to LMH), and

other sponsors listed at www.BOLNET.ca.

We wish to thank Axel Hausmann

(Zoological State Collection Munich) and

Kim Mitter (University of Maryland) for

allowing us access to the Eurasian se-

quences, Allison Shaver (Agriculture and

Agri-Food Canada) for slide preparation

and photography, Gurp Thandi (Natural

Resources Canada, Pacific Forestry Centre)

for preparing the distribution map, the staff

at the Canadian Centre for DNA Barcoding

for their assistance with genetic analysis,

and two anonymous reviewers for helpful

comments on the manuscript. We are also

grateful to Claudia Copley (RBCM), Karen

Needham (UBCZ), Felix Sperling (UASM),

Don Lafontaine (CNC), Jean-Francois

Landry (CNC), Alex Borisenko (BIOUG),

and Greg Pohl (NFRC) for access to the

respective collections in their care.

REFERENCES

Armstrong, K.F., and S.L. Ball. 2005. DNA barcodes for biosecurity: invasive species identification. Phi-

losophical Transactions: Biological Sciences 360: 1813-1823.

Beljaev, E.A. and S.V. Vasilenko. 2002: An annotated checklist of geometrid moths (Lepidoptera: Geomet-

ridae) from the Kamchatka Peninsula and adjacent islands. Entomologica Fennica 13: 195-235.

Bolte, K.B. 1990. Guide to the Geometridae of Canada (Lepidoptera). VI. Subfamily Larentiinae. 1. Revi-

sion of the genus Eupithecia. Memoirs of the Entomological Society of Canada 151: 1-253.

Choi, S.-W. 2000. The occurrence of Lampropteryx suffumata (Denis and Schiffermiiller) (Lepidoptera:

Geometridae) in North America. Pan-Pacific Entomologist 76: 123-125.

Chown, S.L., B.J. Sinclair, and B. Jansen van Vuuren. 2008. DNA barcoding and the documentation of

alien species establishment on sub-Antarctic Marion Island. Polar Biology 31: 651-655.

Danks, H.V., J.A. Downes, D.J. Larson, and G.G.E. Scudder. 1997. Insects of the Yukon: characteristics

and history. In Insects of the Yukon (Eds: H.V. Danks and J.A. Downes; Biological Survey of Canada

(Terrestrial Arthropods), Ottawa). Pp. 963-1013.

deWaard, J.R., N.V. Ivanova, M. Hajibabaei, and P.D.N. Hebert. 2008. Assembling DNA Barcodes: Ana-

lytical Protocols. In Methods in Molecular Biology: Environmental Genetics (Ed: C. Martin; Humana

Press Inc., Totowa, USA). Pp. 275-293.

Hajibabaei, M., J.R. deWaard, N.V. Ivanova, S. Ratnasingham, R. Dooh, S.L. Kirk, P.M. Mackie, and

P.D.N. Hebert. 2005. Critical factors for the high volume assembly of DNA barcodes. Philosophical

Transactions: Biological Sciences 360: 1959-1967.

Hajibabaei M., G.A. Singer, E.L. Clare, and P.D.N. Hebert. 2007. Design and applicability of DNA arrays

and DNA barcodes in biodiversity monitoring. BMC Biology 5: 24.

McDunnough, J. 1954. The species of the genus Hydriomena occurring in America north of Mexico

(Lepidoptera: Geometridae) American Museum Novitates. No. 1592. 17 pp.

McGuffin, W.C. 1967. Guide to the Geometridae of Canada (Lepidoptera). I. Subfamily Sterrhinae. Mem-

oirs of the Entomological Society of Canada 50: 1—67.

McGuffin, W.C. 1972. Guide to the Geometridae of Canada (Lepidoptera). II. Subfamily Ennominae. 1.

Memotrs of the Entomological Society of Canada 86: 1-159.

McGuffin, W.C. 1977. Guide to the Geometridae of Canada (Lepidoptera). III. Subfamily Ennominae. 2.

Memoirs of the Entomological Society of Canada 101: 1-191.

J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008 25

McGuffin, W.C. 1981. Guide to the Geometridae of Canada (Lepidoptera). IV. Subfamily Ennominae. 3.

Memoirs of the Entomological Society of Canada 117: 1-153.

McGuffin, W.C. 1987. Guide to the Geometridae of Canada (Lepidoptera). V. Subfamily Ennominae. 4.

Memoirs of the Entomological Society of Canada 138: 1-182.

McGuffin, W.C. 1988. Guide to the Geometridae of Canada (Lepidoptera). II, [V, and V. Subfamilies Ar-

chiearinae, Oenochrominae, and Geometrinae. Memoirs of the Entomological Society of Canada 145: 1—

56.

Ratnasingham, S. and P.D.N. Hebert. 2007. BOLD: The Barcode of Life Data System (http://

www .barcodinglife.org). Molecular Ecology Notes 7: 355-364.

Sato, R. and M. Kameda. 1997. Discovery of Lampropteryx suffumata (Denis & Schiffermiiller)

(Geometridae: Larentiinae) from Hokkaido, Japan. Yugato 148: 33-37.

Scoble, M.J. (ed.). 1999. A taxonomic catalogue to the Geometridae of the world (Insecta: Lepidoptera). 2

vols. CSIRO Publications. 1016 pp.

Skou, P. 1986. The Geometroid Moths of North Europe (Lepidoptera: Drepanidae and Geometridae). Scan-

dinavian Science Press. 348 pp.

Troubridge, J.T. 1997. Revision of the Nearctic species of the genus Entephria Hiibner (Lepidoptera: Ge-

ometridae, Larentiinae). Entomologica Scandinavica 28: 121-139.

J. ENTOMOL. SOc. BRIT. COLUMBIA 105, DECEMBER 2008

Identification of new aphid vector species

of Blueberry scorch virus

D. THOMAS LOWERY '!, MICHAEL G. BERNARDY|,

ROBYN M. DEYOUNG! and CHRIS J. FRENCH!

ABSTRACT

Transmission of Blueberry scorch virus (BIScV) by the aphid species Ericaphis fimbri-

ata (Richards), Aphis spiraecola (Patch), Aphis pomi DeGeer, Acyrthosiphon pisum

(Harris), Myzus ornatus Laing, Aphis helianthi Monell, Myzus persicae (Sulzer), and

Rhopalosiphum padi (L.), was studied in the laboratory using timed aphid acquisition

feeding periods and known numbers of aphid vectors. Successful infection of Nicotiana

occidentalis Wheeler (Solanaceae), a newly identified herbaceous host, and highbush

blueberry, Vaccinium corymbosum L. (Ericaceae), following brief virus-acquisition

feeds lasting less than 5 min, demonstrated that BIScV was transmitted in a non-

persistent, non-circulative manner. Based on transfer of 10 aphids per plant, the most

efficient vector of BIScV from infected to healthy N. occidentalis was M. ornatus. Com-

pared with this herbaceous host, infection rates for blueberry were much lower even

though higher numbers of aphids (25/plant) were used. The highest rate of infection for

blueberry (20%) was achieved when the green colour form of E. fimbriata was used to

transmit the virus. The relatively low rate of transmission from infected to healthy blue-

berry suggests that BIScV would spread slowly in the field. Planting of certified virus-

free nursery material and aggressive removal of infected plants should help control this

Dae |

economically important disease of highbush blueberries.

Key Words: Blueberry scorch virus, aphid vectors, virus transmission

INTRODUCTION

Blueberry scorch virus (BlScV) was

first reported in New Jersey in the late

1970's as Sheep Pen Hill disease of

highbush blueberry, Vaccinium corymbo-

sum (L.) (Ericaceae) (Podleckis and Davis

1989). Several distinct strains infect

highbush blueberry in the northeastern and

northwestern United States and southwest-

erm British Columbia (Cavileer et al. 1994,

Catlin and Schloemann 2004, Bernardy et

al. 2005, Wegener et al. 2006). BIScV has

also been recently reported from Europe

(Ciuffo et al. 2005). Depending on the virus

strain and blueberry cultivar, infection can

result in a wide range of symptoms. While

some varieties are tolerant to certain strains

and display no visible symptoms, infection

with other strains can result in severe necro-

sis of new leaves, twigs and flower clusters

and almost complete loss of yield over time

(Martin and Bristow 1988, Catlin and

Schloemann 2004, Wegener ef al. 2006).

The latent period between infection and

development of symptoms for established

plants is thought to be one to two years

(Caruso and Ramsdell 1995).

There are relatively few previous studies

on BlScV; these mostly relate to detection,

symptomology and strain differentiation.

Although little is currently known about the

insect vectors of BIScV, carlaviruses as a

group are transmitted primarily by aphids in

a non-persistent, non-circulative manner

(Ng and Perry 2004). Non-persistent virus

transmission is characterized by short ac-

quisition and inoculation feeding times,

' Agriculture and Agri-Food Canada, Pacific Agri-Food Research Centre, 4200 Hwy 97, Box 5000, Sum-

merland, British Columbia, Canada VOH 1Z0

28 J. ENTOMOL. Soc. BRIT. COLUMBIA 105, DECEMBER 2008

lasting from several seconds to a few min-

utes in duration (Raccah 1986). In uncon-

trolled cage studies, Hillman et al. (1995)

were the first to demonstrate aphid trans-

mission of BIScV. An unidentified aphid

collected from blueberry and placed on

infected Chenopodium quinoa Willd.

(Chenopodiaceae), an alternate herbaceous

host for the New Jersey strain of BIScV,

was shown to transmit the virus to unin-

fected C. quinoa. In a similar manner, Bris-

tow et al. (2000) were able to demonstrate

infection of containerized highbush blue-

berry plants in cages supplied with diseased

blueberry leaves infested with Ericaphis

fimbriata (Richards). In the same study,

transfer of individual aphids from infected

blueberry leaves to containerized potted test

plants resulted in a very low rate of infec-

tion, less than one percent. These previous

studies were not designed to determine if

BlScV was transmitted by aphids in a semi-

persistent or non-persistent manner. Two

carlaviruses vectored by aphids are thought

to be transmitted in a semi-persistent man-

ner (Bristow et al. 2000).

A better understanding of BIScV epide-

miology will aid in the development of ef-

fective control measures. To this end, the

purpose of our study was to determine the

mode of transmission of BIScV and com-

pare aphid transmission efficiencies of E.

fimbriata, a species that colonizes blue-

berry, with transmission by several non-

colonizing aphid species. Identification of

effective aphid vectors will also assist in

future laboratory investigations to deter-

mine biological differences between the

various strains of BIScV.

MATERIALS AND METHODS

Plant and aphid culture. Large

highbush blueberry plants from two com-

mercial fields near Abbotsford, British Co-

lumbia (BC), that had previously tested

positive for BIScV by ELISA using poly-

clonal antibodies (Agdia, Elkhart, Indiana)

were potted into large (~ 60 cm x 43 cm

deep) plastic pots and moved to a green-

house at the Pacific Agri-Food Research

Centre, Summerland, BC. These plants also

formed the basis for the isolation and mo-

lecular characterization of two major strains

of BIScV (Bernardy et al. 2005).

Nicotiana occidentalis Wheeler, re-

cently identified as a herbaceous host for

BIScV (Lowery et al. 2005), was grown in

the greenhouse in 20-cm plastic containers

in a 1:1:5 mixture of steam-sterilized field

soil, perlite, and commercial potting soil

(Pro-Mix BX, Premier Horticulture Ltd.,

Dorval, Quebec). Temperatures were vari-

able and ranged from daytime highs of 25 °

C to nighttime lows of 15 °C, with supple-

mental lighting supplied by sodium vapour

lamps to provide a 16-h photophase. Plants

were used at the four- or five-true-leaf

stage. Small BIScV-free blueberry plants cv

‘Berkeley’ were acquired from a commer-

cial supplier (Fall Creek Nurseries, Lowell,

Oregon) and grown in the greenhouse in

3.8-litre plastic pots under the same condi-

tions.

Aphids were maintained in _ vented,

Plexiglas ® cages (50 cm x 50 cm x 33 cm

wide) in a growth room (18 °C, 16-h photo-

phase) on suitable host plants as follows:

red and green forms of EF. fimbriata on

strawberry, Fragaria x ananassa Duchesne

(Rosaceae); spirea aphid, Aphis spiraecola

(Patch) and apple aphid, A. pomi DeGeer,

on apple, Malus domestica L. (Rosaceae);

pea aphid, Acyrthosiphon pisum (Harris),

on garden pea, Pisum sativum L.

(Fabaceae); violet aphid, Myzus ornatus

Laing, and Aphis helianthi Monell on sun-

flower, Helianthus annuus L. (Asteraceae);

green peach aphid, Myzus persicae (Sulzer),

on bok-choi, Brassica rapa _ UL.

(Brassicaceae); and the bird cherry-oat

aphid, Rhopalosiphum padi (L.), on barley,

Hordeum vulgare L. (Poaceae). Host plants

were reared in the greenhouse under condi-

tions outlined above.

Except for E. fimbriata that were origi-

nally collected from commercial fields of

highbush blueberry in the Fraser Valley and

provided by Dr. D.A. Raworth (Agriculture

and Agri-Food Canada, Pacific Agri-Food

J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

Research Centre, Agassiz, BC), all of the

aphid species used in these studies, other

than A. pisum, were collected in Summer-

land, BC, from the hosts on which they

were reared. Acyrthosiphon pisum was col-

lected from garden peas in Armstrong, BC.

Aphids were identified by Dr. R.G. Foottit

(Agriculture and Agri-Food Canada, East-

ern Cereals and Oilseeds Research Centre,

Ottawa, Ontario).

Aphid transmission studies. Fourth

instar and adult apterous aphids from the

laboratory colonies were placed in small

self-sealing petri dishes containing mois-

tened filter paper for a 2- to 3-h pre-

acquisition starvation period. Aphids were

allowed to feed for 5 min on BIScV-

infected leaf pieces in groups of 10 aphids/

petri dish, and then transferred, 25 aphids/

plant for blueberry and 10 aphids/plant for

N. occidentalis, to BIScV-free test plants,

which were then sealed in plastic bags to

prevent the aphids from escaping. Fine,

moistened natural fibre brushes were used

to transfer aphids. At least 1 h after the final

transfer, plants were sprayed with the aphi-

cide pirimicarb (Pirimor 50WP, Chipman

Chemicals Ltd., Stoney Creek, Ontario) to

kill any remaining aphids. Plants were held

in the bags for a further 24 h to ensure that

all aphids were dead. Nicotiana occiden-

talis were then moved to a growth chamber

at 20 °C under fluorescent and incandes-

cent lights (approx. 185 wmol m” s’ PAR)

and a 16-h photophase. After 6 to 8 wk,

plants were tested for BIScV infection by

enzyme-linked immunosorbent assay

(ELISA) using polyclonal antibodies

(Agdia, Elkhart, Indiana). Blueberry plants

were held in the greenhouse under the con-

ditions outlined above and tested for infec-

tion approximately 3 mo later. Plants were

then moved to cold storage at 4 °C for 3 mo

and then re-tested 2 to 3 mo after being

returned to the greenhouse. Virus transmis-

sion studies for all species of aphids and

both species of host plant were conducted

concurrently.

ELISA analysis. The double-antibody

sandwich (DAS) ELISA method used was a

modification of the protocol described by

Clark and Adams (1977). All reagents were

added at 100 wl per well in microtitre

plates. Microtitre plates (EIA Microplate,

ICN Biomedicals, Irvine, California) were

coated with purified immunoglobulin (IgG)

(Agdia, Elkhart, Indiana) diluted (5 yl mI)

in phosphate-buffered saline (PBS) for 4 h

at 37 °C. Plates were washed three times

with PBS. Plant samples (0.25 g) were thor-

oughly ground in Bioreba bags (Bioreba

AG, Reinach, Switzerland) with 1.5 ml

borate buffer (0.1 M boric acid, 0.01 M

sodium borate, 2% polyvinylpyrolidine

(PVP 44,000), 0.2% non-fat milk powder,

0.05% Tween-20, 0.5% nicotine), and the

bags briefly centrifuged at 2000 rpm to aid

pipetting. The liquid extract (25 wl) and

borate buffer (75 41) were added to the mi-

crotitre plates, which were covered in cello-

phane and placed overnight on an orbital

shaker at 600 rpm. After washing the plates

with PBS-Tween and adding a dilute (5 ul

ml') IgG-enzyme conjugate in PBS-

Tween-BSA-polyvinylpyrolidine, plates

were incubated at 37 °C for 2 h. After

plates were washed with buffer, a dilute

(0.5 mg mI’) solution of p-nitrophenyl

phosphate buffer was added. Plates were

incubated at room temperature on an orbital

shaker (600 rpm) for about 1 hr and absorb-

ance was read at 405 nm. A subset of

healthy blueberry nursery plants was tested

by ELISA to verify that they were free of

BIScV.

In order to verify BlScV infections, a

subset of blueberry and N. occidentalis

plants that had tested positive by ELISA

were also tested by reverse transcriptase

polymerase chain reaction (RT-PCR) as

described in Bernardy et al. (2005).

Statistical analysis. Differences in rates

of transmission of BIScV by the various

aphid species were determined by contin-

gency table analysis and multiple compari-

sons for proportions, analogous to a

Tukey’s test (Zar, 1984). Data were ana-

lyzed separately for each combination of

infected source and healthy test plants. In-

fection rates were not included in the analy-

sis if fewer than ten test plants had been

inoculated.

30 J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

RESULTS

By using WN. occidentalis for both

BlScV-infected and healthy test plants, we

were able to compare transmission rates for

several species of aphids not previously

known to vector this disease (Table 1). Sev-

eral species, including A. pomi, M. persi-

cae, and R. padi, were inefficient vectors

that were able to infect N. occidentalis only

at low transmission rates ranging from 2%

to 4%. The highest rate of transmission

from infected to healthy N. occidentalis

occurred when M. ornatus (average trans-

mission rate 69%) or A. helianthi (data not

shown) were used as vectors. Unfortu-

nately, the latter species was not included in

the statistical analysis due to the death of

the colony from a fungal infection before

the tests could be completed. Both the

green and red forms of E. fimbriata trans-

mitted BIScV between N. occidentalis at

intermediate rates of 10% and 8%, respec-

tively. Acyrthosiphon pisum and A. spirae-

cola did not transmit BIScV from infected

to healthy N. occidentalis.

The highest rate of infection of highbush

blueberry (20%) was recorded for the green

form of E. fimbriata, whereas infection

rates for M. ornatus and A. spiraecola were

both 7% (Table 1). Aphis helianthi was not

included in the data analysis, as we were

only able to inoculate six blueberry plants

with BlScV using this species before the

colony collapsed due to a fungal infection.

However, the infection rate for this species,

which does not colonize blueberry, ap-

peared to nearly equal that for the coloniz-

ing species E. fimbriata.

Virus transmission tests from infected

blueberry to N. occidentalis were conducted

to evaluate the acceptability of N. occiden-

talis as a trap plant in field studies of BIScV

epidemiology. No plants became infected

when M. persicae was used to vector the

virus from infected blueberry to N. occiden-

talis (Table 1), but use of the green form of

E. fimbriata resulted in an infection rate of

21%:

The utility of NV. occidentalis, a recently

identified herbaceous host of BIScV

(Lowery et al. 2005), for laboratory studies

of aphid transmission efficiencies was dem-

onstrated in this study. Even though fewer

aphids (10/plant) were used to inoculate N.

occidentalis than blueberry (25/plant), over-

all infection rates were similar. Blueberry

plants had to be held for many months to

demonstrate virus transmission, and ap-

proximately half the plants tested positive

only after an intervening 3-month cold pe-

riod. This was expected since virus titres

are generally low in blueberry compared

with herbaceous hosts, the virus is often

distributed unevenly within blueberry

plants, and detection may vary seasonally

(Martin and Bristow 1988, Wegener ef al.

2006). In comparison, unequivocal ELISA

results could be obtained for infected N.

occidentalis within 6 to 8 wk after infection

and plants then retained a high virus titre

over a period of several months.

DISCUSSION

Carlaviruses were, until recently, one of

the largest and least studied of the plant

virus groups (Foster 1992). Diseases caused

by these viruses often result in latent infec-

tions or they cause indistinct, mild symp-

toms, which resulted in carlaviruses being

largely ignored by pathologists. BIScV is an

exception to this general condition, with

infections resulting in significant loss of

yield and eventual death of certain cultivars

of highbush blueberry. For this reason, a

number of recent studies have investigated

the molecular characteristics, epidemiology,

and aphid transmission of BIScV.

Carlaviruses are transmitted largely by

aphids in a non-circulative, non-persistent

manner (Foster 1992). Certain of them are

thought to be transmitted in a _ semi-

persistent manner, however, and at least

one member of the group, Cowpea mild

mottle virus, is transmitted by whiteflies

(Harris 1983, Ng and Perry 2004). In the

J. ENTOMOL. SOc. BRIT. COLUMBIA 105, DECEMBER 2008

Table 1.

Aphid transmission of Blueberry scorch virus from infected Nicotiana occidentalis or

highbush blueberry, Vaccinium corymbosum, to healthy test plants.

Aphid Species Infected Source Test Species Infected/ % Infection

Total

Acyrthosiphon pisum Nicotiana Nicotiana 0/41 Od

occidentalis occidentalis

Aphis pomi N. occidentalis N. occidentalis 1/48 2c

Aphis spiraecola N. occidentalis N. occidentalis 0/33 Od

Ericaphis fimbriata, N. occidentalis N. occidentalis 4/40 10b

green form

Ericaphis fimbriata, N. occidentalis N. occidentalis 3/40 8bc

red form

Myzus ornatus N. occidentalis N. occidentalis 11/16 69a

Myzus persicae N. occidentalis N. occidentalis 2/48 Abc

Rhopalosiphum padi N. occidentalis N. occidentalis 1/40 3be

Acyrthosiphon pisum blueberry blueberry 0/18 Ob

Aphis pomi blueberry blueberry 0/15 Ob

Aphis spiraecola blueberry blueberry 1/14 Ta

Ericaphis fimbriata, blueberry blueberry 5/2) 20a

green form

Ericaphis fimbriata, blueberry blueberry 0/7 -

red form

Myzus ornatus blueberry blueberry 1/14 Ta

Myzus persicae blueberry blueberry 0/24 Ob

Rhopalosiphum padi blueberry blueberry 0/23 Ob

Ericaphis fimbriata, blueberry N. occidentalis 4/15 Zia

green form

Myzus persicae blueberry N. occidentalis 0/30 Ob

' For each combination of infected source and healthy plant species, infection rates followed by

the same letter are not significantly different based on contingency table analysis and multiple

comparisons for proportions (Zar 1984).

present study, the results of earlier uncon-

trolled cage studies that demonstrated trans-

mission of BIScV by E. fimbriata (Bristow

et al. 2000) were confirmed. Utilizing

timed acquisition feeding periods, we found

that BIScV is indeed transmitted by aphids

in a non-persistent manner, as might be

expected for a member of the carlavirus

group. Aphids were able to acquire the vi-

rus during brief acquisition-feeding periods

lasting less than 5 minutes. Additionally,

aphids that do not colonize blueberry, such

as M. ornatus, were equally efficient virus

vectors compared to the colonizing species

E. fimbriata. A pre-acquisition fasting pe-

riod and short virus-acquisition probes in-

crease transmission of non-persistent vi-

ruses, while prolonged feeding leads to

greatly reduced transmission rates

(Maramorosch 1963). For this reason, under

field conditions, non-colonizing aphids are

often more important vectors of viruses

such as BIScV. Due to low virus titres,

however, a slightly longer acquisition feed-

ing period might improve transmission effi-

ciencies when BlIScV is acquired from

highbush blueberry. Although EF. fimbriata

was not the most efficient vector of BIScV

from N. occidentalis to N. occidentalis, it

was the best vector when the virus was ac-

32 J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

quired from highbush blueberry, possibly

because this species was observed to settle

and feed more readily on this host plant.

Thus, E. fimbriata might contribute signifi-

cantly to the spread of this virus within in-

fected fields, particularly in years with large

numbers of these colonizing aphids.

Infection of N. occidentalis by virulifer-

ous aphids in this study occurred at a level

comparable with that for infections of her-

baceous plants with non-persistently trans-

mitted potyviruses. In similar controlled

studies, transmission of Potato virus Y from

infected to healthy sweet pepper by M. per-

sicae resulted in an 89% infection rate

(Lowery et al. 1997), whereas in another

study involving several species of aphids

the maximum rate of infection of rutabaga

with Turnip mosaic virus was 55% (Lowery

1997). The highest rate of infection of N.

occidentalis with BIScV falls within these

range of values (Table 1). Successful trans-

mission of BlScV from blueberry to N.

occidentalis by E. fimbriata has also been

used successfully to help purify and am-

plify virus in strain determination studies

(Bernardy et al. 2005). Compared with

highbush blueberry, this herbaceous host

should prove useful as a trap or sentinel

plant in studies of BIScV epidemiology. It

will be necessary, however, to first show

that NV. occidentalis is uniformly susceptible

to all strains of BIScV.

During a two year study, Raworth et al.

(2006) captured alate aphids of 87 species

in water pan traps placed in commercial

blueberry fields in the Fraser Valley, BC.

Our results suggest that many of these spe-

cies are likely vectors of BIScV. Future

virus transmission studies involving aphid

species that were captured in large numbers

from the middle of June to the middle of

July when most trap plants became infected

(Raworth et a/. 2008), which would include

several species such as Euceraphis betulae

(Koch) that develop on trees (Raworth et al.

2006), might help identify some of the

other major vectors contributing to the

spread of BIScV and suggest possible man-

agement strategies.

Based on our laboratory results, BIScV

is transmitted between highbush blueberry

at a rate similar to that for other non-

persistent, aphid-borne viruses of woody

perennial plants. In comparable transmis-

sion tests using 50 M. persicae per plant a

‘D’ strain of Plum pox virus, a member of

the Potyviridae, was transmitted from in-

fected peach, Prunus persicae L., to healthy

peach seedlings at an average infection rate

of 22% (D.T.L. unpublished data). In the

present study with blueberry, but using only

25 aphids per plant, a maximum infection

rate of 20% was recorded for E. fimbriata

(Table 1). The relatively low rate of trans-

mission from blueberry to blueberry as

compared with infection of herbaceous

hosts suggests that a number of years would

be required for BIScV to spread throughout

a blueberry field from an initial infection

locus. Accordingly, mapping of disease

incidence in three commercial blueberry

fields in the Fraser Valley, BC, showed that

BlScV spread only slowly (Wegener ef al.

2006). Similarly, spread of BIScV through-

out two commercial fields of blueberry in

the northwestern United States required

between 5 to 8 years (Bristow ef al., 2000),

and Raworth ef al. (2008) recorded a low

rate of BIScV infection for highbush blue-

berry and N. occidentalis bait plants placed

weekly throughout the summer in highly

infected commercial blueberry fields, indi-

cating a low rate of natural spread. Given

the relatively slow spread of the virus under

field conditions, these findings suggest that

planting only certified virus-free nursery

material and aggressive removal of diseased

plants might provide an effective means of

control of BIScV under field conditions.

ACKNOWLEDGEMENTS

This work was funded by the BC Blue-

berry Council and Agriculture and Agri-

Food Canada (AAFC) through the Match-

ing Investment Initiative. We wish to thank

Lisa Wegener and Mark Sweeney, BC Min-

istry of Agriculture and Lands, for supply-

J. ENTOMOL. SOc. BRIT. COLUMBIA 105, DECEMBER 2008 33

ing the original infected blueberry plants, _ for identifying the aphid species. Additional

and Dr. R.G. Foottit, AAFC Eastern Cere- technical assistance was provided by Mi-

als and Oilseeds Research Centre, Ottawa, — chael Bouthillier and Sonya Goulet.

REFERENCES

Bernardy, M.G., C.R. Dubeau, A. Braun, C.E. Harlton, A. Bunckle, L.A. Wegener, D.T. Lowery and C.J.

French. 2005. Molecular characterization and phylogenetic analysis of two distinct strains of blueberry

scorch virus from western Canada. Canadian Journal of Plant Pathology 27: 581-591.

Bristow, P.R., R.R. Martin and G.E. Windom. 2000. Transmission, field spread, cultivar response, and

impact on yield in highbush blueberry infected with blueberry scorch virus. Phytopathology 90: 474-479.

Caruso, F.L. and D.C. Ramsdell. 1995. Compendium of Blueberry and Cranberry Diseases. APS Press, St.

Paul, MN.

Catlin, N.J. and S.G. Schloemann. 2004. Blueberry Scorch Virus (BIScV). UMass Amherst Extension Fact-

sheet, F-129-2004, University of Massachusetts, Amherst, MA.

Cavileer, T.D., B.T. Halpern, D.M. Lawrence, E.V. Podleckis, R.R. Martin and B.I. Hillman. 1994. Nucleo-

tide sequence of the carlavirus associated with blueberry scorch and similar diseases. Journal of General

Virology 75: 711-729:

Ciuffo, M., D. Pettiti, S. Gallo, V. Masenga and M. Turina. 2005. First report of blueberry scorch virus in

Europe. Plant Pathology 54: 565.

Clark, M.F. and A.N. Adams. 1977. Characteristics of the microplate method of enzyme-linked immunosor-

bent assay for the detection of plant viruses. Journal of General Virology 34: 475-483.

Foster, G.D. 1992. The structure and expression of the genome of carlaviruses. Research Virology 143:

103-112.

Harris, K.F. 1983. Sternorrhynchous vectors of plant viruses: virus-vector interactions and transmission

mechanisms. Advances in Virus Research 28: 113-140.

Hillman, B.I., D.M. Lawrence and B.T. Halpern. 1995. Characterization and detection of blueberry scorch

carlavirus and red ringspot virus caulimovirus. Journal of Small Fruit and Viticulture 3: 83-93.

Lowery, D.T. 1997. Turnip Mosaic Virus Monitoring and Research Program Final Report. Ontario Ministry

of Agriculture, Food, and Rural Affairs, Guelph, ON.

Lowery, D.T., K.C. Eastwell and M.J. Smirle. 1997. Neem seed oil inhibits transmission of Potato virus Y

to pepper. Annals of Applied Virology 130: 217-225.

Lowery D.T., C.J. French and M.G. Bernardy. 2005. Nicotiana occidentalis: a new herbaceous host for

blueberry scorch virus. Plant Disease 89: 205.

Maramorosch, K. 1963. Arthropod transmission of plant viruses. Annual Review of Entomology 8: 369-

414,

Martin, R.R. and P.R. Bristow. 1988. A carlavirus associated with blueberry scorch disease. Phytopathology

78: 1636-1640.

Ng, C.K. and K.L. Perry. 2004. Transmission of plant viruses by aphid vectors. Molecular Plant Pathology

5: 505-511.

Podleckis, E.V. and R.F. Davis. 1989. Infection of highbush blueberries with a putative carlavirus. Acta

Horticulturae 241: 338-343.

Raccah, B. 1986. Nonpersistent viruses: epidemiology and control. Advances in Virus Research. 31: 387-

429.

Raworth, D.A., C.J. French, D.T. Lowery, M.G. Bernardy, M. Bouthillier, S. Mathur, C.-K. Chan, R.G.

Foottit, E. Maw, L.A. Wegener and M. Sweeney. 2008. Temporal trends in the transmission of Blueberry

scorch virus in British Columbia, Canada. Canadian Journal of Plant Pathology 30: 345-350.

Raworth, D.A., C.-K. Chan, R.G. Foottit and E. Maw. 2006. Spatial and temporal distribution of winged

aphids (Hemiptera: Aphididae) frequenting blueberry fields in southwestern British Columbia and impli-

cations for the spread of Blueberry scorch virus. The Canadian Entomologist 138: 104-113.

Wegener, L.A., R.R. Martin, M.G. Bernardy, L. MacDonald and Z.K. Punja. 2006. Epidemiology and iden-

tification of strains of blueberry scorch virus on highbush blueberry in British Columbia, Canada. Cana-

dian Journal of Plant Pathology 28: 250-262.

Zar, J.H. 1984. Biostatistical Analysis. Prentice-Hall, Englewood Cliffs, NJ.

J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

Sex attraction in Polistes dominulus (Christ) demonstrated

using olfactometers and morphological source extracts

JOHN K. MACKENZIE’, PETER J. LANDOLT™

and RICHARD S. ZACK°

ABSTRACT

Y-tube and parallel tube olfactometers were used to test for attraction between and

within genders of the European paper wasp, Polistes dominulus (Christ). In the Y-tube

olfactometer, unmated females were attracted to male odour, while males were repelled

by unmated female odour. Males and females were not attracted to the odour of the

same sex in this experiment. In the parallel tube olfactometer, females were attracted to

male odour, while males were not attracted to female odour. Morphological sources of

potential sex attractants were tested using an arena bioassay design. Males rubbed their

mandibles and gaster on the substrate when exposed to extracts of unmated female or

male tagmata, female or male legs, or the male seventh gastral sternite. We did not see

overt behavioural responses by females to male or female extracts.

Key Words: attractant, pheromone, Polistes dominulus, paper wasp

INTRODUCTION

Sex pheromones are chemicals that

elicit behaviour related to mate-finding,

mate-selection, and copulation in insects,

including vespid wasps (Wilson 1971;

Shorey 1977; Landolt et al. 1998). Close-

range attractants and copulatory incitants or

aphrodisiacs have been demonstrated be-

tween males and females of the social

wasps Polistes exclamans Viereck (Post

and Jeanne 1984; Reed and Landolt 1990a),

Polistes fuscatus (F.) (Post and Jeanne

1983a, 1984), Belonogaster petiolata De-

geer (Keeping et al. 1986), Vespula

squamosa Drury (Reed and Landolt 1990b),

and Vespa spp. (Batra 1980; Ono and Sa-

saki 1987). Despite such demonstrations, no

vespid sex pheromone chemical structure

has been identified. Sexual behaviour of

Polistes dominulus (Christ) (Hymenoptera:

Vespidae) has not been described or quanti-

fied in controlled experiments. Knowledge

of behavioural responses to putative phero-

mones is necessary for accurate pheromone

characterization.

Behavioural evidence in Polistes paper

wasps suggests that sex pheromones from

exocrine glands in the mandibles, legs, and

gastral sterna may be involved in mate at-

traction (Landolt and Akre 1979; Jeanne ef

al. 1983; Beani and Turillazzi 1988; Beani

and Calloni 1991a,b; Beani et a/. 1992). In

several of these species, mating often oc-

curs away from the nest (Noonan 1978) on

perching substrates that are at prominent

locations such as on hilltops (Bean and

Turillazzi 1988; Mathes-Sears and Alcock

1986). Males, in some species of Polistes,

scent-mark by dragging their posterior gas-

tral sternites (Post and Jeanne 1983b; Reed

and Landolt 1991) and by rubbing their

mandibles (Wenzel 1987; Reed and Landolt

1991) on the perching substrate. Four spe-

' 4316 Pioneer Court, Abbotsville, BC, Canada V2S 7Z1

: USDA-ARS, 5230 Konnowac Pass Road, Wapato WA 98951, USA

* Department of Entomology, Washington State University, Pullman WA 99164, USA

i Correspondence: USDA, ARS, 5230 Konnowac Pass Road, Wapato, WA 98951 USA, Tel. 509 454

6570, fax 509 454 5646, e-mail peter.landolt@ars.usda.gov

36 J. ENTOMOL. SOc. BRIT. COLUMBIA 105, DECEMBER 2008

cies, including P. dominulus, are shown to

drag their hind legs on the substrate (Beani

and Calloni 1991a). Polistes dominulus

males have ducted class III gland cells that

open onto the cuticle in their legs, as well

as onto the seventh abdominal sternite

(Downing et al. 1985; Beani and Calloni

1991a). Although females of Polistes spp.

are not known to show overt scent-marking

behaviour, P. exclamans females possess a

pheromone in the venom that elicits sexual

behaviour in both conspecific and_het-

erospecific males (Post and Jeanne 1984).

Additionally, a surface pheromone on the

thoracic and gastral cuticle in P. fuscatus 1s

important for male recognition of conspeci-

fic females (Post and Jeanne 1984).

The objective of this study was to inves-

tigate orientation and behavioural responses

of P. dominulus to potential sex odours

from a variety of morphological sources.

Klinotaxic (turning orientation) and ortho-

taxic (forward orientation) (Fraenkel and

Gunn 1940; Wyatt 2003) responses to male

odour and unmated female odour were

tested using Y-tube and parallel tube olfac-

tometers, respectively. An arena-type bioas-

say was used to test for behavioural re-

sponses to extracts of male and unmated

female tagmata and glands. These studies

are foundational in aiding the overarching

objectives of determining sex pheromone

signaling systems in this species, and deter-

mining sources of those sex pheromones.

MATERIALS AND METHODS

Colony Collection and Maintenance.

Polistes dominulus nests with pupae were

collected in the field, and placed in plastic

screened cages (30.5 x 30.5 x 30.5 cm) dur-

ing late August and early September in

2003 and 2004. Nests were collected at this

time because males were abundant, indicat-

ing that reproductive females would be

emerging from nests, and not worker fe-

males. Collected nests were monitored

daily to remove and segregate male and

female adults that emerged, in order to

minimize encounters between the sexes and

exposure to sex pheromones. Male and fe-

male wasps obtained in this manner were

assumed to be sexually inexperienced since

mating is reported to occur away from the

nest (Noonan 1978). These wasps were

used for the preparation of extracts and for

behavioral assays conducted in Pullman and

Yakima, Washington.

In Pullman (Whitman County), Wash-

ington, USA, the newly emerged wasps

were kept in a laboratory at 24°C, 40% RH,

under a natural light regime (14 hours of

light and 10 hours of dark), until testing in

the Y-tube olfactometer and in the parallel

tube olfactometer. All Y-tube olfactometer

tests were conducted in Pullman, while one

half of bioassay replicates for each experi-

ment conducted with parallel tube olfacto-

meters were conducted in Pullman and the

other half at the USDA, ARS Yakima Agri-

cultural Research Laboratory near Yakima

(Yakima County), Washington, USA.

Wasps used in olfactometer tests and the

arena bioassays conducted in Yakima were

kept in a glass greenhouse under natural

lighting at 30 + 3 °C and 35% RH. At both

sites, wasps in cages were provided water

and a 1:10 molasses:water solution on cot-

ton balls for nutrition. Water and the solu-

tion of molasses were refreshed or replaced

daily. Wasps used in all assays were be-

tween 2 and 14 days old; they were ran-

domly selected for each trial and were not

reused in other trials for at least 48 hours.

Y-tube Olfactometer Bioassay. Un-

mated male and female wasps were tested

for klinotaxic responses to unmated female

odour and male odour in the Y-tube olfacto-

meter. The inside diameter of the glass Y-

tube was 2.5 cm and the length of the tube,

from stem base to Y-juncture, was 18 cm.

Airflow of 100 ml/min was measured with

a flowmeter (Aalborg Instruments, Monsey,

NY) before and after passing through the

480 ml jars housing the treatments. Air

passed through the treatment jar containing

3 “bait wasps” of the sex being assayed,

then through one arm of the Y-tube, and out

the stem of the Y-tube. Simultaneously, air

J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

passed through the empty control jar,

through the other arm of the Y-tube, and

out the stem of the Y-tube.

A paper wasp was placed in the stem of

the Y-tube and observed for a maximum of

five minutes. If the wasp moved upwind to

the Y-juncture and then moved completely

beyond the juncture into either of the arms

(with treatment airflow or with control air-

flow), that assay was ended and the re-

sponse was recorded. Ten wasps were

tested individually and in succession using

the same “bait wasps” in the treatment jar.

To eliminate a potential left or right turning

bias, the positioning of the treatment and

control was switched after the first 5 wasps

had been tested. A clean olfactometer sys-

tem was then set up and a fresh set of bait

wasps was placed in the treatment jar. This

experimental protocol was conducted four

times to provide a total of 40 wasps (5

wasps in series x 2 treatment positions x 4

= 40) tested for responses. Wasps entering

the treatment arm, the control arm, or nei-

ther arm of the Y-tube were recorded. For

each experiment, the numbers of wasps that

entered the treatment arm or the control arm

were compared using the Chi-square good-

ness-of-fit test with Yates correction for

continuity at P < 0.05 (Zar, 1974).

The olfactometer system was placed

horizontally 50 cm beneath two 1.2 m long,

34W fluorescent bulbs (Osram Sylvania

Corp., Danvers, MA) and one 160W mer-

cury vapour bulb (Osram Sylvania Corp.,

Danvers, MA). Temperature at the olfacto-

meter surface was 31°C. Air moving

through the Y-tube olfactometer was sup-

plied by an aquarium air pump, purified

through a hydrocarbon trap (Alltech Asso-

ciates Inc., Deerfield, IL), and humidified

with a gas diffusion bottle. All glassware

(Ace Glass, Inc. Vineland, NJ) and steel

tubing were washed in hot water with Mi-

cro-90 cleaning solution (International

Products Corp., Burlington, NJ), and then

rinsed serially with deionized water, ace-

tone, and then hexane. Glassware was sub-

sequently placed in a drying oven at 180°C

overnight before used again in assays.

Parallel Tube Olfactometer Bioassay.

Unmated females were tested for orthotaxic

responses to male odour and males were

tested for orthotaxic responses to unmated

female odour in a parallel or “straight tube”

olfactometer design. This design is based

on that of Tobin et a/ (1981) and was re-

ported by Landolt et al. (1988). The olfac-

tometer set up was the same as the Y-tube

set up, except for the replacement of the Y-

tube with two straight glass tubes. Each

straight tube was supplied 100 ml/min of

metered, purified humidified airflow that

was passed through a glass jar housing an

odour source, separate from the other tube

and odour source. This setup was placed on

a laboratory table with fluorescent lighting

above and natural lighting from windows.

A wasp was placed in a straight glass tube,

2.5 cm diameter and 18 cm long, downwind

from the treatment airflow (3 bait wasps)

and another wasp was placed in an identical

tube downwind from the control airflow

(empty). For each wasp, the time it took to

cover the full 18 cm distance of the tube

was recorded, if indeed it completed the full

distance upwind. This assay was conducted

with 10 pairs of wasps (treatment and con-

trols paired), and the glassware for treat-

ment and control were switched after 5

pairs of wasps were tested. This experiment

was then replicated eight times, (N = 80)

and treatment mean times were separated

from control mean times using a paired (-

test at P < 0.05. Also, the mean percents of

those that travelled the entire lengths of the

treatment and control tubes, within the five

minute time limit, were separated using a

paired f-test at P< 0.05.

One half of the parallel tube olfactome-

ter replicates were conducted under the

same conditions as the Y-tube olfactometer

bioassays, in Pullman. The other replicates,

in Yakima, were conducted as they were in

Pullman, with these slight modifications:

(1) the bioassays were conducted in a con-

trolled environment room at 24 °C and 65%

RH; (2) airflow was from a compressed air

source; (3) light was supplied by two, 1.2 m

long, 34 W fluorescent light bulbs (Osram

Sylvania Corp., Danvers, MA) 50 cm above

the olfactometer. A J16 Digital Photometer

38 J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

(Tektronix Inc., Beaverton, OR) measured

at 27,663 lux (lumens/m’) at the olfactome-

ter surface; (4) after cleaning, the olfacto-

meter glassware and tubing were placed in

a drying oven for 24 h. Data from the paral-

lel tube olfactometer bioassays in Pullman

and Yakima were pooled and analyzed to-

gether. We did not expect the minor differ-

ences in assay conditions to alter the behav-

iour of the wasps and a preliminary analysis

of the results indicated similar responses in

the assays.

Tagmata and Extract Preparations.

Dissecting and grinding tools and equip-

ment were washed in hot water with Micro-

90 cleaning solution, and then rinsed with

deionized water, acetone, and methylene

chloride. Samples of 40 female heads, 40

male heads, 40 female thoraces, 40 male

thoraces, 40 female gasters, and 40 male

gasters, all from freshly freeze-killed

wasps, were each ground with a mortar and

pestle in methylene chloride. Additionally,

40 female venom sacs with acid sting

glands, 40 female alkaline glands, legs of

40 females, 40 male mandibles with ectal

glands, 40 male seventh gastral sternites

with glands, and legs of 40 males were dis-

sected or removed and then extracted with

methylene chloride. All tagmata and gland

extracts were reduced to 4 ml under a N>

stream and kept in a freezer at -15 °C, pro-

viding concentrations of one wasp-

equivalent per 100 uL of extract.

Tagmata and Gland Bioassay. Arena

bioassays were conducted in the same

greenhouse environment in which the

wasps were housed. The assays occurred

over the course of 3 weeks in September

between 10:00 and 16:00 hr. Light intensity

at the table was 16,758 + 795 (mean + S.E.)

lux, measured at 20 different times through-

out the bioassays.

On a table covered with white paper, a

wasp was placed under the bottom half of

an upside down, plastic, 8.5 cm diam. Petri

dish for one minute before experiencing

extract odour. Petri dishes and paper were

discarded after each bioassay. Immediately

prior to conducting the assay, 100 ul of the

treatment extract or 100 ul of the methylene

chloride control were applied to 4 wedges

of 5.5 cm diameter, #3 Whatman Filter Pa-

per (Whatman International Ltd., Maid-

stone, England). The methylene chloride

was evaporated before the filter paper was

placed under the Petri dish with the wasp.

Wasps were observed for two minutes

while in the presence of the extract or sol-

vent blank, after which they were placed

into holding cages to ensure they were not

used again in the assay. At the end of the

assay period on any given day, all wasps

were returned to cages that constituted the

general pools of male and female wasps

from which random selections were made

for subsequent experiments.

Male and female wasps were tested for

responses to extracts of tagmata and glands

from both sexes. Each of the tests was repli-

cated 20 times. In bioassays of male and

female tagmata, the sequence was: blank,

head, thorax, and gaster. In bioassays of

female gland bioassay the sequence was:

blank, venom, legs, and alkaline gland. In

bioassays of male glands the sequence was:

blank, mandibles, legs, and seventh sternal

gland. For each assay, a record was kept of

continuous movement, no movement, and

the number of times a wasp showed stop &

go movement, antennal contact with the

filter paper, grooming fore legs through

mandibles and then rubbing antennae,

grooming fore legs through mandibles then

rubbing thorax, grooming gaster with hind

legs, rubbing hind legs together, rubbing

mandibles on substrate, and rubbing gaster

on substrate. For each behaviour and each

sex, a 2x2 contingency table was con-

structed to make comparisons of the num-

ber of times a behaviour was observed for

the control versus each of the tagmata and

gland extracts. For each behavior, contin-

gency tables were analyzed using the Chi-

square statistic with Yates correction for

continuity at P< 0.05 (Zar, 1974).

J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

RESULTS

Y-tube Olfactometer Bioassays (Table

1). Females significantly more often turned

toward male odour and away from the con-

trol (P < 0.001). Males significantly more

often turned away from female odour and

toward the control (P < 0.05). Neither

males nor unmated females turned toward

odour from males (P > 0.10) and females

(P > 0.5), respectively, compared to the

control.

Parallel Tube Olfactometer Bioassays.

The mean time (4+ S.E.) of female move-

ment toward male odour (33.7 + 4.4 sec, df

= 79, P = 0.002) was significantly lower

than toward the control (49.1 + 4.1 sec).

There was no significant difference in the

mean percent (+ S.E.) of females that trav-

eled the length of the treatment (89.0 +

4.3%, df = 7, P = 0.087) and control (81.0 +

4.3%) tubes within five minutes. The mean

time (+ S.E.) for male movement toward

female odour (88.2 + 14.3 sec, df = 79, P=

0.141) was not statistically different com-

pared to the control (68.1 + 12.0 sec). This

was also true for the mean percent (+ S.E.)

of males that travelled the length of the

treatment (70.0 + 7.3%, df = 7, P = 0.361)

and control (78.8 + 6.9%) tubes within five

minutes.

Tagmata Experiments. Females did

not behave differently in the presence of

extracts of female or male tagmata com-

pared to the blank. Males rubbed their man-

dibles on the substrate more often when

treated with extract of 2 thorax (P < 0.05)

compared to the blank (Table 2). They also

rubbed their gaster on the substrate more

often when treated with extracts of 9 tho-

rax (P < 0.005) or 2 gaster (P < 0.05) com-

pared to the blank (Table 2). When pre-

sented with extracts of 4 thorax and ¢

gaster, males rubbed their mandibles on the

substrate more often (P < 0.05) compared to

the blank (Table 2).

Gland Experiments. Unmated females

did not show any significant behavioural

differences in the presence of extracts of

female or male glands compared to the

blank. Males rubbed their mandibles (P <

0.05) and gaster (P < 0.005) on the sub-

strate more often when exposed to 9 leg

extract (Table 3). When treated with 3 leg

extract, males continuously moved (P <

0.001) and rubbed their mandibles on the

substrate (P < 0.05). They also rubbed their

mandibles (P < 0.005) and gaster (P <

0.005) on the substrate when exposed to ¢

seventh sternite with gland extract (Table

3). Lastly, males rubbed their gaster on the

substrate (P < 0.005) more often when ex-

posed to @ mandibles with ectal mandibular

gland extract (Table 3).

DISCUSSION

Polistes males are known to use mating

strategies such as marking behaviour by

rubbing their mandibles, gaster, and legs on

perch sites (Beani and Calloni 1991a; Polak

1993). It is hypothesized that such scent-

marked perch sites attract females (Landolt

and Akre 1979; Post and Jeanne 1983b,c;

Wenzel 1987; Reed and Landolt 1990a).

Our orientation bioassay results using olfac-

tometers support the hypothesis that fe-

males are attracted to a pheromone of males

since male odour elicited positive turning

and increased speed of forward movement

from females. Female attraction to males in

olfactometers has also been shown for the

paper wasp species P. fuscatus (Post and

Jeanne 1983a) and P. exclamans (Reed and

Landolt 1990a).

The observed repellency of females to

males in the Y-tube, but not the parallel-

tube olfactometer, assays is puzzling and

calls for possible explanation. It is assumed

that females would control release of any

sex attractant or courtship pheromone and

we have no means of knowing if and when

females were “calling” when they were

tested in olfactometers. Thus, females may

be attractive to males under circumstances

not met by our assay conditions. Also, fe-

male venom may possess alarm pheromone

(Bruschini et al. 2006), which could com-

plicate assay results when females have

40 J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

Table 1.

Numbers of European paper wasps, Polistes dominulus, turning towards airflow from over

conspecific wasps compared to control airflow, in a Y-tube olfactometer'.

Bioassay Ntreatment Neontrol X experimental P-value

? to Y ay 18 0.225 P>0.5

Oto d 31 9 11.025 P<0.001

3 to 2 13 27 4.225 P<0.05

3 to d 16 24 e225 P>0.10

' Analyzed using Chi-square goodness-of-fit test with Yates correction for continuity where

1 theoretical(1, 0.05) = 3.841.

Table 2.

Numbers of unmated male European paper wasps, Polistes dominulus, responding to unmated

male and female tagmata in an arena type assay’.

Behaviour. Blank Head Thorax QGaster GHead CGThorax GGaster

A 12 14 ile) IZ 1] 11 14

B 3 4 0 2 4 2 2

Cc a 2 5 l 5 7 4

D 5 8 10 10 4 4 10

E 12 8 13 8 7 10 7

F 5 0 i, 3 a 6 8

G 2 4 4 3 3 3 3

H 4 0 ) | 0 4

I 2 7 Q* 7 4 Q* on

J 0 2 9 6* 2 5 4

' Within a row, treatments compared only to blank in a 2x2 contingency table analyzed using

the Chi-square statistic with Yates correction for continuity are significant at P < 0.05. Num-

bers with an asterisk are significantly different from the blank.

? A — continuous movement, B — no movement, C — stop & go, D — antennate paper, E— groom

fore legs in mandibles; rub antenna, F — groom fore legs in mandibles; rub thorax, G — groom

gaster with hind legs, H — rub hind legs together, I — rub mandibles on substrate, J — rub gaster

on substrate.

been handled and disturbed. Caution must

then be exercised in interpreting negative or

apparently conflicting results.

Female P. dominulus in this study

showed neither a positive nor negative ori-

entation response toward female odour.

Overwintering females in search of hiberna-

tion sites might be expected to respond to

aggregations of females (Reed and Landolt

1991). Heterospecific aggregations of paper

wasp gynes in hibernacula have been re-

ported (Rau 1930, 1938; Bohart 1942;

Hermann et al. 1974; Gibo 1980; Reed and

Landolt 1991) as well as purely conspecific

aggregations (Rau 1938; Strassmann 1979).

Polistes dominulus queens appear to over-

winter in multi-colony groups (Starks

2003). However, there may be multiple

factors (such as temperature and day

length) that stimulate females to seek out

hibernacula and other overwintering fe-

males. The wasps used in these bioassays

were housed under summer-like tempera-

tures and day length. Hence, we can make

no conclusions regarding the presence or

absence of a female aggregation phero-

mone.

Venom is thought to include a sex

pheromone that elicits copulatory behaviour

in males of P. fuscatus and P. exclamans

J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

Table 3.

Numbers of unmated male European paper wasps, Polistes dominulus, responding to unmated

male and female glands in an arena assay .

Behaviour’ Blank QVenom QLegs SMandibles GLegs CGSternite Blank § Alkaline

A 8 13 14 15

B 3 Z

C 9 6 + +

D 6 1] Ld 13

E 9 8 15 14

F 0 3 2,

G 6 2 10 7

H 4 2 7 6

I 2 6 o* 6

J | 4 10* o*

Loe IS 1] 12

0 2 2 5

3 7 3

12 13 S 7

2) 9 1] 2

2 4 Z

6 6 7 5

4 5 4 6

yd ie 4 2

5 10* 3 5

' Within a row, treatments compared only to blank in a 2x2 contingency table analyzed using

the Chi-square statistic with Yates correction for continuity (significant at P < 0.05, df = 1).

Numbers of responses with an asterisk are significantly different from the blank.

? A — continuous movement, B — no movement, C — stop & go, D — antennate paper, E — groom

fore legs in mandibles; rub antenna, — groom fore legs in mandibles; rub thorax, G — groom

gaster with hind legs, H — rub hind legs together, I — rub mandibles on substrate, J — rub gaster

on substrate

(Post and Jeanne 1983a; 1984). Odorants on

the female cuticle are thought also to be a

conspecific sex pheromone in these species

(Post and Jeanne 1983a, 1984). Our olfacto-

meter bioassays did not evaluate copulatory

behaviour, but rather orientation behaviour.

Males of P. dominulus did not orient by

turning or increasing their speed of move-

ment toward female odour in either the Y-

tube or parallel tube bioassays, respectively.

We did observe behavioural responses such

as increased movement and possible scent

marking by males to extracts of female

gasters and venom glands in an arena assay,

but again we did not study copulatory re-

sponses. Although venom seems to play a

role in mediating courtship or copulation in

some paper wasps, it is not known to serve

as a sex attractant. In our experiments, we

cannot rule out possible female release of

alarm chemicals due to handling stress,

complicating the interpretation of behav-

ioural assay results, although Freisling

(1943) was unable to demonstrate an alarm

pheromone in P. dominulus.

Polistes dominulus females have ducted

type III gland cells in their legs (Beani and

Calloni 1991a). Polistes fuscatus males

responded to female tagmata extracts in a

wind tunnel, but the greatest response was

to female thorax extract (Reed and Landolt

1990a). During our behaviour bioassays,

males responded to extract of 2 legs in the

same manner as to extract of the entire tho-

rax, by rubbing their mandibles and gaster

on the substrate. Additionally, males rubbed

their gaster on the substrate when exposed

to 2 gaster extract. These behavioural re-

sponses have been observed in males of

other Polistes species and described as

scent-marking (Post and Jeanne 1983b;

Wenzel 1987). Female odour may not elicit

orientation responses, but female extracts of

thoraces and legs do appear to elicit scent-

marking in males.

Scent-marking behaviour in males is

believed to attract females for mating pur-

poses, and to signal territorial ownership to

other males (Post and Jeanne 1983b; Beani

and Calloni 1991a). Male odour may deter

other males of the same species since terri-

torial males will actively chase intruders

away and, afterwards, increase grooming

and scent-marking behaviour (Post and

42 J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

Jeanne 1983b; Beani and Calloni 1991a).

Males of other Polistes species are known

to patrol and defend territorial perches near

hibernacula and nest sites by gaster drag-

ging and mandible rubbing on the substrate

(Post and Jeanne 1983b; Wenzel 1987;

Reed and Landolt 1991). The dragging of

the male gaster has also been in observed in

Mischocyttarus spp. (Litte 1979, 1981).

Nineteen of 20 males that were presented

with 3 leg extract continuously moved for

the two-minute assay; they also rubbed

their mandibles on the substrate more often

than when presented with the blank. Males

rubbed their mandibles on the substrate

when presented with CG gaster extract.

When the seventh sternal gland was dis-

sected and presented to males, they per-

formed gaster dragging in addition to rub-

bing their mandibles on the substrate. Al-

though males did not respond to 3 head

extract, they did drag their gaster on the

substrate when presented with 4 mandible

extract. Therefore, we quantified and report

behaviours that are consistent with previous

behavioural studies of different Polistes

species.

The orientation and behavioural evi-

dence reported herein supports the previ-

ously stated hypothesis that males use

scent-marking for mating and territorial

defense purposes by depositing pheromone

from well-developed exocrine glands in the

gaster, mandibles, and legs onto the perch

substrate (Landolt and Akre 1979; Jeanne et

al. 1983; Beani and Calloni 1991b). The

quantification of orientation and behav-

1oural responses to potential sex attractant

odours between and within genders is foun-

dational to the larger work of isolating,

identifying, and testing sex pheromones in

Vespidae.

ACKNOWLEDGEMENTS

Daryl Green (USDA-ARS) provided

technical assistance. The Washington State

Commission on Pesticide Registration and

Sterling International Inc. provided funding

for this work.

REFERENCES

Batra, S. W. 1980. Sexual behaviour and pheromones of the European hornet, Vespa crabro germana

(Hymenoptera: Vespidae). Journal of the Kansas Entomological Society 53:461-469.

Beani, L., and C. Calloni. 1991a. Leg tegumental glands and male rubbing behaviour at leks in Polistes

dominulus (Hymenoptera: Vespidae). Journal of Insect Behaviour 4:449-462.

Beani, L., and C. Calloni. 1991b. Male rubbing behaviour and the hypothesis of pheromonal release in

polistine wasps (Hymenoptera: Vespidae). Ethology Ecology and Evolution (Special Issue) 1:51-54.

Beani, L., and S. Turillazzi. 1988. Alternative mating tactics in males of Polistes dominulus (Hymenoptera:

Vespidae). Behavioural Ecology and Sociobiology 22:257-264.

Beani, L., R. Cervo, M. C. Lorenzi, and S. Turillazzi. 1992. Landmark-based mating systems in four

Polistes species (Hymenoptera: Vespidae). Journal of the Kansas Entomological Society 65:211-217.

Bohart, G. E. 1942. Notes on some feeding and hibernation habits of California Polistes. Pan-Pacific Ento-

mologist 18:30.

Bruschini, C., R. Cervo, and S. Turillazzi. 2006. Evidence of alarm pheromones in the venom of Polistes

dominulus workers (Hymenoptera: Vespidae). Physiological Entomology 31: 286-293.

Downing, H. A., D. C. Post, and R. L. Jeanne. 1985. Morphology of sternal glands in male polistine wasps

(Hymenoptera: Vespidae). Insectes Sociaux 32:186-198.

Fraenkel, G. S., and D. L. Gunn. 1940. The Orientation of Animals: Kineses, Taxes and Compass Reac-

tions. Oxford University Press, Oxford, United Kingdom.

Freisling, J. 1943. Zur Psychologie der Feldwespe. Z. Tierpsychol. 5:439-463.

Gibo, D. L. 1980. Overwintering of Polistes fuscatus in Canada: use of abandoned nests of Dolichovespula

arenaria. Journal of the New York Entomological Society 88:146-150.

Hermann, H. R., D. Gerling, and T. Dirks. 1974. Hibernation and mating activity of five polistine wasp

species. Journal of the Georgia Entomological Society 9:203-204.

Jeanne, R. L. 1996. The evolution of exocrine gland function in wasps, pp. 144-159, in S. Turillazzi and M.

J. West-Eberhard (eds.). Natural History and Evolution of Paper-Wasps. Oxford University Press, Ox-

J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008 43

ford, United Kingdom.

Jeanne, R. L., H. A. Downing, and D. C. Post. 1983. Morphology and function of sternal glands in polistine

wasps (Hymenoptera: Vespidae). Zoomorphology. 103:149-164.

Keeping, M. G., D. Lipschitz, and R. M. Crewe. 1986. Chemical mate recognition and release of male sex-

ual behaviour in polybiine wasp, Belonogaster petiolata Degeer (Hymenoptera: Vespidae). Journal of

Chemical Ecology 12:773-779.

Landolt, P. J., and R. D. Akre. 1979. Occurrence and location of exocrine glands in some social Vespidae

(Hymenoptera). Annals of the Entomological Society of America 33:45-57.

Landolt, P. J., R. W. Hofstetter, and P. S. Chapman. 1998. Neonate codling moth larvae (Lepidoptera: Tor-

tricidae) orient anemotactically to odor of immature apple fruit. Pan Pacific Entomologist 74: 140-149.

Landolt, P. J., R. L. Jeanne, and H. C. Reed. 1998. Chemical communication in social wasps, pp. 216-235,

in R.K. Vander Meer, M.D. Breed, M.L. Winston, and K.E. Espelie (eds.). Chemical Communication in

Social Insects. Westview Press, Boulder, Colorado.

Litte, M. 1979. Mischocyttarus flavitarsis in Arizona: social and nesting biology of a polistine wasp.

Zeitschrift fur Tierpsychologie 50:282-312.

Litte, M. 1981. Social biology of the polistine wasp Mischocyttarus labiatus: survival in a Columbian rain

forest. Smithsonian Contributions in Zoology No. 327.

Mathes-Sears, W., and J. Alcock. 1986. Hilltopping behaviour of Polistes commanchus navajoe

(Hymenoptera: Vespidae). Ethology 71: 42-53.

Noonan, K. M. 1978. Sex ratio of parental investment in colonies of the social wasp Polistes fuscatus. Sci-

ence 199:1354-1356.

Ono, M., and M. Sasaki. 1987. Sex pheromones and their cross-activities in six Japanese sympatric species

of the genus Vespa. Insectes Sociaux 34:252-260.

Polak, M. 1993. Competition for landmark territories among male Polistes canadensis (L.) (Hymenoptera:

Vespidae): large-size advantage and alternative mate-acquisition tactics. Behavioural Ecology 4:325-331.

Post, D. C., and R. L. Jeanne. 1983a. Venom: Source of a sex pheromone in the social wasp Polistes fusca-

tus (Hymenoptera: Vespidae). Journal of Chemical Ecology 9:259-266.

Post, D. C., and R. L. Jeanne. 1983b. Male reproductive behaviour of the social wasp Polistes fuscatus

(Hymenoptera: Vespidae). Zeitschrift fur Tierpsychologie 62:157-171.

Post, D. C., and R. L. Jeanne. 1983c. Sternal glands in males of six species of Polistes (Fuscopolistes)

(Hymenoptera: Vespidae). Journal of the Kansas Entomological Society 56:32-39.

Post, D. C., and R. L. Jeanne. 1984. Venom as an interspecific sex pheromone and species recognition by a

cuticular pheromone in paper wasps (Polistes Hymenoptera: Vespidae). Physiological Entomology 9:65-

7d.

Rau, P. 1930. The behaviour of hibernating Polistes wasps. Annals of the Entomological Society of Amer-

ica 23:461-466.

Rau, P. 1938. Studies in the ecology and behaviour of Polistes wasps. Bulletin of the Brooklyn Entomologi-

cal Society 33:224-235.

Reed, H. C., and P. J. Landolt. 1990a. Sex attraction in paper wasp, Polistes exclamans Viereck

(Hymenoptera: Vespidae), in a wind tunnel. Journal of Chemical Ecology 16:1277-1287.

Reed, H. C., and P. J. Landolt. 1990b. Queens of the southern yellowjacket Vespula squamosa, produce sex

attractant (Hymenoptera: Vespidae). Florida Entomologist 73:687-689.

Reed, H. C., and P. J. Landolt. 1991. Swarming of paper wasp (Hymenoptera: Vespidae) sexuals at towers

in Florida. Annals of the Entomological Society of America 84:628-635.

Shorey, H. H. 1977. Interactions of insects with their chemical environments, pp. 1-5, in H. H. Shorey and

J. J. McKelvey (eds.). Chemical Control of Insect Behaviour: Theory and Application. Wiley-

Interscience, New York, New York.

Starks, P. T. 2003. Natal nest discrimination in the paper wasp, Polistes dominulus. Annales Zoologici Fen-

nici 40: 53-60.

Strassmann, J. E. 1979. Honey caches help female paper wasp (P. annularis) survive Texas winters. Science

204:207-209.

Tobin, T. R., G. Seelinger, and W. J. Bell. 1981. Behavioral responses of Periplaneta americana to pe-

riplanone B, a synthetic component of the female sex pheromone. Journal of Chemical Ecology 7: 969-

979.

Wenzel, J. W. 1987. Male reproductive behaviour and mandibular glands in Polistes major (Hymenoptera:

Vespidae). Insectes Sociaux 34:44-57.

Wilson, E. O. 1971. The Insect Societies. Harvard University Press, Cambridge, Massachusetts.

Wyatt, T. D. 2003. Pheromones and Animal Behaviour: Communication by Smell and Taste. Cambridge

University Press, Cambridge, UK.

Zar, J. H. 1974. Biostatistical Analysis. Prentice-Hall, Inc. Englewood, New Jersey.

J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

J. ENTOMOL. SOc. BRIT. COLUMBIA 105, DECEMBER 2008

Efficacy of Isomate-CM/LR for management of leafrollers

by mating disruption in organic apple orchards

of western Canada

GARY J.R. JUDD!” and MARK G.T. GARDINER!

ABSTRACT

Results of a three-year study demonstrated that Isomate-CM/LR, a polyethylene, single

tube-type pheromone dispenser releasing an incomplete mixture of several species’

multi-component pheromones is an effective management tool that provides multiple-

species mating disruption for Choristoneura rosaceana (Harris) and Pandemis limitata

(Robinson). When applied at a rate of 500 dispensers / ha within the orchard and the

equivalent of 2000 dispensers / ha to trees on the orchard perimeter, levels of control

were adequate for production of organic apples in British Columbia, Canada. Trap

catches with synthetic pheromone lures were reduced by 79 -99% and mating of females

on mating tables was reduced by 87 - 98% in these species. At harvest, damage from

leafrollers in pheromone-treated organic orchards was below organically-acceptable

economic levels (5%) and similar to damage levels observed in insecticide-treated con-

ventional orchards (2%). Over three years, total trap catches of these two leafrollers and

their damage decreased in four of five orchards treated with pheromone, but catches and

damage from leafrollers increased in one orchard. These indices remained relatively

unchanged in paired insecticide-treated conventional orchards over the same three-year

period. In pheromone-treated orchards, levels of damage from leafrollers at harvest were

positively correlated with total leafroller catches in pheromone monitoring traps. Use of

Isomate-CM/LR as a supplemental pest management tactic in organic orchards will help

to reduce damage and economic losses from leafrollers that have been increasing under

the area-wide codling moth sterile insect programme ongoing in this semi-desert, mon-

tane apple production region.

Key Words: leafrollers, multiple-species mating disruption, organic apples

INTRODUCTION

Over the last decade a new paradigm for

integrated pest management in pome fruits

has emerged in western North America.

This transformation was driven by imple-

mentation of area-wide programmes to con-

trol codling moth, Cydia pomonella (L.),

using sterile insect technique (SIT) in Can-

ada (Dyck and Gardiner 1992) and phero-

mone-based mating disruption in the United

States (Calkins 1998). Application of these

species-specific controls for codling moth

has resulted in increasing damage from

secondary pests (Brunner et al. 1994,

Knight 1995, Gut and Brunner 1998). Con-

sistent with earlier prediction (Madsen and

Morgan 1970), several species of leaf-

rolling caterpillars (Lepidoptera: Tortrici-

dae) have become an increasing problem

when broad-spectrum insecticides that tar-

get codling moth, but which provide partial

control of leafrollers, are removed from the

production system (Madsen and Proctor

1985). Although the species complex varies

across production regions, increasing dam-

age from leafrollers in orchards using mat-

ing disruption for codling moth is a recur-

' Agriculture and Agri-food Canada, Pacific Agri-food Research Centre, Box 5000, 4200 Hwy 97, Summer-

land, British Columbia, Canada VOH 1Z0

? Author to whom correspondence should be sent. Tel.: +1 250 494 7711; fax.: +1 250 494 0755; E-mail:

Gary.Judd@agr.gc.ca

46 J. ENTOMOL. Soc. BRIT. COLUMBIA 105, DECEMBER 2008

ring problem seen around the world

(Charmillot 1990, Wearing et al. 1995,

Walker and Welter 2001).

The British Columbia (BC) organic ap-

ple industry is concentrated in the Simil-

kameen Valley (Judd et al. 1997, Mullinix

2005). Although orchards in this region

have been receiving sterile codling moth

since 1994 as part of an area-wide SIT con-

trol programme (Dyck and Gardiner 1992),

producers of organic fruit found it neces-

sary to supplement this programme with

codling moth mating-disruption technology

in spring (Judd and Gardiner 2005). Be-

cause codling moth is presently under con-

trol but profit margins are shrinking under

increased foreign competition, organic ap-

ple producers in BC are seeking alternative

controls for leafrollers to improve quality of

graded export fruit. Before 2005, Bacillus

thuringiensis Berliner (Bt) was the only

organic material available for controlling

leafrollers in Canada. Although Bt 1s effec-

tive, its use is often limited by inclement

spring weather in montane areas of BC, and

while fairly benign to beneficial species, it

can have a significant effect on parasites of

leafrollers if applied at the wrong time

(Cossentine et al. 2003).

Organic pome-fruit producers in BC

have been interested in pheromonal control

of leafrollers ever since pheromones were

first applied to control codling moth (Judd

et al. 1997). The idea of using one mating-

disruption system to simultaneously control

codling moth and leafrollers has been

around for some time (van Deventer ef al.

1992) but few commercial products exist.

Isomate-CM/LR, a multiple-species mat-

ing-disruption product designed to control

codling moth and leafroller species impor-

tant in western North America, was regis-

tered in the United States in 1997 and in

Canada in 2004 (Don Thomson, personal

communication). When Isomate-CM/LR

was used in conjunction with insecticides

(Knight et al. 1997, Knight et al. 2001),

apple orchards had 41% less leafroller dam-

age and received one less spray per season.

These same orchards consistently had less

codling moth damage than orchards receiv-

ing Isomate-C* and supplemental insecti-

cides. Whether leafrollers can be controlled

effectively, or sufficiently, with Isomate-

CM/LR when no supplemental insecticides

are used remains untested.

Judd and Gardiner (2005) reported on

the use of mating disruption as a supple-

mentary tactic for spring control of codling

moth in organic orchards that were part of

the area-wide SIT programme. Herein we

report results using Isomate-CM/LR to con-

trol damage from leafrollers while at the

same time supplementing codling moth

control in those same organic apple or-

chards. The primary objective of this study

was to conduct season-long assessments on

disruption of pheromonal communication

and mating in the leafrollers, Choristoneura

rosaceana (Harris) and Pandemis limitata

(Robinson) in commercial, organic apple

orchards where Isomate-CM/LR was used,

and to report on levels of leafroller damage

in the absence of insecticide inputs. Second,

we wanted to collect data on the relation-

ship between different measures of disrup-

tion and relative density of adult leafrollers

as measured by pheromone traps, because

questions about the efficacy of mating dis-

ruption and population density are rarely

addressed experimentally. Third, we

wanted to determine if pheromone trapping

of C. rosaceana and P. limitata has the po-

tential to be a predictive tool of potential

damage in orchards under pheromone-

based mating disruption (Walker and Wel-

ter 2001).

MATERIALS AND METHODS

Test orchards. All apple orchards used

in this study are located at Cawston, BC, in

the Similkameen Valley (latitude 49° 10.8’

N, longitude 119° 46.2” W, elevation 401

m). The five organic orchards treated with

Isomate-CM/LR and five paired conven-

tional orchards treated with insecticides

(Table 1) were described in detail by Judd

J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008 47

Table 1.

Management, monitoring and fruit sampling details for the leafrollers Choristoneura

rosaceana and Pandemis limitata in each organic (O1 - O5) and paired conventional (C1 - C5)

apple orchard studied in Cawston, BC, Canada, 1997 - 1999,

No. of pheromone’ Yearly insecti- Yearlynumberof Number of fruit

dispensers / ha cide treatments traps foreach sampled for damage

applied each year for leafrollers? _leafroller species’ at harvest each year

Orchard! 1997 1998 1999 1997 1998 1999 1997 1998 1999 1997 1998 1999

Ol 500 500 500 0 0 0 4 + + 3368 2145 2000

O2 500 500 500 0 0 0 6 6 6 5006 1756 2000

O3 500 500 500 0 0 0 4 4 4 4298 2100 2000

O04 500 500 500 0 0 0 Z Z 2 4300 2223 2000

O5 500 500 500 0 0 0 + 4 4 3031 1939 2000

Cl 0 0 0 0 1 _ 2 2 2270 1698 2000

C2 0 0 0 3 3 + _ Z Z 2000 2000 2000

C3 0 0 0 Z 2 3 _ Z Z 2000 2000 2000

C4 0 0 0 l 0 ~ Z Z 2100 2000 2000

C5 0 0 0 Z 3 4 ~ 2 2 2000 1883 2000

‘Orchards O1 to OS were cited by Judd and Gardiner (2005) as orchards Al - A4 and BI, re-

spectively, and orchards C1 - C5 remain the same across studies. Six untreated, organic, com-

parison orchards not listed were monitored but not sampled for damage (Cossentine e¢ al.

2004).

” All trees on the perimeter of each orchard received the equivalent of 2000 dispensers / ha.

*Leafroller sprays consisted of spring (April) sprays of methidathion in dormant oil and sum-

mer (July - August) sprays of Confirm ® (tebufenozide).

* No leafroller monitoring was done in conventional comparison orchards in 1997 but they

were sampled for damage at harvest. Six untreated, organic, comparison orchards (Cossentine

et al. 2004) were each monitored with two traps in 1998 and 1999. Each species was monitored

with a septal lure loaded with 3 mg of a multi-component blend described by Deland ef al.

(1994).

and Gardiner (2005). Six organic apple pyramid shape training system.

orchards that received no treatments for No synthetic insecticides were applied

control of leafrollers were described in de- to any of the organic orchards examined in

tail by Cossentine et al. (2004) as part of a _ this study (Table 1), but one orchard (O3)

study on parasitism of leafrollers in 1998 received Bt sprays (Dipel 2X DF) in 1998

and 1999. The latter untreated orchards and this is noted in the Discussion. All but

were used to compare relative trap catches one conventional orchard also received at

of leafrollers only, as no damage data were least one application of Guthion®

collected in the original study. Briefly, all (azinphosmethy! at 0.84 kg a.i./ha) for cod-

orchards ranged in size from 0.5 - 2 ha and ling moth control in 1997, but in later years

were composed of mixed apple varieties growers used Confirm” (tebufenozide) dur-

planted at densities of 267 - 938 trees/ha ing both spring and summer usually timed

with tree x row spacing of 2.4 - 6.1 x 3.0- for control of leafroller larvae. One caveat

6.1 m, respectively. Trees ranged in height is that conventional orchards did not neces-

from 2.5 - 3.5 m and were pruned using a _ sarily receive similar or optimal insecticide

48 J. ENTOMOL. Soc. BRIT. COLUMBIA 105, DECEMBER 2008

spray programmes because these orchards

were chosen by Judd and Gardiner (2005)

as local comparisons not controlled treat-

ments.

Pheromone’ disruption treatment.

Isomate-CM/LR is a brownish red, single-

tube, polyethylene dispenser, manufactured

by the Shin-etsu Chemical Company Ltd.

(Tokyo, Japan) and marketed by Pacific

Biocontrol Corporation (Vancouver, Wash-

ington). Each Isomate-CM/LR dispenser

contains a 285 mg blend of 36.9% (E£,E)-

8,10-dodecadien-l-ol (codlemone), 1.8%

isomers of codlemone, 6.0% dodecanol and

1.2% tetradecanol for disruption of codling

moth, and 43.5% (Z)-11-tetradecenyl ace-

tate (Z11-14:Ac) and 2.4% (E£)-11-

tetradecenyl acetate (£11-14:Ac) for dis-

ruption of leafrollers, plus 8.2% inert ingre-

dients and pheromone stabilizers (Don

Thomson, personal communication).

Isomate-CM/LR dispensers were deployed

in five organic apple orchards at a rate of

500/ha, however each perimeter tree re-

ceived the equivalent of 2000 dispensers/ha

(Table 1). Dispensers were attached to

branches in the upper third of tree canopies

ca. 1 m below the top of the central leader,

or on the first lateral branch beneath the

central leader. All pheromone dispensers

were deployed a few days before codling

moths were expected to emerge and no later

than 8 May each year (Judd and Gardiner

2005).

Monitoring seasonal flight activity of

moths. In all orchards, seasonal flight ac-

tivity and capture of adult leafrollers were

assessed using traps baited with synthetic

pheromones. Disruption of pheromone

communication in leafrollers was calculated

by expressing catches of moths in phero-

mone-treated organic orchards as a percent-

age of catch in either insecticide-treated

conventional orchards or untreated organic

orchards in 1998 and 1999. No comparisons

were made in 1997 as the latter sets of or-

chards were not monitored that season.

Depending on orchard size and shape, 2

- 6 Pherocon 1-C style open (5-cm side

Spacers) wing traps (Pherotech Interna-

tional, Delta, BC) baited with each species’

multi-component pheromone blend (Deland

et al. 1994; described below) were de-

ployed evenly throughout each orchard

(Table 1). One trap for each species was

hung ca. 1.5 - 2.0 m above ground on dif-

ferent sides of the same tree in 2 - 6 sepa-

rate trees. Positions for all traps remained

fixed within and across years. In 1997 only

the five Isomate-CM/LR-treated organic

orchards were monitored with pheromone

traps. On 30 May 1997, wing traps were

deployed in each of these organic orchards

and checked weekly from 6 June until 18

September. In 1998, wing traps were de-

ployed in these same five Isomate-CM/LR-

treated organic orchards, five paired insecti-

cide-treated conventional orchards and six

untreated organic orchards on 8 May, and

checked weekly from 15 May until 25 Sep-

tember. In 1999, wing traps were deployed

in all orchards on 27 May and checked

weekly from 3 June until 30 September.

Moths were counted and removed weekly

with trap bottoms replaced as needed and

pheromone lures changed every three

weeks.

Synthetic multi-component pheromone

lures for each leafroller species were pre-

pared with chemical components (Aldrich

Chemical Company Inc., Milwaukee, Wis-

consin) of known purity, as confirmed by

gas chromatographic analysis (Z11-14:Ac,

98% with 2% £E11-14:Ac; (Z)-11-

tetradecenal 96%; (Z)-11-tetradecanol;

97%, (Z)-9-tetradecenyl acetate, 96%) us-

ing published ratios (Roelofs et al. 1976,

Vakenti et al. 1988). In making pheromone

lures for each species, 200 ul of each multi-

component pheromone blend was dissolved

in dichloromethane and 3 mg of the phero-

mone blend was loaded into each red rubber

septum (Aldrich Chemical Company Inc.,

Milwaukee, Wisconsin). After loading,

septa were air-dried for ca. 18 h at 23 °C in

a fume hood and stored at 0 °C until pinned

to the inner side of trap lids in the field.

Assessment of mating in leafrollers.

Mating was assessed using laboratory-

reared, virgin, female moths placed in Tef-

lon®-lined mating tables described by

McBrien and Judd (1996). Both C.

J. ENTOMOL. Soc. BRIT. COLUMBIA 105, DECEMBER 2008

rosaceana and P. limitata were reared on a

modified pinto bean-based diet (Shorey and

Hale 1965) at 24 °C and 16:8 h L:D photo-

regime. Female pupae of each species were

placed individually in 150-ml plastic cups

provided with moist cotton wicks until

eclosion. Female moths aged 24 - 72 h were

immobilized at 0.5 °C and one forewing

and a tarsal tip were removed with fine for-

ceps to prevent escape from mating tables.

Females were kept chilled and transported

to field sites in small ice chests.

In 1998, mating activity of both leafrol-

ler species was assessed weekly from 2

June until 3 September in each pheromone-

and insecticide-treated orchard. During

each weekly assessment, one female of

each species was placed in 5 or 10 separate

trees in each, insecticide- or pheromone-

treated orchard, respectively, on Tuesdays,

Wednesdays and Thursdays (n = 15 or 30

females/species/orchard/week). Two mating

tables, each containing an individual female

of each species, were placed in opposite

sectors of the same tree, in the upper third

of the canopy, several rows and trees dis-

tant from any pheromone traps. Females

were placed in the field during the after-

noon and removed the following morning

to minimize predation and escape. Females

recovered from the field <24 h after deploy-

ment were returned to the laboratory and

each was dissected and examined for the

presence of a spermatophore in the bursa

copulatrix which indicates females have

mated. Any females that were dead when

collected from the field were omitted from

the data.

In 1999, we conducted an experiment

during flight of the first summer generation

to determine if the probability of mating

increased with the length of time (24 - 96 h)

females were exposed in the field in either

pheromone- or insecticide-treated orchards.

Starting on Monday, 21 June, 100 female P.

limitata were deployed in one pheromone-

treated orchard and another 100 females

were placed in an insecticide-treated or-

chard. At 24-h intervals for four consecu-

tive days, 25 females were recovered from

each orchard and returned to the laboratory

where their mating status was assessed as

before. This procedure was repeated for

several consecutive weeks.

Fruit damage sampling. Depending on

year and orchard, we examined fruit from

18 to 48 trees in the paired, pheromone- and

insecticide-treated orchards (Table 1) using

a stratified, cluster sampling procedure,

where the outer row of trees and all interior

trees represent two strata, and each tree

represents a cluster of fruit, respectively

(Judd et al. 1997). Sample trees were cho-

sen systematically by crossing each orchard

from corner to corner and edge to edge,

ensuring that each variety and stratum was

sampled. It was necessary to sample irregu-

lar numbers of trees and fruit from year to

year because biennial bearing in organic

orchards resulted in large annual differ-

ences in fruit set. All orchards were sam-

pled during normal periods of harvest for

each variety as fruit maturity and growers

dictated. In most cases, a minimum of 100

fruit were removed from each sample tree

by picking 50 low and 50 high fruit from

south side branches. If there were fewer

than 100 fruit on a tree, all fruit were re-

moved from the sample tree. Early- or late-

season leafroller damage, caused by either

overwintering or summer larvae, respec-

tively, can be distinguished by the degree of

surface tunneling and scarring of fruit. Only

late-season damage caused by summer-

feeding larvae is scored in this study. All

leafroller damage observed in this study

was caused by either C. rosaceana or P.

limitata, as no other species were previ-

ously found in this region (Madsen and

Madsen 1980, Judd and Gardiner 2004).

Our damage comparisons were limited to

pheromone vs. insecticide-treated orchards

because no damage samples were taken in

the untreated organic orchards examined by

Cossentine et al. (2004).

Statistical analyses. For each species,

moth captures from all traps in the same

orchard were pooled and transformed (logio

[x +1]) to normalize the data. Annual mean

total number of moths caught per trap in

untreated, Isomate-CM/LR-treated, and

insecticide-treated orchards were compared

50 J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

using an analysis of variance (ANOVA)

followed by a Student Neuman Keuls’ mul-

tiple comparisons test (Zar 1984), where

orchards are treated as replicates (n = 5 or

6). Mean weekly and seasonal total percent

mating of each species in pheromone- and

insecticide-treated orchards in 1998 were

compared using two-sample f-tests (n = 5).

Linear regression analysis was used to re-

late mean weekly percent mating and mean

weekly trap catches in 1998. The frequency

of mating among females placed in the field

for varying lengths of time in either a

pheromone-treated or insecticide-treated

orchard was compared weekly and season-

ally using contingency tables and y’ tests or

a binomial Z-test (Zar 1984). Mean percent

leafroller damage at harvest for pheromone-

and insecticide-treated orchards was com-

pared annually using a two sample f-test

following an arcsine Vp transformation of

raw data. Linear regression analysis was

used to relate mean percent damage at har-

vest in pheromone-treated orchards with

mean seasonal cumulative moth catches

using all three years of data (n = 15 data

pairs) and to examine changes in moth

catches and damage over time in organic

orchards. All statistical tests were per-

formed using SigmaStat® (Version 3.0.1,

SPSS Software Inc., San Jose, California)

and an experimental error rate of a = 0.05.

RESULTS

Seasonal flight activity of leafrollers.

Mean weekly catches of C. rosaceana and

P. limitata in orchards under different treat-

ment regimes in 1999 are shown in Fig. 1.

Similar weekly catches were seen in 1997

and 1998 but for brevity data are not

shown. Catches of both species reflect two

adult flight periods representing the first

and second generations in this region, re-

spectively. Weekly catches of C. rosaceana

under all treatment regimes tended to be

smaller during second generation than those

during first generation, but this trend was

often reversed in P. limitata (Fig. 1). P.

limitata appeared to be the most abundant

leafroller species in conventional orchards,

but the relative species makeup reversed

itself annually in untreated and pheromone-

treated organic orchards (Table 2).

Disruption of leafroller pheromone

trap catches. Weekly catches of C.

rosaceana were reduced 70 - 100% in the

Isomate—CM/LR treatment relative to both

untreated and insecticide-treated orchards

in 1999 (Fig. 1), and relative reductions

averaged 90.4 and 92.1%, respectively,

across years in 1998 and 1999 (Table 2).

Weekly catches of P. limitata were reduced

67 - 100 % by treatment with Isomate-CM/

LR compared to untreated and insecticide-

treated orchards (Fig. 1), and relative reduc-

tions averaged 92.5 and 87.2 %, respec-

tively, across years (Table 2). Catches of P.

limitata were always higher in insecticide-

treated conventional orchards compared

with untreated organic orchards (Table 2).

Mating in leafrollers. During the entire

1998 season, 35.1% of female C. rosaceana

(n = 757 females recovered) and 57.4% of

female P. limitata (n = 702) mated on mat-

ing tables hung in the insecticide-treated

orchards (Fig. 2). If 1 August is used as an

approximate starting point for second-

generation flight activity in both species

(Fig. 2), then mating of both species tended

to increase during the second-generation

flight period. In the insecticide-treated or-

chards mating of C. rosaceana during the

first (26.8%) and second generation

(54.3%) was significantly different (y? =

23.01, df = 1, P< 0.001). However, mating

of P. limitata during first (53.6 %) and sec-

ond generation (65.3%) was not signifi-

cantly different (y° = 2.05, df = 1, P =

0.152).

During the entire 1998 season only

1.6% of female C. rosaceana (n = 1610)

and 7.4% of female P. limitata (n = 1522)

mated on mating tables hung in the

Isomate-CM/LR-treated orchards (Fig. 2).

As seen in the absence of pheromone dis-

ruption (Fig. 2), significantly more C.

rosaceana mated during second generation

(3.5%) than during first generation (0.75%)

J. ENTOMOL. SOc. BRIT. COLUMBIA 105, DECEMBER 2008

EEL L! AEBAGLERILD? AALS SIEGES SATE LAIT

June 19

orchards

Pandemis fimitata

Mean number of moths caught / trap / week

July 17

Isomate-cM/LR L777]

July 17

Week

Choristoneura rosaceana

De ADE iy SB

Wi aS I sa DLE LEO ELE LOCC SETTLE

SS a a RS A SS SS A OT

” ELLY, {LTD OER ITE TELS, TELA, [LEAL AIS, [ELE DDI ODED ’ SOLEIL SLIT ILD f

Aug 14 Sept 11

ee)

Insecticide

orchards

Untreated

orchards

Aug 14

Sept 11

Figure 1. Mean weekly catches of two leafroller species in their species-specific synthetic

pheromone-baited traps hung in untreated organic apple orchards (n = 6), Isomate-CM/LR-

treated organic apple orchards (n = 5) and insecticide-treated conventional apple orchards (” =

5) in 1999,

(y° = 14.74, df = 1, P < 0.001) and mating

of P. limitata was also significantly greater

in the second generation (15.4%) than in

the first generation (2.9%) in pheromone-

treated orchards (7 = 65.14, df = 1, P <

0.001).

Regression analyses (Fig. 3) indicate

that weekly differences in mating of C.

rosaceana and P. limitata in the insecticide-

treated orchards were partially correlated

with the differences in weekly catches of

males in pheromone-baited traps, respec-

tively. In the Isomate-CM/LR-treated or-

chards, weekly differences in mating of

female P. limitata were explained (see r°

values) by differences in weekly catches of

males (Fig. 3), but both mating and catches

of C. rosaceana were too low in phero-

mone-treated orchards to ascribe any sig-

nificant relationship to these variables (P =

0.06).

Estimates of mating in P. limitata did

not appear to increase with increasing time

deployed in the orchard (Table 3). Compar-

ing results in Table 3 (1999) and Fig. 2

(1998), it appears that percent mating of

a2 J. ENTOMOL. SOc. BRIT. COLUMBIA 105, DECEMBER 2008

Table 2.

Seasonal total number of male leafroller moths, Choristoneura rosaceana and Pandemis limi-

tata, caught in synthetic pheromone-baited traps placed in untreated organic orchards (n = 6),

insecticide-treated conventional orchards (n = 5) and Isomate-CM/LR-treated organic orchards

(n = 5) and relative percent disruption of trap catches.

Mean = SE total number of Relative % trap

moths / trap / year / treatment! catch reduction’

Year Species Untreated Insecticide Isomate- Untreated Insecticide

CM/LR

1997 C. rosaceana — — 124.2 + 187.6 - -

P. limitata _ — 84.7 + 103.7 — -

1998 C.rosaceana 137.0+45.la 56.3 £21.6b 3.84 2,1¢ 93.3 O72

P. limitata 282.6+ 33.5b 460.3 +173.7a 66.2 +35.3c 85.6 76.6

1999 C.rosaceana 211.74+37.la 150.1473.la 27.8 +411.3b 87.5 86.9

P. limitata 101.7428.5b 386.8 +155.2a 2.3 +0.8c 99.4 OTe

Mean C. rosaceana 174.4+437.4a 103.2+46.9a 51.9 +36.8b 90.4446 92.14+5.1

oe P. limitata 192.2+90.5b 423.6436.7a 51.1 +24.9c 92.544.8 87.2 + 10.6

'Means within a row a followed by different letters are significantly different (P < 0.05) by

Student Neuman Keuls’ multiple comparisons test following significant (P < 0.05) ANOVA.

? Percent trap catch reduction in Isomate-CM/LR-treated organic orchards relative to catches in

untreated organic orchards or insecticide-treated conventional orchards.

female P. /imitata in the insecticide-treated age in pheromone-treated organic orchards

orchards was higher during the first genera- ranged from 1.7 - 24.8% in 1997, 0.4 -

tion in 1999 (82.9%) compared with 1998 4.2% in 1998, and 0.4 - 13.7% in 1999 (Fig.

(53.6%), while percent mating (2.9%) dur- 4A). From 1997 - 1999 leafroller damage

ing the first generation in the Isomate-CM/ declined in 4 of the 5 organic orchards and

LR-treated orchards was identical in 1998 in 12 out of 15 orchard-years damage was

(Fig. 2) and 1999 (Table 3). less than 5% at harvest under Isomate-CM/

Disruption of mating in leafrollers. LR treatment. Downward trends in leafrol-

When the entire 1998 season is considered __ler damage over three years appeared corre-

(Fig. 2), Isomate-CM/LR significantly re- lated with downward trends in total catches

duced mating of C. rosaceana by 95.4% (y? of leafrollers in each orchard, respectively

= 378.47, df = 1, P< 0.001) and mating of (Fig. 4B). There was a significant correla-

P. limitata by 87.1% (7 = 376.78, df=1,P tion (r = 0.65, P < 0.009) between total

< 0.001) relative to their mating in insecti- _leafroller trap catches and damage at har-

cide-treated orchards, respectively. Disrup- vest, but catches of leafrollers only ex-

tion of mating in both species tended to be _ plained 42.3% of the variation in harvest

lower during the second-generation flight damage (Fig. 4C). Comparison of leafroller

period, dropping to 93.5% in C. rosaceana damage in five organic orchards under

and 76.4% in P. limitata, respectively. Even management with Isomate-CM/LR, and

though percent mating of P. Jimitata in the _ five conventional orchards under various

insecticide-treated orchards was lower dur- _ insecticide programmes is shown in Fig. 5.

ing the first generation of 1998 (Fig. 2) than Damage levels were not significantly differ-

1999 (Table 3), disruption of mating by — ent between the two groups of pheromone-

treatment with Isomate-CM/LR was similar and insecticide-treated orchards in 1998 and

in 1998 (95.7%) and 1999 (96.4%). 1999 (Fig. 5).

Fruit damage. Summer leafroller dam-

J. ENTOMOL. Soc. BRIT. COLUMBIA 105, DECEMBER 2008

Choristoneura rosaceana

100

Insecticide-treated orchards

Seasonal mean mating = 38.8 + 5.7%

May 22 June 19 July 17 Aug. 14 Sept. 11

[ender]

Mating

100

Insecticide-treated orchards

Seasonal mean mating = 51.6 + 5.6%

8&0

Mean number of moths caught / trap / week

May 22 June 19

July 17

Week

Figure 2. Observed mean weekly percent mating of two leafroller species on mating tables

Aug. 14 = Sept. 11

hung in Isomate-CM/LR-treated organic apple orchards (n

Pandemis limitata

Isomate-CM/LR-treated orchards

Seasonal mean mating = 1.7 + 0.6%

6.0

3.0

May 22 June 19 July17 Aug. 14 Sept. 11

Catches

Isomate-CM/LR-treated orchards

Seasonal mean mating = 6.8 + 2.2%

Mean % mating / week

June 19

Aug. 14 Sept. 11

May 22 July 17

Week

5) and insecticide-treated

conventional apple orchards (7 = 5) and mean weekly catches of moths in synthetic phero-

mone traps in the same orchards in 1998.

DISCUSSION

Previously we showed that Isomate-CM/

LR was a useful spring-time supplement for

the codling moth SIT programme in BC

(Judd and Gardiner 2005), but its additional

benefits as a supplement for control of lea-

frollers in organic orchards was not de-

scribed. This study has demonstrated that

pheromone communication and mating in

sympatric leafroller moths commonly found

infesting organic apples in the Similkameen

Valley of BC can be effectively and simul-

taneously disrupted by releasing a mixture

54 J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

Choristoneura rosaceana

Insecticide-treated orchards

Mean % mating / week

% Mating = 5.56 + (7.06xcatch)

e = F- = 0.36, P= 0.009

0.0 2.0 5.0

Isomate-CM/LR-treated orchards

% Mating = -0.02 + (5.21xcatch)

r=0.31,P=0.06

0 v

0.00 0.25 0.50 0.75

Pandemis limitata

Insecticide-treated orchards

Mean % mating / week

% Mating = 12.53 + (1.06xcatch)

r=0.64, P=0.001

0 29 50

Isomate-CM/LR-treated orchards

% Mating = 0.076 + (1.56xcatch)

r= 0.85, P= 0.001

0 5 10 15

Mean number of moths caught / trap / week

Figure 3. Regression analyses showing relationships between relative moth density (trap

catches) and mating in two leafroller species in Isomate-CM/LR-treated organic apple orchards

(n = 5) and insecticide-treated conventional apple orchards (” = 5) in 1998.

of their major pheromone components from

Isomate-CM/LR. Season-long reductions of

pheromone trap catches of both C.

rosaceana and P. limitata with Isomate-

CM/LR were comparable to levels seen in

several studies examining each species indi-

vidually (Deland ef al. 1994, Agnello et al.

1996, Knight et al. 1998, Knight and

Turner 1999, Trimble and Appleby 2004)

and greater than levels observed in other

studies (Lawson ef al. 1996). There is no

generally accepted level for the reductions

in pheromone trap catches often observed

using mating disruption that correlate with

crop protection, but the observation has

been made that this reduction is usually 97 -

100% in species where disruption appears

to be an effective crop-protection tool

J. ENTOMOL. Soc. BRIT. COLUMBIA 105, DECEMBER 2008 55

Table 3.

Percentage of female Pandemis limitata mating in mating tables when placed in insecticide-

treated conventional and Isomate-CM/LR-treated organic apple orchards for increasing lengths

of time during first-generation flight in 1999.

Insecticide-treated orchard

Hours fenales —rroeeeoe

were infield —» females’ % mated’

24 274 86.5a

48 274 75.9a

2 242 80.6a

96 220 79.5a

Total 1010 82.9

(24 - 96h)

somate-CM/LR-treated orchard % mat-

f les! 9% Pe ing re-

n temales o mate ducton

246 2.03a 97.6*

252 4.36a 94.3*

224 311228 96.1*

220 221A 97.1*

942 2.97 96.4*

“n = total number of live females recovered from field in test period.

* Percentages within a column followed by the same letter are not significantly different (P >

0.05) based on a x’ test of the null hypothesis of equal mating frequencies across time catego-

ries.

> Asterisk indicates a significant (P < 0.001) reduction in mating based on a comparison of

paired proportions of mating within a row using a binomial Z-test (Zar 1984).

(Trimble and Appleby 2004). In many mat-

ing-disruption studies on C. rosaceana

(Reissig et al. 1978, Deland et al. 1994,

Agnello et al. 1996, Lawson et al. 1996,

Knight et al. 1998, Trimble and Appleby

2004), pheromone treatments have resulted

in less than 97% reduction in pheromone

trap catches relative to catches with the

same traps in insecticide-treated orchards.

In our study, a reduction of this magnitude

was achieved in 1998 when fewer than 4

moths were caught / trap / year (Table 2)

and a similar reduction was observed for P.

limitata in 1999 when catches of this moth

averaged fewer than 3 moths / trap / year.

Knight and Turner (1999) found a signifi-

cant negative relationship between mean

catches of Pandemis spp. / trap and percent

reduction of catches in synthetic pheromone

traps. A similar relationship has not been

reported for C. rosaceana, but our data

(Table 2) reflect this type of trend for both

species.

Presumably, reductions in pheromone

trap catches are correlated with reductions

in mating, but this is almost never con-

firmed in mating-disruption studies because

measures of mating are often missing. Ac-

tual reductions in female mating should be

more directly correlated with reductions in

damage from larvae than reductions in

males caught in pheromone traps. We found

relatively large correlations between trap

catches, a relative measure of population

density, and mating of P. /imitata on mating

tables (Fig. 3), but only a weak correlation

was found for C. rosaceana in the insecti-

cide-treated orchards, and no correlation

was found for this species in the phero-

mone-treated orchards (Fig. 3). Given that

catches of P. limitata represented about

50% of the total leafroller catch in organic

orchards during 1997 - 1999, and total

catches of leafrollers declined in Isomate-

CM/LR-treated orchards each year (Table

2), a significant, albeit weak, relationship

between total leafroller catches and damage

may be expected (Fig. 4C). A better rela-

tionship might be observed in orchards hav-

ing populations of P. limitata only. Never-

theless, the relationship shown in Fig. 4C is

consistent with the view that the reduction

in trap catches needs to be close to 99% (ca.

6 moths / trap / season) to ensure damage

from leafrollers is 1% or less, an acceptable

level for organic apple producers in BC.

We acknowledge that the relationship

we have shown between trap catches in

pheromone-treated organic orchards and

damage from summer-feeding leafrollers

may not hold true for conventional orchards

in this region, because there are significant

56 J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

Total % leafroller damage

Total leafrollers / trap / year

Total % leafroller damage

aye

% Damage = -0.5 + (0.6catch)

r? = 0.423 , P = 0.009

100 400

Total number of leafroller moths caught / trap / year

Figure 4. Percentage of damage at harvest caused by summer-feeding leafroller larvae in each

Isomate-CM/LR-treated organic apple orchard by year (A), total number of leafroller moths

(Choristoneura rosaceana and Pandemis limitata) caught in synthetic pheromone traps in each

Isomate-CM/LR-treated organic apple orchard by year (B), and linear regression of the ob-

served relationship between damage and total leafroller catches (log scales) in 1997 - 1999 (C).

differences in the levels of biological con-

trol in these different production systems.

In a_ study running parallel to ours,

Cossentine et al. (2004) found that summer

larval populations of both C. rosaceana and

P. limitata in untreated organic apple or-

chards in the Similkameen Valley experi-

enced parasitism rates as high as 68% dur-

ing 1998 and 1999, and higher levels of

parasitism were observed as_leafroller

populations declined. Given these observa-

tions, if mating disruption was causing lea-

froller populations to decline then it might

be expected to increase the impact of para-

sitoids and reduce damage. The general

absence of these natural control agents in

local conventional orchards (Joan

Cossentine, personal communication) may

invalidate any application of an established

relationship between trap catches and dam-

age from organic orchards where natural

controls are also acting. Mating disruption

of leafrollers in conventional orchards may

have to be augmented with insecticides,

J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

orchards

Mean + SE % summer leafroller damage

1997

LC] Isomate-CM/LR

1998

Insecticide

orchards

1999

Figure 5. Comparison of mean (+ SE) percent damage at harvest caused by summer-feeding

leafroller larvae in Isomate-CM/LR-treated organic apple orchards (m = 5) and insecticide-

treated conventional apple orchards (n = 5) in 1997 -1999. Paired means with the same letter

superscript are not significantly different (P > 0.05) by two-sample -test.

particularly where C. rosaceana is the

dominant species. With one exception

(Trimble and Appleby 2004), limited use of

selective insecticides in combination with

mating disruption has provided a small im-

provement in the control of C. rosaceana

over mating disruption alone (Agnello et al.

1996, Lawson ef al. 1996, Knight ef al.

1998, Knight et al. 2001).

To the best of our knowledge this study

is the only evaluation of mating disruption

of C. rosaceana and P. limitata with

Isomate-CM/LR in organic production sys-

tems where no insecticides were applied.

This technique holds promise for organic

pome fruit producers in the Similkameen

Valley, especially if P. limitata is the domi-

nant species. Our results are in sharp con-

trast to a failure of mating disruption to

keep damage from C. rosaceana below

economic levels in other regions even when

used in conjunction with pesticides

(Agnello et al. 1996, Lawson et al. 1996).

This failure has been attributed to high

population density and potential immigra-

tion of adults and larvae into treatment ar-

eas. Population density should be an impor-

tant factor in limiting the efficacy of mating

disruption, but this has seldom been shown

experimentally. Our data certainly point to

a strong relationship between relative adult

numbers, mating success and harvest dam-

age, but moth catches in our pheromone-

treated orchards were often greater than

those reported elsewhere so it is difficult to

reconcile our results on the basis of adult

population differences alone. Four of the

five organic orchards in this study were

somewhat isolated by having other orchards

located on only one border. Orchard O3

was the only one that was surrounded by

adjacent orchards, particularly cherries,

which were not treated for leafrollers, and it

was the one orchard in which we saw a

significant increase in catches of P. limitata

late in 1998 and damage in 1999 (Fig. 4).

Interestingly this was the only orchard that

received a petal-fall spray of Bt in spring of

1998. As noted by Knight et al. (1998),

immigration may be an important constraint

on use of mating disruption for leafrollers,

but monitoring may help to allay some of

this concern if it can predict immigration of

adults, as it did in orchard O3.

Although damage in the organic or-

chards was comparable to that seen in some

comparison insecticide-treated orchards

(Fig. 5), we make no claim that mating dis-

ruption is as effective as an optimal insecti-

cide-based control programme. However,

we are of the opinion that the efficacy of

mating disruption against any species is

58 J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

best evaluated over several years. Mating

disruption is certainly a less robust control

technique than most insecticides, and more

constrained by population density than the

latter. Suppression of codling moth popula-

tions using mating disruption often takes

several years and this will probably be true

of leafrollers (Fig. 4). Depending on the

comparison orchards chosen, it is possible

for mating-disruption technology to look

very effective, or highly ineffective; a better

approach may be to examine its long-term

effects in the same locations over several

years and compared with standard systems

as has been used for codling moth

(Charmillot 1990, Judd et al. 1997). The

long-term impact of mating disruption on

biological control of leafrollers in orchards

(Cossentine et al. 2004) relative to standard

controls also needs to be considered.

In areas of BC outside the Similkameen

Valley, control of leafrollers using phero-

mone-based mating disruption requires a

multi-species approach if different com-

plexes of sympatric leafrollers are to be

controlled effectively (Judd and McBrien

1995). For example, in the Okanagan and

Creston Valleys of BC, eye-spotted bud

moth, Spilonota ocellana (Denis and Schif-

fermiiller), European leafroller, Archips

rosanus (L.) and fruit-tree leafroller, Ar-

chips argyrospilus (Walker) are also impor-

tant pests of apple. The latter two species

use Z11-14:Ac as the major component in

their multi-component pheromone blends

(Arm ef al. 1982) and small-plot studies

(Deland 1992) demonstrated that phero-

mone communication and mating in A.

rosanus and A. argyrospilus could be dis-

rupted effectively with a pheromone blend

similar to that used in Isomate-CM/LR.

Spilonota ocellana, however, uses (Z)-8-

tetradecenyl acetate as its major pheromone

component (Arn et al. 1982) and would not

be controlled by Isomate-CM/LR, but can

be controlled by mating disruption

(McBrien et al. 1998). An Isomate dis-

penser containing this added ingredient is

currently under study as a _ mating-

disruption system for control of the entire

leafroller complex found in organic pome

fruit orchards in BC. With an organic for-

mulation of spinosad (Entrust®) registered

in Canada during 2005, the combined use

of this insecticide and mating disruption in

organic orchards also warrants study, be-

cause the impact of spinosad on parasites of

leafrollers in these systems needs to be con-

sidered carefully.

ACKNOWLEDGEMENTS

We thank the Similkameen-Okanagan

Organic Producers' Association (SOOPA)

and its cooperating members for allowing

us to conduct trials in their orchards. We

thank Janine Gartrell, Lila DeLury, Karen

Todd, and Nicole Verpaelst for technical

assistance. We especially thank Joan

Cossentine for providing trapping data on

leafrollers in untreated orchards, and Don

Thomson of Pacific Biocontrol Corporation

for making Isomate-CM/LR available for

testing and sharing technical data on this

product. This research was partially funded

by the Washington State Tree Fruit Re-

search Commission, SOOPA and the Agri-

culture and Agri-Food Canada Matching

Investment Initiative.

REFERENCES

Agnello, A.M., W.H. Reissig, S.M. Spangler, R.E. Charlton, and D.P. Kain. 1996. Trap response and fruit

damage by obliquebanded leafroller (Lepidoptera: Tortricidae) in pheromone-treated apple orchards in

New York. Environmental Entomology 25: 268-282.

Arn, H., M. Toth, and E. Priesner. 1982. List of sex pheromones of Lepidoptera and related attractants, ys

Edition. International Organization for Biological Control, Montfavet.

J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008 59

Brunner, J.F., L.J. Gut, and A.L. Knight. 1994. Transition of apple and pear orchards to a pheromone-based

pest management system. Proceedings of the Washington State Horticultural Association 88: 169-175.

Calkins, C.O. 1998. Review of the codling moth area-wide suppression program in the western United

States. Journal of Agricultural Entomology 15: 327-333.

Charmillot, P.-J. 1990. Mating disruption technique to control codling moth in western Switzerland, pp.

165-182. In Ridgway, R.L., Silverstein, R.M., Inscoe, M.N. (Eds.), Behavior-Modifying Chemicals for

insect Management: applications of pheromones and other attractants. Marcel Dekker, Inc., New York

and Basel.

Cossentine, J., L. Jensen, and K. Deglow. 2003. Strategy for orchard use of Bacillus thuringiensis while

minimizing impact on Choristoneura rosaceana parasitoids. Entomologia Experimentalis et Applicata

109: 205-210.

Cossentine, J., L. Jensen, K. Deglow, A. Bennett, H. Goulet, J. Huber, and J. O’Hara. 2004. The parasitoid

complex affecting Choristoneura rosaceana and Pandemis limitata in organically managed apple or-

chards. BioControl 49: 359-372.

Deland, J.-P. 1992. Mating disruption of fruittree leafroller, Archips argyrospilus, and effects on other lea-

froller species. Master of Pest Management Thesis. Department of Biological Sciences, Simon Fraser

University, Burnaby, British Columbia, Canada.

Deland, J.-P., G.J.R. Judd, and B.D. Roitberg. 1994. Disruption of pheromone communication in three sym-

patric leafroller (Lepidoptera: Tortricidae) pests of apple in British Columbia. Environmental Entomol-

ogy 23: 1084-1090.

Dyck, V.A. and M.G.T. Gardiner. 1992. Sterile-insect release programme to control the codling moth Cydia

pomonella (L.) (Lepidoptera: Olethreutidae) in British Columbia, Canada. Acta Phytopathologica et

Entomologica Hungarica 27: 219-222.

Gut, L.J. and J.F. Brunner. 1998. Pheromone-based management of codling moth (Lepidoptera: Tortricidae)

in Washington apple orchards. Journal of Agricultural Entomology 15: 387-405.

Judd, G.J.R. and M.G.T. Gardiner. 2004. Simultaneous disruption of pheromone communication and mating

in Cydia pomonella, Choristoneura rosaceana and Pandemis limitata (Lepidoptera: Tortricidae) using

Isomate-CM/LR in apple orchards. Journal of the Entomological Society of British Columbia 101: 3-13.

Judd, G.J.R. and M.G.T. Gardiner. 2005. Towards eradication of codling moth in British Columbia by com-

plimentary action of mating disruption, tree-banding and sterile insect technique: five-year study in or-

ganic orchards. Crop Protection 24: 718-733.

Judd, G.J.R. and H.L. McBrien. 1995. Pheromone-based management of pome fruit pests in British Colum-

bia: a multiple-species approach, pp. 9-17. Jn Research, development and commercialization of semio-

chemicals in insect pest management in Canada. Proceedings of The Entomological Societies of Mani-

toba and Canada, Workshop, Winnipeg, Manitoba, 1994.

Judd, G.J.R., M.G.T. Gardiner, and D.R. Thomson. 1997. Control of codling moth in organically-managed

apple orchards by combining pheromone-mediated mating disruption, post-harvest fruit removal and tree

banding. Entomologia Experimentalis et Applicata 83: 137-146.

Knight, A.L. 1995. The impact of codling moth (Lepidoptera: Tortricidae) mating disruption on apple pest

management in Yakima Valley, Washington. Journal of the Entomological Society of British Columbia

92: 29-38.

Knight, A.L. and J.E. Turner. 1999. Mating disruption of Pandemis spp. (Lepidoptera: Tortricidae). Envi-

ronmental Entomology 28: 81-87.

Knight, A., S. Cockfield, D. Thomson, and D. Neff. 1997. Developing mating disruption for leafrollers. /n

Proceedings of the 71 Annual Western Orchard Pest and Disease Management Conference, Portland,

Oregon, U.S.A. January 8-10, 1997. Western Orchard Pest and Disease Management Conference 71:

112-114.

Knight, A.L., D.R. Thomson, and S.D. Cockfield. 1998. Developing mating disruption of obliquebanded

leafroller (Lepidoptera: Tortricidae) in Washington State. Environmental Entomology 27: 1080-1088.

Knight, A.L., B.A. Christianson Jr., S. Cockfield, and J. Dunley. 2001. Areawide management of codling

moth and leafrollers. Good Fruit Grower 52(11): 31-33.

Lawson, D.S., W.H. Reissig, A.M. Agnello, J.P. Nyrop, and W.L. Roelofs. 1996. Interference with the

mate-finding communication system of the obliquebanded leafroller (Lepidoptera: Tortricidae) using

synthetic sex pheromones. Environmental Entomology 25: 895-905.

Madsen, H.F. and B.J. Madsen. 1980. Response of four leafroller species (Lepidoptera: Tortricidae) to sex

attractants in British Columbia orchards. The Canadian Entomologist 112: 427-430.

Madsen, H.F. and C.V.G. Morgan. 1970. Pome fruit pests and their control. Annual Review of Entomology

15: 295-320.

Madsen, H.F. and P.J. Procter. 1985. Insects and mites of tree fruits in British Columbia. British Columbia

Ministry of Agriculture and Food. Publication 82-6. Victoria, BC.

60 J. ENTOMOL. Soc. BRIT. COLUMBIA 105, DECEMBER 2008

McBrien, H.L. and G.J.R. Judd. 1996. Teflon®-walled mating table for assessing the efficacy of pheromone-

based mating disruption. Journal of the Entomological Society of British Columbia 93: 121-125.

McBrien, H.L., G.J.R. Judd, and J.H. Borden. 1998. Development of pheromone-mediated mating disrup-

tion for control of the eye-spotted bud moth, Spilonota ocellana. Entomologia Experimentalis et Appli-

cata 88: 101-107.

Mullinix, K. 2005. Pacific Northwest / Colorado / British Columbia - Regional Report, pp. 24-30. Jn Pro-

ceedings of the third national organic tree fruit research symposium, June 6-8, 2005, Chelan, WA, USA.

Reissig, W.H., M. Novak, and W.L. Roelofs. 1978. Orientation disruption of Argyrotaenia velutinana and

Choristoneura rosaceana male moths. Environmental Entomology 7: 631-632.

Roelofs, W., A. Cardé, A. Hill, and R. Cardé. 1976. Sex pheromone of the threelined leafroller, Pandemis

limitata. Environmental Entomology 5: 649-652.

Shorey, H.H. and R.L. Hale. 1965. Mass-rearing of the larvae of nine noctuid species on a simple artificial

medium. Journal of Economic Entomology 58: 522-524.

Trimble, R.M. and M.E. Appleby. 2004. Comparison of efficacy of programs using insecticide and insecti-

cide plus mating disruption for controlling the obliquebanded leafroller in apple (Lepidoptera: Tortrici-

dae). Journal of Economic Entomology 97: 518-524.

Vakenti, J.M., A.P. Guance, K.N. Slessor, G.G.S. King, A.A. Allan, H.F. Madsen, and J.H. Borden. 1988.

Sex pheromone components of the oblique-banded leafroller, Choristoneura rosaceana in the Okanagan

Valley of British Columbia. Journal of Chemical Ecology 14: 605-621.

van Deventer, P., A.K. Minks, L.H.M. Blommers, U. Neuman, and K. Jilderda. 1992. Mating disruption

utilizing lepidopterous sex pheromones: three years of testing in apple orchards in The Netherlands.

Brighton Crop Protection Conference — Pests and Diseases 3: 1193-1198.

Walker, K.R. and S.C. Welter. 2001. Potential for outbreaks of leafrollers (Lepidoptera: Tortricidae) in

California apple orchards using mating disruption for codling moth suppression. Journal of Economic

Entomology 94: 373-380.

Wearing, C.H., J.T.S. Walker, W.P. Thomas, J.R. Clearwater, D.M. Suckling, J.G. Charles, P.W. Shaw, V.

White, and G. Burnip, G. 1995. Pest control for organic apple production in New Zealand. The Orchard-

ist July: 22-27.

Zar, J.H. 1984. Biostatistical Analysis, Second edition. Prentice Hall, Englewood Cliffs, USA.

J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

Climate change and potential selection for

non-diapausing two-spotted spider mites on strawberry

in southwestern British Columbia

D.A. RAWORTH!

ABSTRACT

A validated model of the timing of post-diapause oviposition in the two-spotted spider

mite, Tetranychus urticae Koch, was used to predict when 50% of strawberry leaflets

with 7. urticae also have T. urticae eggs (100s) in each year from 1954 to 2006 at Lang-

ley, British Columbia. This timing was studied in relation to hours of frost occurring

before and after oviposition. Historically, IOo.5 occurred before there were frost-free

days, but there was a clear threshold at 390 h with temperatures < 0 °C after 100.5, which

was not exceeded. This suggests that there is selection pressure for early oviposition, but

also a limit to the extent of selection. The subzero temperature profile ~1 month before

oviposition was clearly different from that after 109.5. The number of hours with subzero

temperatures 1 month before oviposition, and the standard deviation of those estimates,

were negatively correlated with year and indicated that there could be oviposition in

January - rather than February - by 2015. Cumulative hours with temperatures < 0 °C

between 27 November (the empirical estimate of the time when T. urticae begins accu-

mulating degree-days for post-diapause oviposition) and 30 April was negatively corre-

lated with year, and extrapolation of a linear regression suggested that there could be

selection for continuous annual oviposition by 2050. There was considerable variation in

the data, but considered in combination with published evidence for climate change,

these results will be important in developing pest management strategies, and further-

more, will impact many aspects of agriculture in the Fraser Valley of British Columbia.

INTRODUCTION

Global warming (Intergovernmental (B.C.), Canada. This study considers

the

Panel on Climate Change 2007) will proba-

bly affect arthropod ranges (Gray 2004;

Logan and Powell 2004; Gutierrez et al.

2006; Musolin 2007) assuming fixed bio-

logical tolerances to environmental condi-

tions. At the same time, within a home

range, it will probably also affect the life

history characteristics of arthropods

through selection (cf. Bradshaw ef al.

2004). Winter diapause is a key feature of

temperate arthropods (Danks 2006). Dia-

pause characteristics will probably be af-

fected by global warming, and changes may

become evident first in areas that have a

relatively mild but temperate climate, such

as the Fraser Valley of British Columbia

effects of climate change on the timing of

initial, post-diapause oviposition (IO) by

two-spotted spider mites, Tetranychus urti-

cae Koch (Acari: Tetranychidae), on straw-

berry (Fragaria x ananassa_ Duch.

Rosaceae).

T. urticae females have a facultative

reproductive diapause. Diapause is induced

in the pre-imaginal stages, particularly at

the end of the protonymphal instar, by

short-day photoperiods (Veerman 1977a).

Termination is dependent on duration of

cold rest, temperature, and photoperiod

during the first few months of diapause

(Veerman 1977b). During winter, after

photoperiodic sensitivity is gone, diapause

' Agriculture and Agri-Food Canada, Pacific Agri-Food Research Centre, P.O. Box 1000, Agassiz, British

Columbia, VOM 1A0, Canada. e-mail: raworthd@agr.gc.ca

62 J. ENTOMOL. SOc. BRIT. COLUMBIA 105, DECEMBER 2008

would be sustained and prolonged by low

temperatures. Under long days at relatively

high temperatures, diapause can be termi-

nated without a cold rest period (Veerman

1977b), but under short days a cold rest

period is required.

Given a warmer climate there should be

selection for individuals who reproduce

continuously, despite short days, and there

may be selection for individuals who do not

respond to the initial photoperiodic induc-

tion cues. This idea is supported by the fact

that there is considerable variation in dia-

pause induction and termination character-

istics among 7. urticae strains, probably

associated with adaptation to local condi-

tions (Takafuji et al. 1991; Koveos et al.

1999).

Raworth (2007) developed and validated

a FORTRAN program (ProgIO) that deter-

mined the timing of post-diapause oviposi-

tion — the day when 50% of strawberry leaf-

lets with T. urticae also had T. urticae eggs

(100s). The oviposition model for the pro-

gram was based on field samples from

inland and coastal sites, collected during

1988 to 2006; estimates of 10); from 10

populations during 1988 to 2003 were used

to calibrate the model, and six independent

populations during 2004 to 2006 were used

for validation.

Here, ProglO and historical weather

data are used to examine 1) where, histori-

cally, T. urticae has placed 1095 with re-

spect to frost, and 2) the annual historical

trends in hours of frost. These trends are

then used to predict when 7. urticae can be

expected to commence oviposition in Janu-

ary — rather than the current timing in Feb-

ruary — and when 7. urticae may go through

the winter with no reproductive diapause. It

may be objected that viable leaves must be

available for feeding and oviposition; how-

ever, in this system mature green leaves

overwinter, and the mites usually initiate

feeding and oviposition on these leaves in

February and March (Raworth 2007).

MATERIALS AND METHODS

ProglO and meteorological data were

used to predict IOo.5 each year from 1954 to

2006. ProgIO first calculated the tempera-

ture under a strawberry leaf in a commer-

cial field at Langley, B. C. based on hourly

temperature and cloud opacity data meas-

ured by Environment Canada at Abbots-

ford, B. C. since 1953, and calibration

equations (Raworth 2007):

1. Cloud opacity = 1 (full cloud)

a. 0800-1600 h: y = -8.756 + 1.009 ¢ +

1.731 h - 0.013 # - 0.076 h° + 0.015 ¢Xh

R’ = 0.92, P< 0.0001, 425 df.

b. 1700-0700 h: y = -1.104 + 0.864 ¢ - 0.025

h+0.010r

R’ = 0.92, P< 0.0001, 862 d.f.

2. Cloud opacity = 0 (full sun)

a. 0800-1600 h: y = -54.250 + 1.116 ¢ +

9.515 h- 0.370 h°

R* = 0.85, P< 0.0001, 241 df.

b. 1700-0700 h: y = -3.848 + 1.153 ¢ +

0.066 h + 0.023 r

R’ = 0.92, P< 0.0001, 554 df.

where: y = temperature experienced by the

mites; ¢ = temperature in a Stevenson

Screen in the field (¢ = -1.141 + 1.043 tec,

where fec = air temperature at the Environ-

ment Canada station); and / = hour (where:

0800-1600 h = 8, 9,...16; and 1700-0700 h

= 17, 18,...24, 25, 26,...31). Temperatures

under a leaflet in intermediate cloud condi-

tions were determined by linear interpola-

tion between the results provided by the

equations for full cloud and full sun. The

timing of IOo5 was then determined from

thermal summations > 9.4 °C starting on an

empirically-derived day, 27 November

(Raworth 2007), and a thermal requirement

(vy) that was negatively correlated with ac-

cumulated cold-rest hours < 4 °C (x)

summed from 27 November (Raworth

2007):

3. y= 78.3 - 0.0279 x; r° = 0.83, P= 0.01,

4A df.

Equation 3 implies that, for equivalent

rates of thermal summation, the spider

mites will stay in diapause longer if they

have had insufficient cold rest. The timing

J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

of IOo.5; was compared graphically with the

cumulative daily hours of frost below 0 °C

summed between 27 November and 30

April of the following year.

To determine what temperature condi-

tions the mites have avoided, annual fre-

quency was plotted against the number of

hours of frost below 0, -1, -2,...-10 °C that

remained after IOo,5 (3-d plot), and the

hours of frost below 0, -1, -2,...-10 °C that

occurred one month before mite eggs would

be observed in the field, 10 January to 10

February (3-d plot). To determine when

conditions would be suitable for earlier

occurred between 10 January and 10 Febru-

ary were regressed (SAS Institute 2004)

against year and the regression was ex-

trapolated beyond 2006. Changes in vari-

ability as a function of year were deter-

mined by pooling the latter data for each

decade and regressing 1 SD for mean hours

below a given temperature in that decade

against the median year. Finally, a similar

technique was used to determine when con-

ditions would be suitable for T. urticae to

pass through the winter without reproduc-

tive diapause based on the annual sum of

the hours of frost below 0 °C between 27

oviposition by 7. urticae, the annual sum of | November and 30 April.

hours of frost below 0, -2, -4,...-10 °C that

RESULTS AND DISCUSSION

Cumulative hours of frost below 0 °C

between 27 November and IO, ;, from 1954

to 2006 varied between 400 and 1600 h

(Fig. 1B, triangles). Despite this variation,

100.5 always occurred before frost-free days

had begun (Fig. 1A, B), but with a clear

upper threshold of 390 h of frost remaining

after 1095. This indicates sensitivity to

frost, because all the estimates of IOp;5 are

clustered below 390 h; it also indicates

some selection pressure for early oviposi-

tion, because the mites do not wait until

frost-free conditions occur. As long as a

female’s progeny can survive and go on to

reproduce, a female that initiates reproduc-

tion early in the season should have a nu-

merical advantage over one that initiates

reproduction later. This result should be

qualified. The 390 h threshold was deter-

mined from a model of post-diapause ovi-

position in southwestern B. C.; it would not

be expected to apply to 7. urticae popula-

tions that are adapted to different local con-

ditions in other temperate regions. Such a

generalization would require further re-

search.

Comparison of the subzero temperature

profiles before and after IOo.5 suggests that

T. urticae in southwestern B. C. is able to

tolerate >200 h at -2 °C, but only 20 h at -6

°C (Fig. 2). Lower temperatures for longer

periods (Fig. 3) have been avoided. How-

ever, the number of hours with tempera-

tures below 0, -2, ... -10 °C, 1 month before

T. urticae normally commences oviposi-

tion, have declined significantly since 1954

(Fig. 4). Taking -8 °C as a critical tempera-

ture associated with no oviposition (Fig. 2),

the data indicate that an average year will

have zero hours <-8 °C by 2015 (Fig. 4). At

this time there could be reduced selection

pressure against emergence, and hence ovi-

position in January rather than February.

Because the regression predicts hours at a

given temperature in an average year, one

would expect some hours at temperatures

<-8 °C in some years, and possible mortal-

ity of early-emerging mites. However, the

variation in subzero temperatures during

this 1 month period has also declined since

1954 (Fig. 5) so that there will be less un-

certainty about subzero conditions in 2015

than there was in 1957, and reduced selec-

tion pressure against early emergence. The

regression of total hours of frost below 0 °C

from 27 November to 30 April against year

suggests that the number of hours of frost

will decrease to the threshold of 390 h by

~2050 (Fig. 6). At this point, selection for

early oviposition could result in reproduc-

tion by some individuals right through the

winter in southwestern B.C.

The objective of this study was to exam-

ine general patterns in the initiation of post-

800-4:;.'

Hours < 0°C

400

eae rp ! z

eon

Pi he Or

Tc. ma Eset a

ct OE 3.)

sift tee a et

gees:

t%e, ° Pee ‘Pun, °

a cae EES eet ae,

c teed. peat rin Tae . ee.

J. ENTOMOL. SOC

Apr

. BRIT. COLUMBIA 105, DECEMBER 2008

Figure 1. Cumulative hours with temperatures < 0 °C under a strawberry leaflet in a commer-

cial field from: (A) start date to 30 April versus start date (dots); (B) 27 November to the date

when 50% of strawberry leaflets with 7. urticae also have T. urticae eggs (IOs) versus IOo.5

(triangles); and (B) 109.5 to 30 April versus IOo,5 (circles), each year from 1954 to 2006. The

horizontal line represents a threshold number of hours < 0 °C remaining after IOg.s.

Number of years

0 aN Z\

34 Ow,

2, RSA

Sk SS SSS

em: SSSSSSSS SSS EUS

%, SSS

Figure 2. Number of years with a given number of hours at subzero temperatures under a

strawberry leaflet in a commercial field, after the date when 50% of strawberry leaflets with T.

urticae also have T. urticae eggs.

J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008 65

}

fee)

Number of years

: 4

mm 2\

rS)

% -A KY

©, <S

Ny, XO 0

“OX (Coen. <

1020” 18 pours (x20)

Figure 3. Number of years with a given number of hours at subzero temperatures under a

strawberry leaflet in a commercial field, ~1 month before the date when T. urticae commences

Oviposition (10 January to 10 February).

500

400--..

1004

1960 1980 2000 2020 2040

Figure 4. Hours with subzero temperatures under a strawberry leaflet in a commercial field,

between 10 January and 10 February, versus year. Solid lines are linear regressions (7* = 0.10,

0.08, 0.08, 0.09, 0.10, 0.14 and P = 0.02, 0.03, 0.03, 0.02, 0.02, 0.004) for temperatures < -10,

-8, ... 0, respectively; data for temperatures <-4 °C are shown.

66 J. ENTOMOL. SOc. BRIT. COLUMBIA 105, DECEMBER 2008

200

—

Hours (1 SD)

1960 1980 2000 2020

Figure 5. One SD of mean hours with subzero temperatures under a strawberry leaflet in a

commercial field, between 10 January and 10 February, versus median year. Solid lines are

linear regressions (7 = 0.95, 0.92, 0.89, 0.87, 0.93, 0.80 and P = 0.0008, 0.002, 0.005, 0.007,

0.002, 0.02) for temperatures < -10, -8, ... 0, respectively.

Hours < 0°C

1960 1980 2000 2020 2040

Figure 6. Hours with temperatures < 0 °C under a strawberry leaflet in a commercial field,

from 27 November to 30 April, versus year. Solid line is a linear regression y = 19600. — 9.37

x; 1° = 0.22; P< 0.001.

diapause oviposition in 7. urticae in rela- populations in southwestern B. C.: a maxi-

tion to field temperatures. A number of mum of 390 h of frost after 109.5; post-IO9,5

thresholds were observed for spider mite tolerance to >200 h at -2 °C, but only 20 h

J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

at -6 °C; and -8 °C was taken as the critical

temperature after 1095 associated with no

oviposition. These thresholds could be con-

firmed by further work, but this was not the

point of the study; regardless of the exact

value of the thresholds, the patterns are

clear. Oviposition is initiated before frost-

free days occur — in ProgIO and ipso facto

in the field data used to build ProglO; the

number of hours of frost 1 month before

Oviposition, and variation in those estimates

among years, has decreased significantly

during the last half century; and the number

of hours of frost from 27 November

through 30 April have also decreased sig-

nificantly over the years.

It is clearly risky to extrapolate from a

linear regression based on data with consid-

erable scatter (Figs. 4, 6); the relationship

may be negative but asymptotic at, for ex-

ample, 600 h frost (Fig. 6). However, there

is additional evidence. Although it is diffi-

cult to attribute observed temperature

changes to natural or human causes at

smaller than continental scales because

factors such as land use change and pollu-

tion complicate the picture (Inter-

governmental Panel on Climate Change

2007), the trends observed in the current

study are consistent with the global warm-

ing scenario. The mechanism driving global

warming, namely increasing levels of

greenhouse gases (GHG) (N20, CHg, and

CO) is well established, and ‘With current

climate change mitigation policies and re-

lated sustainable development practices,

global GHG emissions will continue to

grow over the next few decades’ (Inter-

governmental Panel on Climate Change

2007); a global temperature change of +0.2

°C per decade is projected. Therefore, the

linear extrapolations in Figs. 4 and 6 may

be reasonable. Time will tell, however if

correct, oviposition in January should be

observable within the next decade.

Despite the many uncertainties in this

study, there is sufficient evidence for earlier

post-diapause oviposition in 7. uwurticae

within the relatively near future in south-

western B. C. to at least consider it in plan-

ning spider mite monitoring and manage-

ment activities. This would evolve naturally

into planning for continuous annual ovi-

position should that occur in 4 to 5 decades.

Furthermore, a relatively rapid reduction in

the number of hours of subzero tempera-

tures during the winter will have significant

implications for many aspects of agriculture

in the Fraser Valley, including arthropod

pest and disease management, crop produc-

tion, and crop selection in both field and

greenhouse environments. These effects

need to be considered carefully by growers,

pest managers, researchers, and government

planners at Provincial and Federal levels

with studies and approaches to address po-

tential problems evolving as trends become

increasingly certain.

ACKNOWLEDGEMENTS

I thank M. Petrou (Environment Can-

ada) for meteorological data, and two

anonymous reviewers for their thoughtful

comments. Pacific Agri-Food Research

Centre contribution # 774.

REFERENCES

Bradshaw, W.E., P.A. Zani and C.M. Holzapfel. 2004. Adaptation to temperate climates. Evolution 58:

1748-1762.

Danks, H.V. 2006. Key themes in the study of seasonal adaptations in insects II. Life-cycle patterns. Ap-

plied Entomology and Zoology 41: 1-13.

Gray, D.R. 2004. The gypsy moth life stage model: landscape-wide estimates of gypsy moth establishment

using a multi-generational phenology model. Ecological Modelling 176: 155-171.

Gutierrez, A.P., T. D'Oultremont, C.K. Ellis and L. Ponti. 2006. Climatic limits of pink bollworm in Ari-

zona and California: effects of climate warming. Acta Oecologica 30: 353-364.

68 J. ENTOMOL. Soc. BRIT. COLUMBIA 105, DECEMBER 2008

Intergovernmental Panel on Climate Change. 2007. Fourth Assessment Report, Climate Change 2007: Syn-

thesis Report, AR4 SYR — Topic 6. http://www. ipcc.ch/pdf/assessment-report/ar4/syr/ar4_syr_topic6.pdf

Koveos, D.S., A. Veerman, G.D. Broufas and A. Exarhou. 1999. Altitudinal and latitudinal variation in

diapause characteristics in the spider mite Tetranychus urticae Koch. Entomological Science 2: 607-613.

Logan, J.A. and J.A. Powell. 2004. Modelling mountain pine beetle phenological response to temperature.

Information Report Pacific Forestry Centre, Canadian Forest Service (BC X 399): 210-222.

Musolin, D.L. 2007. Insects in a warmer world: ecological, physiological and life-history

responses of true bugs (Heteroptera) to climate change. Global Change Biology 13: 1565-1585.

Raworth, D.A. 2007. Initiation of oviposition after winter diapause in the spider mite Tetranychus urticae

(Acari: Tetranychidae): prediction and historical patterns. Population Ecology 49: 201-210.

SAS Institute. 2004. User’s manual, version 9.1. SAS Institute, Cary, NC.

Takafuyi, A., P.M. So and N. Tsuno. 1991. Inter- and intra-population variations in diapause attributes of the

two-spotted spider mite, Tetranychus urticae Koch, in Japan. Researches on Population Ecology 33: 331-

344.

Veerman, A. 1977a. Aspects of the induction of diapause in a laboratory strain of the mite Tetranychus

urticae. Journal of Insect Physiology 23: 703-711.

Veerman, A. 1977b. Photoperiodic termination of diapause in spider mites. Nature 266: 526-527.

J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008 69

Biology and management of bark beetles (Coleoptera:

Curculionidae) in Washington cherry orchards

MICHAEL D. DOERR'”, JAY F. BRUNNER'

and TIMOTHY J. SMITH”

ABSTRACT

The biology and management of bark beetles (Coleoptera: Curculionidae, Scolytinae) in

Washington cherry orchards was investigated from 2003-2005. Two dominant species

were identified attacking cherry (Prunus spp.) orchards: the shothole borer, Scolytus

rugulosus Miller, and an ambrosia beetle, Xyleborinus saxeseni Ratzeburg. S. rugulosus

was the species most implicated in damage to healthy trees. Two distinct periods of S.

rugulosus activity occur in Washington, with a possible partial third in some locations.

The first activity period begins in late April and peaks in late May to early June, with the

second beginning in mid-July and peaks in late July to early August. Yellow sticky traps

(unbaited apple maggot traps) were effective tools to monitor S. rugulosus activity but

ethanol-baited intercept-style traps were necessary to monitor X. saxeseni activity.

Movement of S. rugulosus into orchards was closely associated with emergence from

outside hosts, generally a pile of recently pruned or cut wood placed outside the orchard.

S. rugulosus readily moved distances of 10-50 m to attack trees on orchard borders, but

did not move more than two or three rows into a healthy orchard. A residue bioassay

technique demonstrated that several insecticides caused mortality of S. rugulosus adults.

A pyrethroid, esfenvalerate, was the most active 21 d after treatment. Azinphos-methyl

was acutely toxic to S. rugulosus, but for only seven d. Endosulfan and the neonicoti-

nyls, thiamethoxam and acetamiprid, were somewhat toxic to S. rugulosus.

Key Words: bark beetles, Scolytus rugulosus, Coleoptera, Curculionidae, Scolytinae,

ambrosia beetle, Xy/eborinus saxeseni

INTRODUCTION

Bark beetles (Coleoptera: Curculioni-

dae, Scolytinae) have historically been re-

ported as pests of pome and stone fruit

(Kirk 1969, Linsley and MacLeod 1942,

Mendel et al. 1987, Payne 1977, Smith

1932). They are commonly described as

attacking weakened trees and causing limb

or even tree death if present in high enough

numbers (Lindeman 1978). Nutritionally

stressed trees, or those damaged by sun

scald or winter freezing may provide points

of access into orchards for opportunistic

beetles (Bhagwandin 1992). Health of trees

is important to the natural plant defense

against bark beetle attack. High plant cell

turgor pressure through proper soil water

availability may allow trees to mechanically

flood out or chemically repel potential colo-

nizers through increased sap flow at the site

of attack (Rudinsky 1962, Berryman 1972).

The use of synthetic organic insecticides

has likely mitigated problems with bark

beetles in tree fruit orchards, and until re-

cently they have been considered sporadic

and localized pests (Beers ef al. 1993).

However, the reported incidence of injury

from bark beetles in Washington stone fruit

orchards has been increasing (Brunner

'Washington State University Tree Fruit Research and Extension Center, 1100 N. Western Ave., We-

natchee, WA 98801, USA

: Washington State University Cooperative Extension, 303 Palouse St., Wenatchee, WA 98801, USA

: Corresponding author. E-mail: mdoerr@wsu.edu

70 J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

2003). Anecdotal reports attributed injury

to beetles moving into orchards, especially

cherry orchards, from outside hosts and

attacking healthy trees. The damage most

often noted is the boring of beetles at the

base of buds by pioneer beetles, causing

affected buds to die. Continuous and re-

peated attacks eventually weaken otherwise

healthy trees and make them susceptible to

secondary attacks from either conspecifics

or possibly other wood boring beetles. Es-

pecially vulnerable are new plantings of

young cherry trees. Economic difficulties in

Washington’s fruit industry during the late

1990s likely contributed to the bark beetle

problem through an increased occurrence of

neglected or abandoned orchards providing

suitable host material for bark beetle repro-

duction (Warner 2006, Mendel e¢ a/. 1987).

Initial observations (2001-02) of bark

beetle damage in cherry orchards indicated

a need to further explore certain aspects of

their biology and management. With little

or no published information from Washing-

ton, species identifications and verification

of life histories were required for all bark

beetles infesting Washington cherry or-

chards. Preliminary observations indicated

that the main beetle species was the shot-

hole borer, Scolytus sp., however at least

one other species, possibly an ambrosia

beetle, was also involved in attacking

healthy cherry trees. While many other

wood boring beetles were observed in and

around infested hosts, their role in damage

to healthy trees was either unknown or

unlikely based on what was known of their

natural history.

Pests invading orchards from an exter-

nal host represents a significant challenge

to the timing of chemical controls. Knowl-

edge of the pest’s development in host

plants is needed along with its ability to

migrate from these hosts into orchards.

Adult traps have proven useful for monitor-

ing bark beetles in other orchard or natural

systems (Kovach and Gorsuch 1985, Mar-

kalas and Kalapanida 1997), but research is

needed to identify and optimize monitoring

systems for tree fruit pest management pro-

grams. Information is also needed on how

much of an orchard requires protection

from bark beetles, over what time periods,

and which insecticides would be effective

at providing required protection.

This paper provides new knowledge that

will help Washington cherry growers man-

age bark beetle problems. The key species

involved in attacking pome and stone fruit

trees were identified along with a clear un-

derstanding of their seasonal life history.

We also developed methods of monitoring

bark beetles, and determined the distance

bark beetles moved from a natal food host

to attack healthy orchard trees. A bioassay

technique was developed for assessing rela-

tive toxicity of candidate insecticides, and

we documented successful control strate-

gies used to manage bark beetles in heavily

infested orchards.

MATERIALS AND METHODS

Bark beetle identification, monitor-

ing, and life history. Bark beetles and

other wood boring Coleoptera infesting

Washington cherry orchards were identified

by a combination of rearing adults from

host wood infested with immature larvae in

emergence cages and trapping adults near

suspected host sites and along orchard bor-

ders. All insects collected in the following

trials were stored in alcohol and later identi-

fied to family (Dr. Christian Krupke, Pur-

due University; Dale Whaley, Washington

State University; Michael Doerr, Washing-

ton State University). All Scolytinae and

associated parasitoids were sent to Malcom

Furniss, (Entomologist Emeritus, Univer-

sity of Idaho) for identification. All Coleop-

tera collected from emergence cages and

adult traps were identified in 2003. In 2004

only Scolytinae were submitted for identifi-

cation. By 2005 it was apparent that Scoly-

tus spp. were the dominant bark beetles

present in Washington cherry orchards so

identification was further limited to those

J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

species.

Emergence cages were used to identify a

species:host relationship. Infested wood

from four sites was collected during the

spring and summer in 2003, placed in

opaque cardboard boxes (50 x 50 x 30 cm),

and held under laboratory conditions (22 +

2 °C). One glass vial (2.5 cm diameter x 10

cm) was placed through a hole in each

emergence box. Emerging beetles (and Hy-

menopteran parasitoids) where attracted to

the light coming through the opening in the

box and entered the vial. Beetles and natu-

ral enemies that entered the vial were re-

moved daily.

Adult traps were placed near infested

wood piles outside of orchard blocks

(referred to below as ‘outside hosts’) and on

the orchard border closest to the outside

host at sixteen locations in north-central

Washington from 2003-05 (twenty one or-

chard-yr equivalents). No specific protocols

were followed across all sites, but rather an

effort was made to ensure that trap place-

ments sufficiently covered the threatened

area of each orchard border and encircled

outside hosts. Considerations had to be

made depending on the size of each loca-

tion. Generally, traps were placed approxi-

mately 10 m apart on orchard borders and

hung directly in the trees at a height of 2 m.

At least four traps were placed around a

suspected outside host. If circling a host

was not possible, traps were placed ap-

proximately 10 m apart across the length of

the host. Most often traps were hung di-

rectly from host material, but it was some-

times necessary to hang them from a 2 m

tall post that was placed adjacent to the

host. Monitoring efforts in 2003 focused on

identifying the best available trap and lure

system. Commercially available intercept-

style traps (Lindgren Funnel Trap, 8-

funnnels, Phero Tech, Inc., Delta, British

Columbia, Canada; Pane Intercept Trap,

IPM Technologies, Inc., Portland, OR),

either with or without an ethanol attractant,

and un-baited yellow sticky traps (Pherocon

AM, Tréceé, Inc., Adair, OK) were evalu-

ated in trials replicated across several loca-

tions for their ability to monitor adult activ-

ity at an outside host and at a nearby or-

chard. A 12.5 cm? DvDP kill strip

(Vaportape II insecticidal strips, Hercon

Environmental Co, Emigsville, PA) was

placed in the collection container of the

intercept-style traps to kill beetles and pre-

vent their escape.

In 2003, a direct comparison was made

between the Lindgren Funnel Trap and the

Pane Intercept Trap at six locations. Each

trap type was baited with the respective

manufacturer’s commercially available

ethanol lure. Traps were placed on 15 April

and monitored every seven d until 15 Oct.

Lures were replaced at six-wk intervals,

based on manufacturer recommendations. A

direct comparison to evaluate the effective-

ness of the ethanol attractant in the

Lindgren Funnel Trap was also made at

four locations in 2003. Traps were placed

on 15 April and monitored every seven d

for six weeks. In 2003 and 2004, a direct

comparison was made between a Lindgren

Funnel Trap baited with an ethanol lure and

an unbaited yellow sticky trap at ten loca-

tions. In 2004, traps were placed on 1 Mar

and monitored every seven d through Octo-

ber. The ethanol lures were replaced at six-

wk intervals. Two traps of each treatment

were placed in an alternating pattern at all

locations. Season-long captures of the

dominant Scolytinae species were averaged

for the two traps at each location. Due to

high variability in populations between

locations, trap capture data from the paired

comparisons were analyzed by a Wilcoxon

Rank Test (P=0.05) (Wilcoxon 1945) using

JMP statistical software (JMP v. 5.1.2

2004).

Adult trap data gathered from the loca-

tions with the highest populations (15 or-

chard-yr for S. rugulosus and five orchard-

yt for X. saxeseni) were used to plot cumu-

lative emergence of the dominant species

for each of the generations. With no tem-

perature-dependent developmental (degree-

day) data available, the only point of refer-

ence between years was Julian days. Cumu-

lative emergence at each date was averaged

and plotted with the raw data from each

orchard site. Julian days were then con-

72 J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

verted back to calendar days for ease of

reference.

Scolytus rugulosus migration and

damage distribution. Two orchards in

2004 and two in 2005 were identified where

host wood piles that were heavily infested

with S. rugulosus were threatening nearby

healthy orchards (<50 m). Yellow sticky

traps were placed by the hosts located out-

side the orchards to track adult emergence

and on the orchard borders to monitor 1m-

migration. Cumulative capture percentiles

for an entire S. rugu/osus generation at the

outside host and on the orchard border were

plotted together for each study site. If cu-

mulative percentiles were identical at the

host and the orchard this would suggest that

adult dispersal to a suitable feeding or re-

productive site occurred immediately after

emergence. However, dramatic shifts in

cumulative percentiles would indicate e1-

ther a delay in migration from the outside

host or a constant or prolonged immigration

into the orchard from multiple outside

hosts. Each location could only be moni-

tored for one generation because we al-

lowed growers to remove the natal host and

protect their orchard following our observa-

tions.

At the same locations described above,

damage to healthy trees in the orchard was

monitored by visually inspecting trees.

Every tree along the border row and then

every tree in subsequent rows moving into

the orchard away from the outside host

were monitored for S. rugulosus damage

until no further damage was noted. The

total number of trees sampled varied at each

orchard (Site 1 — 3 rows x 19 trees, Site 2 —

5 rows x 8 trees, Site 3 — 4 rows x 12 trees,

Site 4 — 7 rows x 15 trees). Twenty growing

shoots were randomly selected from each

tree and the total number of shoots exhibit-

ing wilting or flagging foliage (visually

confirmed to be caused by S. rugulosus

burrowing) was recorded. We calculated

the total number of damaged shoots at each

site, then noted what percentage of that

total was found in the row closest to the

outside host (row one) and each subsequent

row moving away from the outside host.

Scolytus rugulosus insecticide screen-

ing. Insecticides were evaluated using

newly emerged Scolytus rugulosus adults in

2004. The insecticides outlined in the Re-

sults and Discussion section included the

majority of those recommended for use on

cherries in Washington (Smith et al. 2004).

Although the insecticides chosen for this

trial were those available on cherry, mature

Delicious apple trees at WSU-TFREC were

the only trees readily available for this test.

The trees were treated with various insecti-

cides at the manufacturers’ recommended

rates. All treatments were applied with a

handgun sprayer at 300 psi to drip, simulat-

ing a full dilute spray. Treatments were

applied to one-tree plots replicated three

times in a randomized complete block. A

one-tree buffer (unsprayed tree) was left

between each replicate to reduce over-spray

and drift. Treated apple branches, approxi-

mately 15 cm long x 1.25 cm diameter sec-

tions of two-yr-old wood, were collected at

1, 7, 14 and 21 d after treatment (DAT),

returned to the laboratory and stored at 2 °C

until new adults could be collected. Branch

sections were placed into 1 L deli cups

(Prime Source PS232, Dallas, TX). Un-

treated apple branches were used as con-

trols for each sample date. Five arenas were

prepared for each treatment. Five S. rugulo-

sus adults, collected from emergence cages

described above, were added to an arena

and survival was recorded after 24 h (25

adults/treatment/DAT). It was assumed that

the adults appearing in the vials were newly

emerged, but that could not be verified.

Both males and females were used in the

bioassay, with no effort made to segregate

by sex. We did not generate enough adults

to run the entire screening at one time so

adults were added to the insecticide arenas

in the following order: one replicate from

each treatment followed by an untreated

control replicate for the one DAT samples.

All replicates from this initial collection

date were completed before beginning

evaluations on the next series of samples

(seven DAT). The process was repeated

until all samples were completed. Rearing

conditions were 22 °C, 16:8 L:D. Average

J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

survival and standard error of the means

were reported for each treatment.

Successful Scolytus rugulosus man-

agement practices. During the course of

this study, we documented S. rugulosus

control efforts in four heavily infested or-

chards. In each situation we were contacted

by growers who were already experiencing

severe injury to cherry orchards. We

worked with growers to monitor potential

hosts, whether inside or outside of an or-

chard, with adult traps in an effort to iden-

tify the sources of infestation. Dissections

of suspected host material (removing bark

to expose live larvae) were conducted to

verify S. rugulosus were currently utilizing

the material as a natal host. We also con-

ducted damage evaluations throughout the

orchards to isolate the areas that required

intervention. Once the S. rugulosus situa-

tion was completely described, growers

implemented their own sanitation programs.

We continued to monitor the orchards with

adult traps throughout the clean up process

and subsequently conducted post treatment

damage evaluations to document the effi-

cacy of these efforts. The methods used in

these damage evaluations were consistent

with those described in the trials above.

RESULTS AND DISCUSSION

Bark beetle identification, monitor-

ing, and life history. A total of 17,116

adult Scolytinae were collected from in-

fested fruitwood, yellow sticky traps, and

ethanol-baited intercept traps. The domi-

nant Scolytinae found throughout Washing-

ton was the shothole borer, S. rugulosus

Miiller (ver. Malcom Furniss) (Table 1). An

ambrosia beetle, Xyleborinus saxeseni Ratz-

burg (ver. Malcom Furniss), was present in

high numbers at only one location, a cherry

orchard abandoned for several years. More

than one species of Scolytinae were de-

tected at each location where identification

was not limited to Scolytus spp. A second

Scolytus sp. (S. multistriatus) was found

infesting a pile of cherry wood at only one

site. Cherry has not been reported as a host

(Furniss and Johnson 2002) for S.

multistriatus, and in this case, S. multistria-

tus infested only the pile of cherry wood

and was not detected moving into the

neighbouring cherry orchard. Many other

wood decomposing beetles were reared

from infested fruitwood. In fact, the major-

ity of Coleoptera species collected were

associated with dry, older wood (dead for

more than 18 mo). Buprestid (Buprestidae)

and powderpost beetles (Lyctidae) were the

primary beetles associated with dry wood

(Table 1). S. rugulosus and X. saxeseni

were the primary attackers of weakened

trees or recent cuttings (<18 mo). S. rugulo-

sus was the species most implicated in dam-

age to healthy orchards, whereas X. saxe-

seni was found attacking only trees that had

been previously damaged or weakened.

Initial observations from laboratory emer-

gence cages indicated that there was a high

rate of parasitism (approximately 50%) of

S. rugulosus larvae by Cheiropachus quad-

rum (Hymenoptera: Pteromalidae) (ver.

Malcom Furniss) based on their relative

abundance in vials from emergence cages.

No statistically significant difference

was noted between commercially available

intercept-style traps in their ability to cap-

ture adult S. rugulosus (Chi-Square 3.103,

df 1, P = 0.078) but X. saxeseni captures

were slightly higher in Lindgren Funnel

Trap than Pane Intercept Trap (Chi-Square

4.021, df 1, P = 0.045) (Table 2). Both in-

tercept-style traps should be suitable moni-

toring systems for S. rugulosus and X. saxe-

seni adults. The addition of an ethanol lure

significantly enhanced captures of both S.

rugulosus (Chi-Square 5.333, df 1, P =

0.021) and X. saxeseni (Chi-Square 5.333,

df 1, P = 0.021). Although ethanol lures

significantly increased captures of both

species, this may be an area where monitor-

ing systems could be improved. Synergistic

plant volatiles (Montgomery and Wargo

1983) and/or aggregation pheromones

(Lindgren et al. 1983, Pitman ef al. 1975,

Schroeder and Lindelow 1989, Peacock ef

74 J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

Table 1.

Wood-boring beetle collections from infested fruitwood, yellow sticky traps, and ethanol-

baited intercept traps from Washington, 2003-05.

Location Yr Host Material

Wenatchee 2003 Dead cherry

Wenatchee 2003 <1-yr-old cuttings

Mallot 2003 1-yr-old pushed

over apple

Okanogan 2003 <1l-yr-old pushed

over cherry

Oroville 2003 <1-yr-old cuttings

Oroville 2003 Neglected apple

E. Wenatchee 2003 <2-yr-old cuttings

Wapato 2003 Neglected cherry

W. Valley 2003 Neglected cherry

Cowiche 2003 Neglected cherry

Orondo 2004 <2-yr-old cuttings

Wenatchee 2004 Dead cherry

E. Wenatchee 2004 <2-yr-old cuttings

Okanogan 2004 <l-yr-old cuttings

Bridgeport 2004 <l-yr-old cuttings

Tonasket 2004 <l-yr-old cuttings

Okanogan 2005 <2-yr-old cuttings

Orondo 2005 <2-yr-old cuttings

E. Wenatchee 2005 <2-yr-old cuttings

Moses Lake 2005 Neglected cherry

| __.. Insects not collected for identification.

al. 1972) have been used to improve moni-

toring systems for some bark beetle species,

unfortunately not all species aggregate in

response to pheromone production (Macias-

Samano et al. 1998). It is unclear if S. rugu-

losus or X. saxeseni produce aggregation

pheromones.

Adult S. rugulosus were more highly

attracted to the yellow sticky traps (Chi-

Square 9.143, df 1, P = 0.003), than the

dark coloured ethanol-baited intercept-style

traps. Yellow sticky traps proved to be easy

to deploy and read, and were relatively eco-

nomical compared to the intercept-style

traps. The ethanol-baited intercept-style

traps were necessary to monitor X. saxeseni

Total annual captures in all traps

S. rugulosus X. saxeseni Buprestidae Lyctidae

ie 369 5 6

7 16 27 20

357 a 530 25

8654 4 25 287

695 = = a

26 12 fl 224

247 31 4 357

141 = = is

637 ne - _

284 ae ees =

125 32 a Bet

374 196 = aes

1361 | oe =

218 21 == es

57 6 = eee

6 l aed ae

170 2 = be

1247 = = =

743 = oe a

292 = eat is

761 ae i S22

activity (Chi-Square 12.799, df 1, P =

0.0003), but our experience has been that

this species is a minor contributor to dam-

age in commercial cherry orchards.

Two distinct periods of S. rugulosus

activity occurred in Washington (Fig. 1). S.

rugulosus activity was first noted in late

April or early May and continued through

June. The second adult flight was detected

in mid- to late July and continued through

August and into late-September. Adult S.

rugulosus were trapped through the entire

growing season from initial adult emer-

gence through the end of October. A slight

increase in trap captures occurring at the

end of each season suggested the possibility

J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008 75

Table 2.

Mean (+ SEM) S. rugulosus and X. saxeseni captures using commercially available trap and

lure systems, 2003.

Mean adults/trap (SEM)!

S. rugulosus

Pane Intercept Lindgren Funnel

e 2

X. Saxeseni N

Pane Intercept With Lindgren Funnel With

With Ethanol Lure With Ethanol Lure Ethanol Lure Ethanol Lure 6

68.3 (34.7) 121.3 (59.3) 21.3 (16.4) 31.0 (22.2)*

Lindgren Funnel Lindgren Funnel Lindgren Funnel Lindgren Funnel

With Ethanol Without Lure With Ethanol Without Lure 4

109.1 (66.2)* 59:5:(50:3) 33.9 (29.4)* 7.0 (6.3)

Lindgren Funnel Yellow Sticky Card Lindgren Funnel Yellow Sticky Card

With Ethanol With Ethanol 10

34.9 (27.0) 226.6 (188.6)* 22.7 (14.4)* 0.6 (0.4)

'Means followed by ‘*’ are significantly different (Wilcoxon Rank Test, P = 0.05)

?\N, number of sites in study

Cumulative adult SHB emergence

4/6 4/20 5/4

5/18 6/1 6/15 6/29

7/13

7/27) = 8/10 8/24 9/7 9/21 10/5 10/19

Figure 1. Cumulative emergence of S. rugulosus adults (n=5,675) in Washington, 2003-05.

Open circles represent cumulative emergence from each location, black dots represent the aver-

age emergence on each date. SHB = shothole borer.

of a partial third generation.

Adult X. saxeseni activity occurred

throughout the entire growing season, with

peaks suggesting the presence of three to

four generations (Fig. 2). Adult X. saxeseni

activity was initially noted in late March or

early April. A second peak of activity oc-

curred in early June, with a third noted in

July and early August. A slight increase X.

saxeseni activity was observed in Septem-

ber and early October, although at reduced

numbers. It is unclear if this activity repre-

sented part of a fourth generation or prolon-

gation of the third.

Traps were useful in identifying peak

activity periods of S. rugulosus but it was

not clear if they would be useful in setting

thresholds for treatments. We had trouble

locating S. rugulosus sources with various

population sizes near neighboring cherry

orchards that were allowed to remain un-

treated. Since insecticides were applied

76 J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

100 2°? @ eae Ds e

Cumulative adult AB emergence

3/17) 3/31) 4/14 4/28) «=5/12) 5/26 «6/9 6/23). 7/7 7f21 8/4 8/18 9/19/15

9/29 10/13

Figure 2. Cumulative emergence of X. saxeseni adults (n=805) in Washington, 2003-04. Open

circles represent cumulative emergence from each location, black dots represent the average

emergence on each date. AB = ambrosia beetle.

frequently in cherry orchards for control of

other pests, it was difficult to establish a

consistent relationship between trap cap-

tures and subsequent damage. However, our

observations indicated that if an S. rugulo-

sus host was located near a cherry orchard

and any significant emergence was detected

with yellow traps, some control interven-

tion would be justified to prevent damage.

Scolytus rugulosus migration and

damage distribution. Movement of S. ru-

gulosus into healthy orchards was closely

associated with emergence from a nearby

infested host, generally a pile of recently

pruned or cut wood placed outside the or-

chard (Fig. 3). Cumulative captures of

adults at the outside host and at the border

of the nearby orchard were closely associ-

ated at each study site. There was no con-

sistent pattern of either a lag in percentiles,

or prolonged captures at the orchard border.

In other words, observations at the more

heavily infested outside hosts were repre-

sentative of what was occurring at the or-

chard borders. Further, S. rugulosus activity

was easier to monitor at the host than in

healthy trees (1819 and 258 total S. rugulo-

sus adults, respectively) as a very large

number of adults emerged from a relatively

small area and dispersed immediately.

These data indicated that growers should be

able to focus their efforts at locating and

monitoring suspected outside hosts, under-

standing that as adult activity increased at

the hosts, immigration to healthy orchards

was occurring simultaneously. It appeared

that recently emerged S. rugulosus adults

were highly dispersive and readily moved

distances of at least 50 m from infested

outside hosts to healthy trees in orchard

borders. After dispersal, adult activity in

and around suitable natal hosts continued

where behaviour appeared to be associated

with the construction and care of maternal

galleries. This aspect of S. rugulosus behav-

iour needs to be explored further.

S. rugulosus adult feeding damage to

healthy trees was most commonly associ-

ated with movement from infested hosts.

Generally, S. rugulosus damage was in

close proximity to that host. It appeared that

S. rugulosus moved readily to and along an

orchard border, but did not move more than

two or three rows into a healthy orchard.

On average, 74% of the total S. rugulosus

damage that was detected in healthy or-

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78 J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

chards occurred on the border row closest

to the natal host (Fig. 4). Twenty percent of

the total damage was found on the second

row. Damage in subsequent rows was mi-

nor and scattered. These data indicated that

monitoring and control efforts should be

focused on determining the natal host re-

sponsible for S. rugu/osus infestation, usu-

ally piles of recently cut wood near or-

chards, and then protecting the area of the

orchard immediately adjacent to that host.

If control is neglected, trees will become

weakened and S. rugulosus will success-

fully colonize and reproduce in the weak-

ened trees. Once this occurs immigration 1s

not the sole source of beetles and damage

will spread further into the orchard thereby

complicating management efforts.

Scolytus rugulosus insecticide screen-

ing. Once introduced into the treatment

arenas, adult S. rugulosus began feeding or

attempting to colonize the limb sections

immediately. Although the purpose of this

behaviour was not known (feeding or ovi-

position), the beetles were very active on

the treated wood. The average survival of S.

rugulosus adults on untreated wood was

96% after 24 h. This level of survival indi-

cated any significant mortality could be

attributed to pesticide exposure and not a

problem with the bioassay method.

Many insecticides caused mortality of S.

rugulosus in the bioassays (Table 3). A

pyrethroid, esfenvalerate, was the most

active through 21 d. Azinphos-methyl was

acutely toxic to S. rugulosus, but for only

seven d. Endosulfan and the neonicotinyls,

thiamethoxam and acetamiprid, were some-

what toxic to S. rugulosus. Malathion, in-

doxacarb, and spinosad all caused mortal-

ity, but not at levels expected to provide

adequate control under field conditions.

Additional insecticide efficacy trials are

necessary to understand the full potential of

each insecticide to control S. rugulosus

under field conditions. The repeated use of

these insecticides against other pests, pri-

marily the cherry fruit fly, Rhagoletis indif-

ferens Curran (Diptera: Tephritidae), during

the early part of the growing season is

likely sufficient to suppress damage in most

commercial orchards, especially during the

first S. rugulosus generation. Cherry or-

chards may become more susceptible to

injury in the post-harvest period when in-

secticide programs for cherry fruit fly and

leafroller (Lepidoptera: Tortricidae) have

ceased. Second-generation S. rugulosus

adults would then be able to move into un-

protected orchards.

Successful Scolytus rugulosus man-

agement practices. Sanitation has gener-

ally been touted as the key to control, with

wood or brush piles identified as contribu-

tors of beetles that migrate and attack other

trees (Bhagwandin 1992, Beers et al. 1993,

Payne 1977). In the winter of 2003-04, we

monitored an effort to clean up a large in-

festation of S. rugulosus emerging from an

outside host that had resulted in significant

damage to young, healthy cherry trees. In

2003, approximately 55% of shoots in the

trees along the orchard border at this loca-

tion were damaged despite several insecti-

cide applications, including repeated appli-

cations of endosulfan. The outside host was

a firewood and brush pile that was replen-

ished each year. While the damage was

high, it was fairly well isolated from the

orchard border rows adjacent to the host.

During the winter of 2003-04, the orchard

was pruned heavily, removing all weakened

or damaged branches and the grower made

a concerted effort to clean up all host mate-

rial (firewood, brush piles, and current-yr

cuttings) and maintain a clean area near the

orchard. During the 2004 growing season,

the orchard was monitored with yellow

sticky traps and ethanol-baited intercept-

style traps. Insecticide applications were

planned to coincide with increased trap

captures. However, a total of only four S.

rugulosus and nine X. saxeseni adults were

trapped in five yellow traps and four inter-

cept-style traps and therefore no insecticide

applications specifically for control of bark

beetles were needed. No S. rugulosus dam-

age was noted at any time during the 2004

season demonstrating that the sanitation

efforts were the only control measure

needed.

In 2005, we monitored efforts at three

J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008 79

Mean % of total damage

Row 1 Row 2 Row 3 Row 4 Row 5 Row 6 Row 7

Distance from SHB host

Figure 4. Mean (4SEM) percentage of total damage noted in each orchard (n=4) at the row

adjacent to the source of S. rugulosus (Row 1), and subsequent rows moving into the orchard,

2004-05. SHB = shothole borer.

Table 3.

Mean (4SEM) S. rugu/osus survival on field-aged residues of insecticides, 2004.

ceive Rate gm Mean no. live S. rugulosus (SEM) -— 24h

Insecticide Ingredient —ai/ha 1DAT!' 7DAT 14DAT 21DAT

Asana XL 0.66EC Esfenvalerate 46.8 0.0(0.0) 0.4 (0.2) 0.0(0.0) 0.8 (0.4)

Guthion 50WP — Azinphosmethyl 1,135.0 0.0(0.0) 0.4(0.2) 2.4(0.7) 4.00.3)

Actara25WDG Thiamethoxan 80.0 0.6(0.6) 1.0(0.3) 2.0(0.9) 1.8 (0.7)

Assail 70WP Acetamiprid 168.7 0.6(0.6) 1.4 (0.6) 1.8(0.6) 0.8 (0.4)

Thiodan 3E Endosulfan 259235 0.7 (0.3) 1.3 (0.3) 1.7 (0.3) 1.3 (0.9)

Avaunt 30WDG _ Indoxacarb 127.8 2.0(0.5) 2.0 (0.7) 0.8(0.4) 2.6(1.1)

Malathion 50%EC Malathion 300.0 2.2 (0.8) 2.8 (0.6) 4.2(0.4) 3.8 (0.7)

Success 2SC Spinosad 106.4 3.8 (0.2) 3.6 (0.7) 4.2 (0.4) 3.6 (0.7)

Untreated 5.0(0.0) 4.8 (0.2) 5.0(0.0) 4.4 (0.2)

'DAT, Days After Treatment

locations near Okanogan, WA to clean up _ cherry blocks. These sites were brought to

easily identifiable infested hosts of S. rugu- our attention after first generation beetles

losus located just outside of cherry blocks had caused serious damage to trees in the

exhibiting signs of recent feeding damage. _—_ orchard borders. Insecticidal control options

In addition to the external hosts, weakened were limited as one of the blocks was man-

limbs and one-yr-old cuttings left in the aged organically, and the conventional

orchards were also serving as host material blocks were experiencing damage levels of

for S. rugulosus reproduction within the 50% shoot infestation despite a history of

80 J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

border sprays. Although yellow traps used

to determine what host material was serving

as the source of the S. rugulosus infesta-

tions were placed near the end of first gen-

eration activity, captures in the first seven d

averaged 116 S. rugulosus per trap across

all three locations. Subsequently, the grow-

ers removed all possible host material

within the orchard, including the previous

winter’s cuttings and weakened branches or

limbs. This wood was added to the host

material located outside the orchard and

targeted with an intensive insecticide treat-

ment program. Endosulfan was applied by a

handgun sprayer on a 10-14 d retreatment

interval for the rest of the season with care

taken to thoroughly soak the entire wood

pile. Following this action, no second-

generation beetle activity was noted at any

of the three sites, and no new damage was

detected inside the orchard.

Healthy cherry trees can repel initial

colonization efforts by S. rugulosus adults

by flooding attacked sites with resin, but

with repeated attacks even healthy trees

will eventually become weakened, allowing

successful colonization by secondary at-

tacks from conspecifics (Bauernfeind 1996,

Payne 1977). Our experience with S. rugu-

losus management indicates orchard sanita-

tion is the most important factor contribut-

ing to a reduction in S. rugulosus popula-

tions and damage to healthy cherry trees. If

recent feeding damage is noted on other-

wise healthy trees, adult traps can be placed

on the orchard borders or in suspected host

areas to verify the source of infestation.

Sanitation programs must include removing

potential host material (weakened limbs or

recent cuttings) from within the orchard and

eliminating any host material outside the

orchard. Beetle host material outside the

orchard can be eliminated by burning or by

thoroughly soaking the wood with an effec-

tive insecticide delivered by a handgun

sprayer. The increased volume of water

delivered by handgun applications appears

to be an important factor in insecticide effi-

cacy. We do not believe growers can rely

on traditional insecticide applications via an

air-blast sprayer to control infestations that

originate from within the orchard, or protect

orchard borders from massive immigration

originating from a nearby heavily infested

host.

ACKNOWLEDGEMENTS

We would like to thank Christian

Krupke, Purdue University Department of

Entomology, for reviewing an earlier draft

of this manuscript. This project was funded

by the Washington Tree Fruit Research

Commission.

REFERENCES

Bauernfeind, R. 1996. Shothole Borers. Kansan State Univ. Coop. Ext., MF-2218.

Beers, E.H., J.F. Brunner, M.J. Willett, and G. M. Warner. 1993. Orchard Pest Management: A Resource

Book for the Pacific Northwest. Good Fruit Grower, Yakima, WA. 276 pp.

Berryman, A.A. 1972. Resistance of conifers to invasion by bark beetle fungus associations. BioScience.

22: 598-602.

Bhagwandin, H.O. 1992. The Shothole borer: An ambrosia beetle of concern for chestnut orcharding in the

Pacific Northwest, p. 168-177. Jn 93" Annual Report of the Northern Nut Growers’ Assn., Western

Chestnut Growers Assn.

Brunner, J.F., J. Dunley, W. E. Jones, E. H. Beers, G. V. Tangren, C Xiao, and G. G. Grove. 2003. Pesticide

use and IPM practices in Washington's pear and cherry orchards. Agriculture and Environmental News.

No 208. http://aenews.wsu.edu/Aug03 AENews/Aug03AENews.htm

Furniss, M.M., and J.B. Johnson. 2002. Field Guide to the Bark Beetles of Idaho and the Adjacent Re-

gions. Idaho Forest, Wildlife and Range Experiment Station, Univ. of Idaho. Station Bulletin 74.

JMP statistics and graphics guide, version 5.1.2. 2004. ed by SAS Institute, Cary, North Carolina.

Kirk, V.M. 1969. A list of beetles of South Carolina. Part 1 — Northern Coastal Plains Technical Bulletin

1033. South Carolina Agricultural Experiment Station, Clemson University. 124 pp.

J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008 81

Kovach, J., and C.S. Gorsuch. 1985. Survey of ambrosia beetle species infesting South Carolina peach or-

chards and a taxonomic key for the most common species. Journal of Agricultural Entomology. 2(3):

238-247.

Lindeman, G.V. 1978. Means of adaptation of bark beetles (Scolytus kirschi, Scolytus japonicus, Scolytus

rugulosus, Scolytus moravitzi) to a habitat on slightly weakened trees. The Soviet Journal of Ecology. 9

(6): 538-543.

Lindgren, B.S., J.H. Borden, L. Chong, L.M. Friskie, and D.B. Orr. 1983. Factors influencing the efficiency

of pheromone-baited traps for three species of ambrosia beetles (Coleoptera: Scolytidae). The Canadian

Entomologist. 115: 303-313.

Linsley, E.G., and G.F. MacLeod. 1942. Ambrosia beetles attacking deciduous fruit trees in California.

Journal of Economic Entomology. 35 (4): 601.

Macias-Samano, J.E., J.H. Borden, R. Gries, H.D. Pierce, and G. Gries. 1998. Lack of evidence for phero-

mone-mediated secondary attraction in the fir engraver, Scolytus ventralis (Coleoptera: Scolytidae). Jour-

nal of the Entomological Society of British Columbia. 95: 117-125

Markalas, S., and M. Kalapanida. 1997. Flight pattern of some Scolytidae attracted to flight barrier traps

baited with ethanol in an oak forest in Greece. Journal of Pest Science. 70(3): 55-57.

Mendel, Z., S. Ben- Yehuda, R. Marcus, and D. Nestel. 1987. Distribution and extent of damage by Scolytus

spp. to stone and pome fruit orchards in Israel. Insect Science and its Application. 17(2): 175-181

Montgomery, M.E., and P.M. Wargo. 1983. Ethanol and other host-derived volatiles as attractants to bee-

tles that bore into hardwoods. Journal of Chemical Ecology. 9:181-190.

Payne, P.B. 1977. Shothole borer (Scolytus rugulosus). Leaflet / Division of Agricultural Science, Univer-

sity of California Berkeley, Cooperative Extension. Volume 2943, pp 1-2.

Peacock, J.W., R.M. Silverstein, A.C. Lincoln, and J.B. Simeone. 1972. Laboratory investigations of the

frass of Scolytus multistriatus (Coleoptera: Scolytidae) as a source of pheromone. Journal of Environ-

mental Entomology. 2(3): 355-359.

Pitman, G.B., R.L. Hedden, and R.I. Gara. 1975. Synergistic effects of ethyl alcohol on the aggregation of

Dendroctonus pseudotsugae (Coleoptera: Scolytidae) in response to pheromones. Zeitschrift fiir ange-

wandte Entomologie. 78:203-208.

Rudinsky, J.A. 1962. Ecology of Scolytidae. Annual Review of Entomology. 7: 327-348.

Schroeder, L.M. and A, Lindelow. 1989. Attraction of scolytids and associated beetles by absolute amounts

and proportions of alpha-pinene and ethanol. Journal of Chemical Ecology. 15:807-817.

Smith, L.M. 1932. The shot-hole borer. University of California, Agricultural Extension Service. Circula-

tion 64.

Smith, T.J., J.-E. Dunley, E.H. Beers, J.F. Brunner, G.G. Grove, C.L. Xiao, D. Elfving, F.J. Peryea, R.

Parker, M. Bush, C. Daniels, T. Maxwell, S. Foss, and S. Roberts. 2004. Crop protection guide for tree

fruits in Washington. Washington State University, Cooperative Extension. Bulletin EB 0419.

Warner, G.M. 2006. Pest control important in market access. Good Fruit Grower. 57(6): 30-31.

Wilcoxon, F. 1945. Individual comparisons by ranking methods. Biometrics. 1: 80-83.

J. ENTOMOL. Soc. BRIT. COLUMBIA 105, DECEMBER 2008

J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

Extracts of Ginkgo biloba or Artemisia species reduce feeding

by neonates of codling moth, Cydia pomonella (Lepidoptera:

Tortricidae), on apple in a laboratory bioassay

KEVIN DURDEN’, JOHN J. BROWN?

and MACIEJ A. PSZCZOLKOWSKI'”

ABSTRACT

In a simple bioassay, alcohol extracts (91% v/v ethanol, 4% v/v methanol, 5% v/v iso-

propanol) from Ginkgo biloba L., Artemisia absinthium L., A. arborescens L. x A. ab-

sinthium L., and A. ludoviciana Nutt. “Valerie Finnis”, significantly reduced feeding of

neonatal codling moth larvae on apple. Extracts from A. californica Less. and A. vul-

garis L. had no effects.

Key Words: insect feeding, larvae, repellent, deterrent

INTRODUCTION

Codling moth, Cydia pomonella (L.), is

a cosmopolitan pest that primarily attacks

apple, Malus domestica Borkh., a commod-

ity worth more than US$1 billion per year

in the northwestern United States and Brit-

ish Columbia, Canada. Codling moth fe-

males oviposit mostly on foliage (Jackson

1979). Newly hatched neonates travel over

apple branches and foliage in search of the

fruit (Jackson 1982) and finally burrow into

it (Tadic 1957). Manipulation of neonate

searching behaviour may provide an alter-

native or complementary approach to cur-

rent strategies for codling moth manage-

ment (Pszczolkowski 2007).

Neonate codling moth responses to

feeding stimulants have previously been

explored (Pszczolkowski 2007). Suomi et

al. (1986) and Landolt et al. (1999) studied

deterrent effects of, respectively, plant ex-

tracts and plant essential oils. Of the 25

species tested by Suomi ef al. (1986), five

showed promise as feeding deterrents to

neonatal codling moth larvae: absinthe

wormwood, Artemisia absynthium L.; rab-

bitbrush, Chrysothamnus nauseosus (Pall.

ex Pursh) Britton; false hellebore, Veratrum

californicum Durand; garlic, Alium sativum

L.; and tansy, Tanacetum vulgare L. Of the

27 species tested by Landolt et al. (1999),

the greatest arrestment of neonates was

achieved with oils of lavender, Lavandula

officinalis Chaix ex Vill; pennyroyal, Men-

tha pulegium L.; and cypress, Cupressus

sempervirens L. Oijls of rue, Ruta

graveolens L.; garlic; patchouly, Pogoste-

mon cablin (Blanco) Benth.; and tansy were

the most repellent to neonates (Landolt er

al. 1999). Extracts from the ginkgo tree,

Ginkgo biloba L., which have deleterious

effects on codling moth neonates

(Pszczolkowski and Brown 2005), were not

studied by Suomi ef a/. (1986) or Landolt e¢

al. (1999).

In this paper, we use a simple modifica-

tion of the assay designed by Suomi et al.

(1986) to test the effects of extracts of G.

biloba and of five members of the genus

Artemisia on feeding by codling moth neo-

nates. Our assay allows small volumes of

plant extracts on apple plugs to be pre-

sented to individual neonates. We compare

‘Department of Agriculture, Missouri State University, 9740 Red Spring Road Mountain Grove, MO

65711, USA

? Department of Entomology, Washington State University, FSHN 166, Pullman, WA 99164, USA

> Corresponding author. Tel. 417-547-7507. E-mail: MPszczolkowski@missouristate.edu

84 J. ENTOMOL. Soc. BRIT. COLUMBIA 105, DECEMBER 2008

neonates’ response to gingko extract in the

apple plug assay with their response to

whole apples using an assay previously

described in Pszczolkowski and Brown

(2005).

MATERIALS AND METHODS

Insects. Codling moths originated from

USDA-ARS Yakima Agricultural Research

Laboratory in Wapato, WA, the same

source as used by Suomi ef al. (1986). This

laboratory has maintained the codling moth

colony for about 40 years, comprising more

than 480 generations. Moths were held at

25 °C, 70-80% RH, under a 16L:8D light-

dark regime, and allowed to oviposit on

polypropylene foil. Neonates were collected

0.5-1 h post-hatch, and used for experi-

ments.

Plant extracts. The extracts were pre-

pared from foliage of G. biloba, A. absin-

thium, Artemisia arborescens L. x Ar-

temisia absinthium L., Artemisia ludovici-

ana Nutt. “Valerie Finnis, Artemisia cali-

fornica Less. and Artemisia vulgaris L.

Dried A.absynthium foliage was pur-

chased from a local pharmacy (London

Apothecary, Mansfield, MO). Remaining

plant material was collected in the gardens

of Missouri State University Research

Campus, Mountain Grove, MO, in July

2008.

Dehydration alcohol (91% v/v ethanol,

4% v/v methanol, 5% v/v _ isopropanol;

EMD Chemicals Inc., Gibbstown, NJ) was

used to prepare all extracts. Plants were

dehydrated using an Open Country food

dehydrator (Nesco/American Harvest®,

Two Rivers, WI) at 35 °C for 48 hours. The

dry foliage was ground in a coffee grinder.

Approximately 0.5 ml of dry plant powder

was placed in a plastic centrifugation tube,

500 ul of dehydration alcohol added, then

the tube was vortexed and left at room tem-

perature for 10 min. The tube was then cen-

trifuged at 2000 G for 10 min and 300 ul of

liquid fraction was transferred to a pre-

weighed plastic test tube and allowed to air

dry overnight. The test tube with the resi-

due was re-weighed the next morning and

enough dehydration alcohol was added to

make a 10 mg/ml solution of each plant

extract. The extracts were prepared imme-

diately before testing.

Modified assay using apple plugs. For

each test arena, four plugs were procured

from the same Golden Delicious apple,

using a length of plastic soda straw (Fig.

1A), such that the straw covered the pulp,

but not the epidermis of the apple. The

crevice between the plug and the edge of

the straw was sealed with paraffin wax ap-

plied with a warm spatula (Fig. 1B). The

straws were then placed in a holder, apple

plug facing up, and 5 ul of test solution

were applied to each plug. The plugs were

allowed to air dry, and four plugs were

placed in a 60 x 15 mm polystyrene Petri

dish (Fig. 1C). Small pieces of modeling

clay held the plugs in place. New clay was

used for each assay. A glass rod (1.3 mm

diameter, 25-27 mm long) was positioned

such that each end of the rod touched both

the control and the treated member of the

plug pair (Fig. 1C). One neonate was

placed, using a camel-hair artistic brush, in

the middle of the glass rod and the Petri

dish was covered with a lid. The entire as-

sembly was covered with a half of a white

plastic RipBall (TM & Enor Corp., North-

vale, NJ) to provide a white, slightly

opaque cupola (Fig. 1D) and placed on the

testing bench illuminated by fluorescent

tubes and Soft White 60 general purpose

bulbs (General Electric Canada, Cleveland,

OH). Such an arrangement provided dis-

persed, non-directional light of uniform

luminosity (900-920 lux) over each test

arena, which was important because cod-

ling moth neonates exhibit mild phototro-

pism (Jackson, 1982). Prior to every experi-

ment, glass rods and Petri dishes were

washed sequentially in tap water, double

distilled water, alcohol, then dried.

Preliminary experiments showed that

neonates could be expected to locate a plug

and begin feeding within 20 hours, and that

J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008 85

monaannncnnnnaniin va ‘eee

out of apple. B. Crevice between the plug and the straw is sealed with paraffin wax. C. a, c.

Plugs treated with plant extracts; b, d. control plugs treated with dehydration alcohol. D. Test

arena is covered with plastic cupola to disperse light and eliminate possible effects of direc-

tional lighting.

neonates showed no preference for any plug

position in the arena. In the experiments

reported here, 30 neonates were individu-

ally exposed to ginkgo extracts, and 18-23

neonates were individually exposed to ex-

tracts of each Artemisia species. All neo-

nates were tested simultaneously. All apple

plugs were examined under a dissecting

microscope after 20 h for feeding indicators

such as abraded epidermis, presence of ex-

crement, or a feeding cavity. If evidence of

feeding was found, the plug was removed

from the straw and dissected to reveal the

larva.

Whole-fruit assay. To test whether or

not neonates behaved similarly when pre-

sented with ginkgo extracts on whole fruit,

we compared neonate behaviour in the ap-

ple-plug assay with behaviour in our previ-

ously described whole-fruit assay

(Pszczolkowski and Brown, 2005). Unin-

fested thinning apples (Red Delicious;

about 20 mm diameter) from Mountain

Grove experiment orchards were used for

whole-fruit assays. Apples were submerged

in 10 mg/ml extract of ginkgo in dehydra-

tion alcohol or in dehydration alcohol only

(about 200 ul of test solution per apple),

86 J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

and allowed to air dry. Two apples (one

ginkgo-treated and one alcohol-treated con-

trol) were placed 0.5 cm apart in a 70-mm

diameter Pyrex glass crystallizing dish. One

neonate was gently placed with a fine

camel-hair artistic brush in the space be-

tween the fruits and the crystallizing dish

was covered with a glass Petri dish. To pre-

vent airflow that could bias the results of

the assay, the entire assembly was placed in

a semi-translucent 473-ml high-density

polypropylene container and covered with a

transparent lid. The testing bench and test

arenas were illuminated as described for the

apple-plug assay. Before each test, glass-

ware and polypropylene containers were

washed sequentially in tap water, double

distilled water, alcohol, then dried.

Thirty neonates were tested individually

in this assay. Both apple-plug assay and

whole-fruit assay were conducted at the

same time After 24 h, all apples were ex-

amined under a dissecting microscope for

evidence of feeding, as described above for

the apple-plug assay.

Statistical analysis. Exact Fisher’s test

(a=0.05) was used in all assays to test the

null hypothesis that neonates do not dis-

criminate between plugs or apples treated

with plant extract and those treated with

alcohol (i.e., 50% of the neonates choose

treated plugs or apples and 50% of the neo-

nates choose control plugs or apples).

RESULTS

Effects of ginkgo in apple-plug assay

and whole-fruit assay. In both the apple-

plug and the whole-fruit assays, the major-

ity (29 of 30, and 28 of 30, respectively) of

neonates avoided fruit treated with 10 mg/

ml of ginkgo extract (P< 0.001). For every

neonate, feeding indicators such as abraded

epidermis, presence of excrement, or feed-

ing cavities were found on only one apple

plug or one apple out of two members of

one pair. We conclude that this is evidence

that each neonate larva, upon arrival at a

ginkgo-treated plug or fruit, did not attempt

to feed and was deterred or repelled, or

both, by ginkgo extract.

Effects of Artemisia extracts in apple-

plug assay. Extracts from three of the five

Artemisia species discouraged neonates

from burrowing into apple plugs (Table 1).

Extracts from A. absinthium, A. arbores-

cens x A. absinthium and A. ludoviciana

“Valerie Finnis” were active (P<0.01). Ar-

temisia vulgaris and A. californica had no

effect. As in the case of ginkgo, for every

neonate, feeding indicators were found on

only one apple plug out of two members of

one pair. We conclude that Artemisia ex-

tracts were either repellent or deterrent or

both.

DISCUSSION

Deterrent or repellent activity of ginkgo

extracts toward codling moth neonates is a

novel finding. Surprisingly, information

about insect deterrent activity of this plant

is scarce in the literature, but what exists

provides indirect supportive evidence. Two

studies showed that extracts from ginkgo

foliage reduce feeding by two insect pests

of cabbage: Pieris brassicae (Fu-Shun et al.

1990) and P. rapae (Matsumoto and Sei

1987). Addition of anacardic acids (an al-

kylphenol found in ginkgo) reduced intake

of artificial diet in Colorado potato beetle,

Leptinotarsa decemlineata (Schultz et al.

2006). Other biologically active compo-

nents of ginkgo foliage include flavonoids

and there is evidence that some flavonoids

have deterrent and antifeedant activity in

insects (Simmonds 2001). At the current

stage of our study, we do not know what

constituents of ginkgo extract discourage

codling moth larvae from burrowing into

apple plugs.

The finding that A. absinthium deters

codling moth neonates corroborates the

results of Suomi ef al. (1986). In their ex-

periments, only 9% of larvae bored into

apple plugs treated with a 1% extract ob-

J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

Table 1.

Effect of 10 mg/ml extracts obtained from plants in the genus Arthemisia on feeding by cod-

ling moth neonates.

Plant species used for apple plug Number of Number of larvae feeding

Artemisia absinthium pA Pius

Artemisia arborescens x A. absinthium 16 aad

Artemisia ludoviciana “Valerie Finnis” 17 sta

Artemisia vulgaris 16 4

Artemisia californica 18 71

| ** P<().01, Fisher’s exact test

?* no statistical significance

tained from this plant. Our results showing

that extracts from other members of Ar-

temisia genus also have deterrent properties

against codling moth neonates are novel,

but not surprising in the light of other data

from the literature. For instance, the essen-

tial oil of A. annua has repellent activities

against two economically important stored-

product pests, the red flour beetle Tribolium

castaneum (Herbst) and the cowpea weevil

Callosobruchus maculatus (Tripathi et al.

2000). The compound 1,8-cineole isolated

from the same plant has feeding deterrent

activity against 7. castaneum (Tripathi et

al. 2001). Essential oils from A. vulgaris

also repel T. castaneum beetles (Wang et al.

2006). The fact that different plants from

the same genus have different biological

activity against codling moth neonates may

facilitate isolation of their active compo-

nents by comparative chemical analysis.

We think that our findings warrant fur-

ther studies on effects of ginkgo and Ar-

temisia extracts on codling moth neonates.

Active constituents of these extracts should

be identified, and their potential as codling

moth feeding deterrents or repellents — as-

sessed. Perhaps, if manufactured on a larger

scale, those constituents could be used as

organic alternatives to conventional insecti-

cides for management of codling moth on

apples.

ACKNOWLEDGEMENTS

We thank Ms. Pam Anderson and Mr.

Jim Harris, USDA, ARS, Wapato WA,

USA for provision of insects for this study.

Technical assistance of Miss Samantha

Sellars is greatly acknowledged. A part of

this project was funded by Missouri Life

Science Research Board, grant No. F0741

162001.

REFERENCES

Fu-Shun, Y., K.A. Evans, L.H. Stevens, T.A. van Beek and L.M. Schoonhoven. 1990. Deterrents extracted

from the leaves of Ginkgo biloba: effects on feeding and contact chemoreceptors. Entomologia Experi-

mentalis et Applicata 54: 57-64.

Jackson, M.D. 1979. Codling moth egg distribution on unmanaged apple trees. Annals of the Entomological

Society of America 72: 361-368.

Jackson, M.D. 1982. Searching behavior and survival of 1*-instar codling moths. Annals of the Entomologi-

cal Society of America 75: 284-289.

Landolt, P.J., R.W. Hofstetter, and L.L. Biddick. 1999. Plant essential oils as arrestants and repellents for

neonate larvae of the codling moth (Lepidoptera: Tortricidae). Environmental Entomology 28: 954-960.

Matsumoto, T. and T. Sei. 1987. Antifeedant activities of Ginkgo biloba L. components against the larva of

Pieris rapae crucivora. Agricultural and Biological Chemistry 51: 249-250.

Pszczolkowski, M.A. 2007. How to trick codling moth? Prospects of manipulation with neonate larvae

88 J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

behavior to the benefit of the grower. Annals of Warsaw University of Life Sc. - SGGW, Horticulture

And Landscape Architecture 28: 7-18.

Pszczolkowski M.A. and J.J. Brown. 2005. Single experience learning in host fruit selection by lepidopteran

larvae. Physiology and Behavior 86: 168-175.

Schultz D.J., C. Olsen, G.A. Cobbs, N.J. Stolowich, and M.M. Parrott. 2006. Bioactivity of anacardic acid

against colorado potato beetle (Leptinotarsa decemlineata) \arvae. Journal of Agricultural and Food

Chemistry 54: 7522-7529.

Simmonds, M.S.J. 2001. Importance of flavonoids in insect-plant interactions: feeding and oviposition.

Phytochemistry 56: 245-252.

Suomi, D., J.J. Brown and R.D. Akre. 1986. Responses to plant extracts of neonatal codling moth larvae,

Cydia pomonella (L.), (Lepidoptera: Tortricidae: Olethreutinae). Journal of Entomological Society of

British Columbia 83: 12-18.

Tadic, M. 1957. Jabucni Smotavac (Carpocapsa pomonella L.). Biologija kao osnova za njegovo suzbi-

janije. Univerzitet u Beogradu, Belgrade, Yugoslavia. 100 pp.

Tripathi, A.K., V. Prajapati, K.K. Aggarwal, S.P.S. Khanuja and S. Kumar. 2000. Repellency and toxicity

of oil from Artemisia annua to certain stored-product beetles. Journal of Economic Entomology 93: 43-

47.

Tripathi, A.K., V. Prajapati, K.K. Aggarwal and S. Kumar. 2001. Toxicity, feeding deterrence, and effect of

activity of 1,8-cineole from Artemisia annua on progeny production of Tribolium castaneum (Coleoptera:

Tenebrionidae). Journal of Economic Entomology 94: 979-983.

Wang, J., F. Zhu, X.M. Zhou, C.Y. Niu and C.L. Lei. 2006. Repellent and fumigant activity of essential oil

from Artemisia vulgaris to Tribolium castaneum (Herbst) (Coleoptera: Tenebrionidae). Journal of Stored

Product Research 42: 339-347.

J. ENTOMOL. Soc. BRIT. COLUMBIA 105, DECEMBER 2008

Management of dandelion to supplement control of western

flower thrips (Thysanoptera: Thripidae) in apple orchards

S.D. COCKFIELD’” and E.H. BEERS!

ABSTRACT

We evaluated whether management of the broadleaf weed dandelion, Taraxacum offici-

nale F.H. Wigg. ager., affected damage to apples by western flower thrips (Frankliniella

occidentalis (Pergande)). Four commercial apple orchard blocks in central Washington

having high densities of dandelion were monitored over a 3-yr period. Herbicides were

applied to the drive rows in one-half of each orchard for each year of the study. A 92%

reduction in dandelion densities in the low-weed plots was achieved by the third year of

the experiment. The number of thrips per dandelion plant did not change as dandelions

became less numerous. This resulted in an overall reduction in western flower thrips per

unit area on dandelions throughout the course of the trial. However, the number of west-

ern flower thrips in the apple flowers and shoots were not affected by the treatment.

Estimated western flower thrips population density per ha on apple and dandelion indi-

cated that dandelions harboured a much smaller pool of western flower thrips in com-

parison to apple. No significant reduction in fruit injury was detected in any year. Thus,

reduction or elimination of dandelion from the orchard floor appears to be of limited

value in managing western flower thrips in apple orchards.

INTRODUCTION

Western flower thrips, Frankliniella

occidentalis (Pergande), is a sporadic, di-

rect pest of apple (Beers ef al. 1993). The

most conspicuous injury consists of an ovi-

position puncture, which leaves a small,

rugose scar, and a series of white spots sur-

rounding it, commonly known as pansy

spot (Newcomer 1921). Injury is most ap-

parent on green or blush cultivars (Madsen

and Jack 1966). Densities of adult thrips in

apple blossom clusters increase in late April

and May as the flowers open; oviposition in

fruit occurs from the end of bloom until

fruit reaches 30 mm in diameter (Cockfield

et al. 2007a).

Beginning with the earliest investiga-

tions of western flower thrips in apple or-

chards, broadleaf plants in the groundcover

have been assumed to greatly influence the

population dynamics of this pest. Venables

(1925) collected western flower thrips from

tumbling mustard (Sisymbrium altissimum

L.), a common weed, and alfalfa, Medicago

sativa L., from apple orchards in British

Columbia. These host plants harboured the

pest throughout the summer. Childs (1927)

stated that the choice and management of

cover crops is a major component to west-

ern flower thrips management. He recom-

mended cultivating or plowing the orchard

ground in early spring to eliminate weeds

and to disrupt overwintering thrips in the

soil. After the practice of growing cover

crops such as alfalfa was no longer com-

mon, Madsen and Jack (1966) determined

that dandelion, Taraxacum officinale F.H.

Wigg. aggr., was the most abundant under-

story host of western flower thrips before

and after apple bloom, and wild mustard,

Brassica kaber (DC.) L.C. Wheeler, and

asparagus, Asparagus officinalis L., were

important alternate hosts in the summer.

Pearsall and Myers (2000, 2001) found that

not only did a number of broadleaf weeds

'Tree Fruit Research and Extension Center, Washington State University, 1100 N. Western Avenue, We-

natchee, WA 98801

* Corresponding author: PO Box 1461, Malott, WA 98829. E-mail: pest@bossig.com

90 J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

harbour western flower thrips, but orchards

with the highest population of weeds had

some of the highest numbers of western

flower thrips caught in sticky traps.

Two recent experiments done on

groundcover management have shown

mixed results. Hubscher (1983) found that a

temporary perturbation, such as mowing

dandelions in spring (intended to redirect

pollinators to apple blossoms) had no meas-

urable effect on the population of western

flower thrips in apple, or damage to fruit.

Cossentine ef al. (1999) showed that elimi-

nation of weeds such as dandelion in small

plots reduced thrips populations in trees,

but the effect was temporary. The magni-

tude of the effect may have been limited by

the temporal and spatial scale of both stud-

ies; thus, a more substantial, permanent

effect may result from eliminating weeds

for a number of seasons in larger orchard

blocks. If successful, this method could

reduce the need to control western flower

thrips around the bloom period, or reduce

fruit damage as part of a multi-tactic IPM

program. The purpose of this investigation

was to determine the effect of a cultural

control method, long-term management of

dandelion, on populations of western flower

thrips in large blocks of commercial apple

orchards and to determine the effect of

thrips population changes on fruit damage.

MATERIALS AND METHODS

Site Description. This experiment was

conducted over the course of three years in

four commercial apple orchards near the

towns of Bridgeport, Brewster, Pateros, and

Quincy in Washington State, U.S.A. The

orchards were selected because of a history

of significant thrips damage to fruit and for

their high densities of dandelions (>10/m’).

Trees in the Bridgeport orchard block were

cv. ‘Granny Smith’, while the Brewster and

Pateros blocks were cv. ‘Braeburn’. The

Quincy block was planted with alternating

two rows of cv. ‘Fuji’ and ‘Braeburn’.

Trees were 7-15 yr old. Each block was

approximately 2-4 ha and was surrounded

by other orchards. There were two treat-

ments: 1) low-weed, where herbicide appli-

cations were made in the drive rows (the

section of the orchard floor between the tree

rows left as mixed grass and weed ground-

cover to facilitate use of equipment) and 2)

high-weed (drive rows untreated with herbi-

cides). Each of the four orchards was di-

vided in half, and treatments assigned ran-

domly, with each site serving as a replicate.

Herbicides. A _ vegetation-free strip

about 2 m wide was maintained beneath the

trees in both halves of the block. All or-

chards in the study had a similar program

for weed control in the vegetation-free tree

rows. Pre-emergent herbicides, and some-

times 2,4-dichlorophenoxyacetic acid (2,4-

D), were applied in early spring, followed

by individual sprays or mixtures of

paraquat, 2,4-D, and glyphosate after

bloom. If necessary, repeat applications

were made in June.

The drive rows of low-weed plots were

sprayed with 1.1-1.6 kg Al/ha of 2,4-D

(Weedar 64, Nufarm Inc., St. Joseph, MO)

+ 370 ml/100 litres of R11 surfactant

(Wilbur-Ellis Co., San Francisco, CA). One

or two applications were made in the spring

with a weed sprayer calibrated to deliver

234 litres/ha. Any surviving weeds were

treated individually with a 9% vol:vol solu-

tion of 2, 4-D with a 15.1 litre backpack

sprayer (Wil-Gro, Wilbur-Ellis Co., San

Francisco, CA).

Insecticides. Insecticides applied for

apple pests in a given site were the same

across the entire block (high-weed and low-

weed plots). Densities of lepidopteran pests

were generally low in the study orchards,

and required minimal treatment. These

pests were managed with applications of

chlorpyrifos at delayed dormant, spinosad

at petal fall, and azinphos-methyl or meth-

oxyfenozide during early summer. Because

of the history of fruit injury, growers ap-

plied formetanate hydrochloride to control

western flower thrips at full bloom every

year.

Sampling Methods. Dandelion densi-

J. ENTOMOL. Soc. BRIT. COLUMBIA 105, DECEMBER 2008

ties were sampled by counting the numbers

of plants in a marked 1 m’ area in the drive

rows. Ten randomly selected areas were

marked in the middle row of each treatment

block, with each area separated by about 3-

5 m. Within these areas, dandelion plants

(both total numbers and those in flower)

were counted monthly from March or April

through October.

Thrips densities were sampled monthly

on four flowering and four vegetative dan-

delion plants per plot. All thrips samples

were taken the same time of mid-day in

each of the paired plots, although the time

differed between sites. Dandelion plants

(outside the m” marked areas) were severed

at ground level and placed in self-sealing

plastic bags.

Thrips were sampled in the blossoms

and vegetative shoots of apple trees. Blos-

som samples were taken during full bloom

(April or May) about one day before insec-

ticides were applied. Twenty-five open ap-

ple blossoms were selected from eight trees

randomly chosen in the middle row of the

plot. The blossoms were clipped off and

placed in self-sealing plastic bags. The area

sampled was at least 30 m away from the

edges of the plots. In the third and final

year, the sample size was increased to 150-

300 flowers per tree. Thrips in vegetative

shoots were sampled monthly after the ter-

mination of bloom. The tips (3 cm in

length) of 10 growing shoots were collected

from each of eight trees.

Thrips samples from dandelion and ap-

ple were stored under refrigeration and

processed the day after collection. Thrips

were separated from plant material by fill-

ing the bag with water, adding a few drops

of liquid detergent, and agitating for several

minutes. Thrips and plant material were

separated from the soapy water by pouring

through two sieves (Hubbard Scientific Co.,

Northbrook, IL). The larger sieve (#10,

0.25-mm mesh) trapped most of the plant

material, and the finer sieve (#230, 0.0014-

mm mesh) trapped the thrips. Thrips were

then rinsed into a vial of 50% ethanol.

Specimens were first examined under a

dissecting microscope. All Frankliniella

adults were slide-mounted in PVA Mount-

ing Medium (BioQuip Products, Inc., Ran-

cho Dominguez, CA) for identification to

species. A reference collection was sent to

Cheryl O’Donnell, Department of Entomol-

ogy, University of California, Davis, CA,

who confirmed the identity of species. Only

numbers of F. occidentalis adults were re-

corded. The means of the samples taken per

plot were used in the analyses. All data

were expressed as thrips per dandelion

plant, apple flower, or apple shoot. Thrips

per dandelion plant during peak dandelion

flowering were averaged over the three

years of the study.

Population Estimates. The monthly

population density of western flower thrips

per m° in dandelions in the drive rows was

estimated by multiplying the average num-

ber of thrips per sampled nonflowering and

flowering dandelions by the average num-

ber of dandelions of each type per m’. Esti-

mates were also summed for each replicate

over the three years of the study. The esti-

mate at peak dandelion bloom, just before

or during apple bloom, was then used to

calculate the number of western flower

thrips on dandelion on a per-hectare basis.

This was done by multiplying the mean

number of thrips per m* by the number of

m’ of drive rows in each ha. Similarly, esti-

mates were obtained for western flower

thrips per ha in apple flowers. Flower den-

sities were estimated by counting all blos-

soms on 10 trees per plot. Mean flowers per

tree were calculated for each site, then con-

verted to means per ha based on the tree

density per ha. Flowers per ha were multi-

plied by the mean number of thrips per

flower.

Data Analysis. The experimental design

was a randomized complete block with the

four orchards serving as replicates. The

effects of herbicide treatment and month on

dandelion densities were assessed using a

two-way analysis of variance having re-

peated observations through time (=month).

The analyses were done in SAS _ using

PROC MIXED (SAS Institute 2002). Sepa-

rate analyses were done for each of the

three years of the study. We used a square

92 J. ENTOMOL. Soc. BRIT. COLUMBIA 105, DECEMBER 2008

root transformation on the count data before

each analysis to meet ANOVA assump-

tions. In the event of a significant treatment

x month interaction, tests on simple effects

of treatment and month were done using the

SLICE command. Analyses of flowering

dandelions were done only for the sampling

date with the highest number of flowering

plants. Comparisons of flowering dandeli-

ons were made by year using paired t-tests

(SAS Institute 2002). Data for thrips densi-

ties, whether monthly, averaged, or

summed measurements, were analyzed by

date using ANOVA. Treatment means were

separated using a LSD test (high-weed vs

low-weed, a=0.05). Data for proportion (p)

of fruit with pansy spot injury were first

transformed by arcsine(square root

(p+0.001)), then analyzed using ANOVA

(SAS Institute 2002).

RESULTS AND DISCUSSION

Dandelion Densities. Densities of total

dandelions were high in both high-weed

and low-weed treatments at the start of the

experiment (Fig. 1A). A few other broad-

leaf weeds also occurred in the orchard

blocks, most notably alfalfa, but population

densities of these other weeds were zero in

the randomly-sampled areas. All perennial

and annual broadleaf weeds were affected

by the herbicides and were reduced in the

low-weed plots, which resembled mani-

cured lawns. The treatment < month inter-

action terms were highly significant in 2003

(F=4.10; df=7, 42; P=0.0016) and 2004

(F=3.39; df=8, 48; P=0.0037), and signifi-

cant in 2005 (F=2.37; df7, 42; P=0.039),

thus we examined the simple effects tests

(herbicide effect for each month sepa-

rately). Treatment differences became sig-

nificant by July of 2003, with 17.1 (4 11.6

SEM) dandelions/m’ in the low-weed plots

versus 66.1 (+ 23.4) in the high-weed plots.

These differences were sustained through

the remainder of the year. Treatment differ-

ences were only marginally significant for

the first two months of 2004 (P<0.10), but

were re-established by May with lower dan-

delion densities in the low-weed plots

throughout the remainder of the year

(P<0.001). Treatment differences were sig-

nificant throughout 2005 (P<0.001), which

was also reflected in the main effect treat-

ment means for this year (low-weed plots,

2.3 + 0.3; high-weed plots, 29.4 + 2.4 dan-

delions/m*; F=75.79; df=1, 6; P=0.0001).

The treatment differences in 2005 reflected

a 92% reduction in dandelion densities in

the low-weed plots relative to the high-

weed plots.

Dandelions flowered primarily in April

or May; very few plants flowered in the

summer and fall (Fig. 1B). During the peak

period, flowering dandelion density was not

significantly different in the low-weed (5.5

+ 1.6) and high-weed (7.5 + 3.4) plots in

2003 (t=0.60, df=3, P=0.59). The effect of

the herbicide applications was more clearly

seen in 2004 (low weed, 2.8 + 1.8; high

weed, 10.1 + 3.1; 3.74, df=3, P=0.03) and

2005 (low weed, 0.3 + 0.1; high weed, 8.0

+ 1.8; 4.36, df3, P=0.02).

Thrips in Dandelions. The estimated

population density of thrips on dandelion in

the drive rows (on a per m” basis) generally

increased in the high-weed blocks in late

spring, and peaked in April (2005), during

which maximum dandelion bloom occured,

or June (2003, 2004) (Fig. 1C). Adult thrips

decreased in numbers by July and August

and remained at low densities in the fall.

Estimated densities of thrips were highly

variable between sites and monthly com-

parisons were not significantly different;

however, there were higher thrips densities

in dandelions in the high-weed plots (33.7 +

11.0) than in the low-weed plots (2.9 + 0.7)

when the estimates were summed over the

three years (F=7.79; df=1,6; P=0.031).

The three-year average of thrips per

flowering dandelion plant was 0.22 in high-

weed and 0.33 in low-weed blocks, while

thrips per vegetative plant was 0.04 in high-

weed and 0 in low-weed blocks during peak

dandelion flowering (F=0.38; df=1,6;

P=0.56,). No difference in thrips density

(on a per plant basis) was found in flower-

J. ENTOMOL. Soc. BRIT. COLUMBIA 105, DECEMBER 2008 93

100 | m fais

—co— Low-weed

—@— High-weed

Total Dandelions/m?

Flowering Dandelions/m2

Thrips on Dandelion/m@

ee or : (Nx --Q--@

aie a ee | Se ee ae |

Jan Mar May Jul Sep Nov Jan Mar May Jul Sep Nov Jan Mar May Jul Sep Nov Jan

2003 2004 2005

Figure 1. (A) Dandelions per m’ area of orchard ground in low-weed and high-weed orchard

blocks. (B) Blooming dandelions per m” area of orchard ground in low-weed and high-weed

orchard blocks. (C) Western flower thrips population estimates (adults per m*) on dandelions

in low-weed and high-weed orchard blocks. Symbols are means, error bars are SEM. Grey bars

indicate the period of apple bloom.

94 J. ENTOMOL. Soc. BRIT. COLUMBIA 105, DECEMBER 2008

ing dandelions from either low-weed or

high-weed blocks at any sample period.

Therefore, thrips population density per

dandelion was largely not influenced by

dandelion density. The reduction of thrips

per area was determined by reduction of

dandelions.

Thrips in Apple. All of the thrips speci-

mens collected from apple flowers in the

three years of the experiment were western

flower thrips, F. occidentalis. Western

flower thrips has been the dominant thrips

species collected from apple flowers in the

inland Pacific Northwest (Venables 1925,

Childs 1927, Madsen and Jack 1966, Cock-

field et al. 2007b). In contrast, apple shoots

and dandelions contained a mixture of spe-

cies, including F. occidentalis. Western

flower thrips accounted for about 38% of

the thrips in apple shoots sampled in Wash-

ington state (Cockfield et al. 2007b). No

significant differences in adult western

flower thrips populations were found in any

of the apple flower or shoot samples from

high-weed and low-weed plots, even in

2005, after dandelion numbers had been

greatly reduced in the low-weed plots

(Table 1). Thrips densities at full apple

bloom were average compared with other

samples in Washington (Miliczky ef al.

2007). Thrips in the apple shoots may con-

tribute to fruit injury in late May, but more

likely sustain the population in the orchard

throughout the summer and from year to

year (Cockfield et al. 2007a, 2007b). While

thrips densities provide a useful measure of

the treatment effects, the critical measure-

ment for the purposes of management is

fruit damage. In the study orchards, an in-

secticide treatment was inadequate to pre-

vent fruit injury, indicating the need for an

additional management tactic. However,

significant reductions in numbers of dande-

lions did not correspond with a significant

reduction in fruit injury in any of the three

years (Fig. 2). Thus the effort and time in-

vestment needed to manage broadleaf

weeds did not provide a substantial benefit

to fruit damage reduction.

One possible explanation for the lack of

effect is that thrips moved between high-

weed and low-weed plots, in spite of the

large size of the experimental plots. A sec-

ond and more likely explanation is that the

potential contribution of western flower

thrips from dandelion is relatively small

during late spring, when fruit injury occurs.

While dandelion flowers often harbour

large numbers of thrips, western flower

thrips may be <10% of the total individuals

present (Cockfield et al. 2007b). The per-

hectare estimates of thrips densities indi-

cated that 45,000 and 4,000 adult western

flower thrips per ha (high-weed and low-

weed plots, respectively) occurred on dan-

delion, compared to estimates of 141,000

and 157,000 thrips per ha, respectively, on

apple flowers at peak bloom just before the

critical period for fruit damage. The attrac-

tiveness of flowering plants to western

flower thrips is well established (Terry

1991, 1997). During their bloom periods

there were 1.9 million apple blossoms per

ha compared with 43,000 dandelion blos-

soms; even assuming they are equally at-

tractive, the sheer number of apple flowers

could be expected to dominate this interac-

tion during the bloom period.

There is ample evidence this highly

polyphagous species is abundant in many

hosts other than apple, including crop and

non-crop plants. Recent studies (Pearsall

and Myers 2000, Pearsall and Myers 2001,

Cockfield et al. 2007b) clarify that in the

semi-arid interior fruit growing districts of

the Pacific Northwest, multiple species of

the native vegetation serve as a host for

thrips. Further studies indicate that immi-

gration from the native vegetation may af-

fect thrips density and damage in orchard

borders (Miliczky ef a/. 2007). Even though

the orchards in this study were not adjacent

to native vegetation, it is still potentially a

very large source of thrips in the industry.

Removal of one relatively small source,

dandelion blossoms, would constitute only

a minor change in local populations. This,

coupled with the large resource constituted

by apple blossoms, and to a lesser extent,

vegetative tissues, effectively negates any

benefit of dandelion removal.

J. ENTOMOL. Soc. BRIT. COLUMBIA 105, DECEMBER 2008 95

Table 1.

Western flower thrips, mean (SEM), sampled per apple flower and per shoot in high-weed and

low-weed treatments’.

Low-weed High-weed

Date Flowers Shoots Flowers Shoots F P

May 2003 0.148 (0.068)a 0.106 (0.051)a 142 0.320

June 2003 0.194 (0.034)a 0.181 (0.041)a O.15 0.721

July 2003 0.056 (0.030)a 0.069 (0.01l)a 0.24 0.658

Aug 2003 0.028 (0.020)a 0.028 (0.020)a_ 0.0 1.000

Apr 2004 ~=0.035 (0.012)a 0.025 (0.006)a 1.04 0.382

May 2004 0.388 (0.229)a 0.316 (0.126)a 0.43 0.557

June 2004 0.203 (0.123)a 0.121 (0.059)a 1.54 0.303

July 2004 0.156 (0.080)a 0.113 (0.053)a 1.01 0.388

Apr 2005 ~=—_0.074 (0.027)a 0.089 (0.031)a 4.54 0.123

May 2005 0.028 (0.016)a 0.084 (0.028)a 4.96 0.112

June 2005 0.190 (0.112)a 0.200 (0.122)a 6.00 0.092

July 2005 0.058 (0.011 )a 0.058 (0.018)a 0.0 1.000

'Means within rows followed by the same letter are not significantly different, LSD test,

oa=0.05. For all analyses, df=1,3.

a High-weed

% Injured Fruit

! pa

2003 2004

Figure 2. Percentage of fruit with pansy spot in low-weed and high-weed plots. Bars are

means, error bars are SEM. Means with the same letter are not significantly different within

each year. 2003: F=0.72; P=0.4581. 2004: F=2.33; P=0.2241. 2005: F=1.15; P=0.3625. For all

analyses, df=1,3.

ACKNOWLEDGEMENTS

We thank Cheryl O’Donnell, Depart-

ment of Entomology, University of Califor-

nia, Davis, CA, for identification of species

in the reference collection, and for helpful

information on the preservation and identi-

fication of thrips. We thank Angela Schaub

for assistance in the lab and Randy Talley

for assistance in the field. David Horton,

96 J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

USDA-ARS, Wapato, WA, provided valu- _ part through a grant from the Washington

able contributions to an earlier draft of the | Tree Fruit Research Commission.

manuscript. This research was funded in

REFERENCES

Beers, E.H., J. F. Brunner, M. J. Willet, and G. M. Warner (eds.). 1993. Orchard pest management: a re-

source book for the Pacific Northwest. Good Fruit Grower, Yakima, WA.

Childs, L. 1927. Two species of thrips injurious to apples in the Pacific Northwest. Journal of Economic

Entomology 20: 805-809.

Cockfield, S.D., E.H. Beers, D.R. Horton and E. Miliczky. 2007a. Timing of oviposition of thrips

(Thysanoptera: Thripidae) in apple fruit. Journal of the Entomological Society of British Columbia 104:

45-53.

Cockfield, S.D., E.H. Beers and R.S. Zack. 2007b. Phenology of western flower thrips (Frankliniella occi-

dentalis (Pergande) (Thysanoptera: Thripidae) on plant species in and near apple orchards in Washing-

ton. Journal of the Entomological Society of British Columbia 104: 35-44.

Cossentine, J.E., E.J. Hogue and L.B.M. Jensen. 1999. The influence of orchard ground cover and intro-

duced green lacewings on spring populations of western flower thrips in apple orchards. Journal of the

Entomological Society of British Columbia 96: 7-12.

Hubscher, T.L. 1983. An investigation into the relationship between western flower thrips damage and

orchard floor management in British Columbia apple orchards. Master of Pest Management Thesis,

Simon Fraser University, Burnaby, B.C.

Madsen, H.F. and I.D. Jack. 1966. The relation of thrips to pansy spot on apples. The Canadian Entomolo-

gist 98: 903-908.

Miliczky, E., S.D. Cockfield, E.H. Beers and D.R. Horton. 2007. Spatial patterns of western flower thrips in

apple orchards and associated fruit damage. Journal of the Entomological Society of British Columbia

104: 25-33.

Newcomer, E.J. 1921. A thrips injury to apple. Better Fruit 16:10.

Pearsall, I.A. and J.H. Myers. 2000. Population dynamics of western flower thrips (Thysanoptera: Thripi-

dae) in nectarine orchards in British Columbia. Journal of Economic Entomology 93: 264-275.

Pearsall, I.A. and J.H. Myers. 2001. Spatial and temporal patterns of dispersal of western flower thrips

(Thysanoptera: Thripidae) in nectarine orchards in British Columbia. Journal of Economic Entomology

94: 831-843.

SAS Institute. 2002. User's guide. SAS Institute, Cary, NC.

Terry, L.I. 1991. Frankliniella occidentalis (Thysanoptera: Thripidae) oviposition in apple buds: role of

bloom state, blossom phenology, and population density. Environmental Entomology 20: 1568-1576.

Terry, L.I. 1997. Host selection, communication and reproductive behaviour, pp 65-118., pp. 65-118. Jn T.

Lewis (ed.), Thrips as crop pests. CAB International, New York, NY.

Venables, E.P. 1925. Pansy spots on apple (a peculiar form of thrips injury). Proceedings of British Colum-

bia Entomological Society 22: 9-12.

J. ENTOMOL. Soc. BRIT. COLUMBIA 105, DECEMBER 2008

Effect of pesticides on integrated mite management

in Washington State

LUIS MARTINEZ-ROCHA’, ELIZABETH H. BEERS'”

and JOHN E. DUNLEY'

ABSTRACT

The effect of pesticides used against codling moth, Cydia pomonella L., on integrated

mite management was studied for three years in five or six commercial apple orchards in

central Washington. Phytophagous and predatory mites were counted throughout the

season in blocks ranging from 0.4-1.6 ha in size treated with four codling moth insecti-

cides. In one year of the study (2006), five out of six orchards experienced elevated mite

densities relative to the standard. In four orchards, novaluron caused a 3.0-16.9*x in-

crease in mite populations; acetamiprid caused a 2.6-3.4* increase, and thiacloprid

caused a 1.7-13.8x increase. In the fifth orchard, the organophosphate standard had an

extremely high mite population, in addition to all three experimental treatments. In 2005

and 2007, only one or two orchards had elevated mite levels in the novaluron, acetami-

prid, and thiacloprid treatments. Additive effects of codling moth and thinning programs

were evaluated in small plot research trials. Treatments with all three elements [1) cod-

ling moth insecticide; 2) calcium polysulfide; 3) carbaryl] produced the highest levels of

spider mites. Three sulfur-containing products (calcium polysulfide, ammonium thiosul-

fate, and dry flowable sulfur) were evaluated for their effect on Galandromus occiden-

talis (Nesbitt) and apple rust mite, Aculus schlechtendali (Nalepa). All three materials

caused suppressed G. occidentalis numbers. Calcium polysulfide caused the greatest

reduction in apple rust mite numbers, ammonium thiosulfate the least reduction, with

dry flowable sulfur intermediate between the two. Additive effects of codling moth ma-

terials, carbaryl, and sulfur-containing products may be causing increased mite levels in

Washington orchards.

Key Words: spider mite, integrated mite control, apple, acetamiprid, thiacloprid, no-

valuron, carbaryl, calcium polysulfide, lime-sulfur, dry flowable sulfur, ammonium thi-

osulfate

INTRODUCTION

Spider mites are induced pests of pome

fruits, generally occurring in orchards

which have been disrupted by pesticides. In

wild or abandoned trees, spider mites are

usually maintained at low densities by

predators (Glass and Lienk 1971, Croft

1983). However, in heavily sprayed sys-

tems such as commercial apple orchards,

perturbations occur regularly based on the

need to control direct pests. In Washington

State, codling moth, Cydia pomonella L.., is

the key direct pest of apple, and control

measures used against it determine the en-

tire pest management program and structure

the fauna of the agroecosystem.

The history of spider mite management

in apple orchards is characterized by dis-

ruption of mite biological control following

the introduction of new materials for cod-

ling moth control. DDT was introduced

following WWII, and its use in tree fruits

was accompanied by large scale mite out-

'Tree Fruit Research and Extension Center, Washington State University, 1100 N. Western Avenue, We-

natchee, WA 98801 USA

* Corresponding author. Fax: 509-662-8714; email: ebeers@wsu.edu

98 J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

breaks (Newcomer and Dean 1946, Baker

1952, Clancy and McAlister 1956). As a

result of the disruption, acaricide resistance

became widespread (Hoyt and Caltagirone

1971). DDT was replaced by organophos-

phate insecticides, including azinphos-

methyl, which initially were toxic to both

pest and predatory mites. Spider mites be-

came resistant to the organophosphates, but

it was not until the primary predator of spi-

der mites, Galandromus (=Typhlodromus =

Metaseiulus) occidentalis (Nesbitt) also

became resistant that the opportunity for

integrated mite control arose in the western

US (Hoyt 1969). Integrated mite manage-

ment was then implemented on approxi-

mately 90% of the acreage (Whalon and

Croft 1984) by conserving the organophos-

phate-resistant predatory mites, with grow-

ers actively avoiding materials that were

toxic to these valuable predators. This op-

portunity, however, was predicated on con-

tinuing efficacy of azinphosmethy] for cod-

ling moth control.

The integrated mite control program

implemented in the early 1970s in Wash-

ington remained largely effective through

the 1990s. During this time shifts in the

pesticide program occurred, at least for

pests other than codling moth. The efficacy

of the organophosphates against many of

the secondary pests of tree fruit (aphids,

leafhoppers, leafminers, and _leafrollers)

declined steadily through this period, and

new materials were substituted for their

control. The use of carbaryl for fruit thin-

ning was implemented in the late 1970s;

this carbamate insecticide was initially

highly toxic to G. occidentalis, and its use

was restricted to protect predator popula-

tions in integrated control programs. How-

ever, moderate levels of resistance to this

carbamate were documented within a short

period of use (Babcock and Tanigoshi

1988). Other carbamate insecticides, simi-

larly toxic to the predator, were also used

sparingly, typically only when no other

substitute was available. The use of pyre-

throids, notoriously disruptive to integrated

mite control, was largely avoided in Wash-

ington apples in order to protect the inte-

grated mite control program.

However, following almost forty years

of reliance on azinphosmethyl for codling

moth control, shifts in the codling moth

management program began in the mid-

1990s. These changes have been driven

mainly by either the development of or-

ganophosphate resistance in codling moth

or by regulatory issues (Beers ef al. 2005).

In the meantime, new control techniques

and materials have increased in use. Mating

disruption now forms the foundation of

codling moth control in about 80% of

Washington’s apple orchards (J. Brunner,

personal communication). Choices for sup-

plementary insecticides include insect

growth regulators and _neonicotinoids.

While the new materials meet the new stan-

dard for improved worker safety, their ef-

fects on natural enemies and predator/prey

dynamics have not been well explored.

One class of alternative insecticide

chemistry, the neonicotinoids, demonstrated

a tendency to disrupt integrated mite con-

trol even in the early phases of testing

(Beers et al. 2005). These tests, however,

were characterized by small plots, high

rates, and season-long programs. It re-

mained to be seen if the potential for dis-

ruption still existed under commercial use

conditions (large acreages, applications

against a single generation of codling moth

with any given material). Additionally, po-

tential multiple-year effects from continued

use of the same products could not easily be

examined prior to registration.

Other shifts in the Washington pesticide

program occurred during the same period as

the change in the codling moth program,

especially in the crop load regulation and

fungicide programs. The blossom thinner,

sodium dinitro-o-cresylate (Elgetol®), was

withdrawn from the market in the mid

1990s, and its use was later replaced with

calcium polysulfide (lime sulfur). A plant

nutrient, ammonium thiosulfate, also be-

came more widely used; its sulfur content is

similar to calcium polysulfide. The use of

sulfur fungicides increased in part because

of the plantings of mildew-susceptible culti-

vars, and in part as an alternative mode of

J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

action for fungicide resistance management

(FRAC 2008).

The goals of this study were to explore

the effects on integrated mite control of

three newer codling moth insecticides when

used in a commercial setting; to examine

the additive effects of codling moth insecti-

cides, carbaryl, and calcium polysulfide in a

seasonal program; and the comparative

effects of three sulfur-containing com-

pounds used as a blossom thinner, fungi-

cide, and plant nutrient, respectively.

MATERIALS AND METHODS

Large-block experiment. This test was

conducted in five (2005, 2007) or six

(2006) commercial apple orchards from

Bridgeport to Royal City, WA. Plot size

ranged from 0.4-1.6 ha per treatment at

each orchard; treatments were randomly

assigned to one of four plots within an or-

chard, and replicated across the orchards.

All treatments were applied at a finished

spray volume of 935 litres/ha by the or-

chard’s personnel using their own equip-

ment. The dominant cultivar in the blocks

was either ‘Delicious’ or ‘Fuji’. Four of the

orchards received the same treatments for

all three years of the study (orchards ARR,

BAN, QLR, SLH); MZN was treated in

2006-07, while RYL and BTE were treated

only in 2005 and 2006, respectively.

Treatments consisted of one of the four

insecticides used for codling moth control:

acetamiprid, a neonicotinoid (Assail® 70W,

Cerexagri, King of Prussia, PA; 0.17 kg AI/

ha); thiacloprid, also a neonicotinoid

(Calpyso® 4F, Bayer CropScience, Re-

search Triangle Park, NC; 0.21 kg Al/ha);

novaluron, a benzoylurea insect growth

regulator (Rimon® 0.83EC, Chemtura, Mid-

dlebury, CT; 0.23 g Al/ha); and an organo-

phosphate standard. For the organophos-

phate standard, growers could choose either

phosmet (Imidan® 70W, Gowan, Yuma AZ;

3.9 kg Al/ha) or azinphosmethyl (Guthion®

50W, Bayer CropScience, Research Trian-

gle Park, NC; 1.1 kg Al/ha). Applications

of acetamiprid, thiacloprid, and novaluron

were made only during the first generation

of codling moth (May and June). Two ap-

plications of acetamiprid and _ thiacloprid

were made per season, the first timed for

250 codling moth degree days, and the sec-

ond 21 d later. Three applications of no-

valuron were made (based on its ovicidal

activity), the first at petal fall, the second 14

d later, and the third 28 d later.

For the above the treatments, codling

moth control for the second generation

(July and August) consisted of two applica-

tions of the benzoyl hydrazine insect

growth regulator methoxyfenozide

(Intrepid® 2F, Dow AgroSciences, Indian-

apolis, IN; 0.28 kg Al/ha), the first timed

for 1,250 codling moth degree days, and the

second 21 d later.

In the organophosphate standard treat-

ment, two applications were made per gen-

eration, using the same timing as for

acetamiprid and methoxyfenozide for the

first and second generations, respectively.

Mites were sampled every 2-3 wk from

late May through mid-September. One hun-

dred leaves per plot were collected from the

center portion of the plot and kept cool dur-

ing transportation and storage. The mites

were brushed from the leaves using a mite

brushing machine (Leedom, Mi-Wuk Vil-

lage, CA) and collected on a revolving

sticky glass plate. The composite sample on

the plate was counted using a stereoscopic

microscope. Phytophagous and predatory

mites were recorded, including the motile

stages of European red mite, Panonychus

ulmi (Koch); twospotted spider mite,

Tetranychus urticae Koch; McDaniel spider

mite, Tetranychus mcdanieli McGregor

western predatory mite, G. occidentalis; a

stigmaeid predatory mite, Zetzellia mali

Ewing, and apple rust mite, Aculus

schlechtendali (Nalepa).

Additive effects experiment. This

small-plot experiment examined the effect

of adding one or two potentially disruptive

compounds used for crop load regulation to

the same codling moth programs (rates,

timing, and materials) described for the

100 J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

large plot experiment. The compounds used

were a blossom thinner, calci1um polysul-

fide (Rex Lime Sulfur®, Or-Cal, Junction

City, Oregon), and a fruit thinner, carbaryl

(Sevin® 4F, Bayer CropScience, Research

Triangle Park, NC). Both compounds were

used at their respective recommended tim-

ings (Smith et a/. 2006). Calctum polysul-

fide was applied three times (pink, 20% and

80% bloom) at a rate of 8% vol:vol. Car-

baryl was applied twice, when the fruitlets

were 8 and 12 mm in diameter, at rate of

1.7 kg Al/ha.

This test was conducted in a 2 ha block

of mature ‘Oregon Spur’ and ‘Red Spur’

Delicious apples with ‘Golden Delicious’

pollenizers. Plots were five rows by five

trees. The experimental design was a ran-

domized complete block design with 12

treatments and 4 replicates. All applications

were made with an airblast sprayer (Rears

Pak-Blast, Eugene, OR) calibrated to de-

liver 935 litres/ha. Treatment timings and

materials for first and second generation

codling moth control were the same as de-

scribed in the large-block experiment.

Mites were sampled every other week

from May through September by collecting

40 leaves per plot. The leaves were col-

lected, stored and processed as described

above.

Sulfur products experiment. The sec-

ond small-plot experiment examined the

effect of three sulfur-containing products on

G. occidentalis and apple rust mite. For

purposes of comparison, the materials were

applied to an existing population of these

two species in June, rather than at their nor-

mal timing which ranged from prebloom

through the early post-bloom period. The

three sulfur products were calcium polysul-

fide (Rex Lime Sulfur®, Or-Cal, Junction

City, Oregon; 12% vol:vol), ammonium

thiosulfate (a plant nutrient) (Thio-Sul®,

Tessenderlo Kerley, Phoenix, AZ; 3.4%

vol:vol), and dry flowable sulfur (a fungi-

cide) (Kumulus® 80DF, Micro-Flo, Mem-

phis, TN; 10.8 kg Al/ha).

The experimental design was a random-

ized complete block (randomized on the

basis of a pretreatment count) with seven

treatments and four replicates. Each repli-

cate consisted of three trees in a single row,

with one untreated buffer row separating

the treatment rows. Plots consisted of three

cultivars, “Oregon Spur’, ‘Goldspur’, with

‘Red Fuji BC2’ in the center; however, only

the center tree was sampled. Treatments

consisted of either one or three applications

of each of the three sulfur-containing com-

pounds plus an untreated check. Treatments

receiving a single application were applied

26 June 2006; treatments receiving three

applications were made 26 June, 8 July and

19 July. Treatments were applied by air-

blast sprayer at 935 litres/ha. Mite popula-

tions were assessed by collecting 25 leaves

per plot and processed using the method

described above. Counts were made pre-

treatment and weekly after treatment

through late July.

Data analysis. Cumulative mite days

(CMDs) were calculated for tetranychid (P.

ulmi plus T. urticae), predatory (G. occi-

dentalis plus Z. mali), and apple rust mite.

CMDs provide an estimate of population

densities integrated over the course of the

test, and are calculated as the sums of the

average density of mites on two dates mul-

tiplied by the number of intervening days:

CMD = 20.5(P,+P,)Da+

where P, and P, are the population den-

sities (mean mites/leaf) at times a and b,

and D,» is the number of days between

time a and time b.

Data were analyzed using the Statistical

Analysis System (SAS 1988). Data were

tested prior to analysis for homogeneity of

variance using Levene’s (1960) test. Vari-

ances found to be non-homogeneous were

transformed [In(y+0.5)] before analysis.

PROC GLM was used to conduct an analy-

sis of variance, and treatment means were

separated using the Waller-Duncan k-ratio

t-test. Single degree-of-freedom contrasts

were used to compare groups of treatments

in the small plot experiments.

J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

101

RESULTS

Large-block experiment. Spider mite

populations in the experimental blocks con-

sisted primarily of European red mite, with

99, 92, and 78% of the population of motile

forms comprised of this species in 2005,

2006, and 2007, respectively. Twospotted

spider mite was the next most numerous

species; a relatively higher proportion of

this species occurred only in one orchard,

MZN, in 2007, when 57% of the motile

forms were twospotted spider mite. Only

trace numbers of McDaniel spider mite

were found during the course of the study.

There were no statistical differences

among treatment mean CMDs for

tetranychids or for predatory mites in any of

the three years of the study (Table 1). In

2005 and 2006, rust mite populations were

higher in the acetamiprid treatment com-

pared to the standard. In general, the differ-

ences in densities among years and or-

chards were greater than those among treat-

ments.

Despite this variation, some trends in

these data are apparent. In 2005, elevated

mite densities occurred in only one of five

orchards (QLR) (Fig. 1). However, the

highest mite levels occurred in the novalu-

ron (peak density 54 mites/leaf), acetami-

prid (21 mites/leaf), and thiacloprid (20

mites/leaf) treatments, with only a moderate

increase in the organophosphate standard

treatment (11 mites/leaf). Not all of the

population peaks in the treatments with

newer insecticides can be explained by low

predator numbers, although predatory mite

densities were highest in the organophos-

phate treatment (peak density 2.3 predators/

leaf). The thiacloprid treatment peaked at

0.9 predators/leaf, while the acetamiprid

and novaluron treatments never exceeded

predator densities of 0.3/leaf. Apple rust

mite densities were moderate in most of the

treatment (peak density of 150-300 rust

mites/leaf), with the exception of the no-

valuron treatment, which had relatively low

rust mite densities (<10/leaf) for most the

season.

Mite densities were much higher overall

in 2006 than in 2005 (Fig. 1). Five of six

orchards experienced elevated tetranychid

mite densities in one or more treatments.

This may have been due to a cumulative

effect of disruptive products in four of the

orchards; however, 2006 was characterized

by a high frequency of mite outbreaks

throughout the central fruit-growing district

of the state. The novaluron treatment had

elevated tetranychid mite levels in five or-

chards (20-45 mites/leaf at peak density);

the acetamiprid treatment in two orchards

(22-34 mites/leaf); and the thiacloprid treat-

ment in three orchards (13-32 mites/leaf).

One orchard (QLR) had a high peak mite

density in the organophosphate treatment,

as well as the other three treatments; how-

ever, this was the same orchard that had

high levels in several treatments the previ-

ous year. Trends in predatory mite densities

were again difficult to interpret. Although

no statistical differences occurred among

treatment means for the entire season, the

peak densities of predators occurred too late

in the season to prevent the mid-July peak

in tetranychid mites (data not shown).

Mite densities in the experimental or-

chards were much lower in 2007 than in

2006 (Fig. 1), with no treatment exceeding

7 mites/leaf. Only two of the five orchards

experienced a moderate increase in

tetranychid mite levels, with a slight eleva-

tion in the novaluron treatment in one or-

chard (SLH) (6.2 mites/leaf peak density),

and acetamiprid in two orchards (4.4 and

3.5 mites/leaf in MZN and SLH, respec-

tively) and thiacloprid (3.6 mites/leaf) in

one orchard (MZN).

Additive effects experiment. The re-

sults from the experiment examining the

additive effect of several disruptive prod-

ucts during the season showed a distinct

trend toward increased tetranychid mite

densities when one of the newer codling

moth insecticides was used in the same

program with both a blossom and fruit thin-

ner (Fig. 2). The lowest tetranychid mite

densities occurred in those treatments

where only insecticides for codling moth

were used. Treatments where all three com-

pounds were used (codling moth insecticide

102 J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

Table 1.

Seasonal mite densities (cumulative mite days) resulting from four codling moth control re-

gimes, in commercial apple orchards in Washington, 2005-2007

Cumulative mite days!

Treatment Rate(AlI/ha)~ n een s Per ane Merial

2005

Acetamiprid 0.17 kg 5 113+ 99a i242a 10,861 + 2,690a

Thiacloprid 0.21 kg 5 93 + 84a 17+4a 8,563 + 1,718ab

Novaluron 0.23 kg 5 252 + 244a 11 +3a 6,589 + 2,639b

Standard --- 5 49 + 38a 26 + 8a 6,668 + 1,394b

Vas es 4.32, 0.013 1.90, 0.16 12.03, 0.0001

2006

Acetamiprid 0.17 kg 6 267 + 132a 26 + 3a 870 + 372a

Thiacloprid 0.21 kg 6 305 + 149a 28 + 5a 634 + 244ab

Novaluron 0.23 kg 6 520 + 178a 33 2.134 563 + 30lab

Standard --- 6 401 + 343a 27 45a 476 + 197b

ye 4.78, 0.0045 1.98, 0.12 19.80, <0.0001

2007

Acetamiprid 0.17 kg > 55+3la 29 + 13a 719 + 346a

Thiacloprid 0.21 kg > 18+ 10a 35+ 14a 883 + 240a

Novaluron 0.23 kg 5 3742358 16+ 8a 346 + 153a

Standard --- > 25 5a 32 + loa 632 + 229a

ae a 6.17, 0.0032 5.74, 0.0043 3.43, 0.030

'Means within columns not followed by the same letter are significantly different. For 2005

and 2007, df=7, 19; for 2006, df=8, 23.

+ calcium polysulfide + carbaryl) had sig-

nificantly higher tetranychid mite densities

than when codling moth insecticides alone

were used (df=1, F=5.54, P=0.02).

Trends in seasonal densities of preda-

tory mites and apple rust mite were less

clear (Figs. 3, 4). Comparisons of treat-

ments with or without calcium polysulfide

indicated that there was a significant reduc-

tion in the seasonal apple rust mite densities

where calcium polysulfide was included in

the program (df=1, F=5.07, P=0.03), how-

ever, there was no effect on predatory mite

densities (df=1, F=0.70, P=0.41).

Sulfur products experiment. All three

sulfur products used in this study sup-

pressed G. occidentalis to about the same

extent (Fig. 5). There was a 64-74% reduc-

tion in densities of G. occidentalis in the

treatments containing sulfur products in

relation to the check. There was no differ-

ence between treatments with one applica-

tion versus three applications (df=1,

F=0.11, P=0.75), likely because most of the

mortality had occurred from the first appli-

cation, without sufficient time for reinfesta-

tion between applications.

The effect of the three sulfur products

on apple rust mite was more variable. There

was a 30-80% reduction in densities of ap-

ple rust mite in these treatments. The reduc-

tion in apple rust mite numbers was greatest

in the calcium polysulfide treatment (Fig.

6), and least in the ammonium thiosulfate

treatment. As with G. occidentalis, there

were no differences in treatment means

between treatments with one versus three

applications (df=1, F=0.99, P=0.33).

J. ENTOMOL. Soc. BRIT. COLUMBIA 105, DECEMBER 2008

Organophosphate

Novaluron

Thiacloprid

Acetamiprid

"PT IT LILLIE ED LILI CLLLALLL ADL LALA LAL ALL LLL LAL

Organophosphate

Novaluron

Thiacloprid

Acetamiprid

Organophosphate

Novaluron

Thiacloprid

Acetamiprid

400

103

QLR

besgq SLH

600 800 1000

Cumulative Mite Days {Tetranychids)

Figure 1. Seasonal tetranychid mite densities (cumulative mite days) in commercial apple or-

chard blocks treated with four insecticides for codling moth control, 2005-2007.

DISCUSSION

The responses to the two neonicotinoid

insecticides used in the large-block study

confirms previous work done on small plots

(Beers et al. 2005). Mite populations in the

acetamiprid treatments averaged 2.3, 2.2

and 3.0x higher than the standard organo-

phosphate treatment during 2005-2007,

respectively. Mite populations in the thia-

cloprid treatments averaged 1.3, 3.4 and

2.2x higher than the standard. In addition to

the neonicotinoids, this study provides evi-

dence that novaluron also causes disruption

of integrated mite management, although

this trend was not apparent in small-plot

trials (J. Brunner, personal communication).

Mite populations in the novaluron treat-

ments were 2.1, 7.6, and 2.7x higher than

the standard treatment in the three years of

the study.

Although widely observed, the mecha-

104 J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

CaPoly+Carbaryl+Phosmet -

CaPoty+Phosmet

Phosmet -

CaPoly+Carbaryl+Novaluron

CaPoh+Novahiron

Novaluron

CaPoly+Carbaryl+Thiacloprid

CaPoly+Thiacloprid +

Thiacloprid

CaPoly+Carbaryl+Acetamiprid -

CaPolytAcetamiprid +

Acetamiprid

0) OQ 200) OO 400 DOO G00

Cumutative Mite Days (Tetranychids)

Figure 2. Additive effect of thinning materials and codling moth insecticides on seasonal

CaPoly+Carbaryl4+ Phosmet

CaPoly+Phosmet 4 | ab

Phosmet4 a

CaPoly+Carbaryl+Novaluron b

CaPoly+Novalurony

Novaluon+} b

CaPoly+Carbaryl+Thiacloprid ab

CaPoly+Thiacloprid ab

Thiacloprid

CaPoly+Carbaryl+Acetamiprid ab

eee

CaPoly+Acetamiprid | ab

ee os

Acetamiprid

60 80

Cumulative Mite Days (Predatory Mites)

Figure 3. Additive effect of thinning materials and codling moth insecticides on seasonal

predatory mite populations. F=2.05, P=0.054, df=11, 47. Data transformed log(x+0.5) prior to

jas)

(os)

j=)

nism for the neonicotinoid effect has never effect (Ako ef al. 2004, Ako et al. 2006).

been clearly established. Hormoligosis is

thought to play a role in stimulating pest

reproduction (James and Price 2002), but

other studies have found no hormoligosis

Conversely, neonicotinoids have also been

found to stimulate reproduction in benefi-

cial arthropods (James 1997). Repellency

(James 1997) and suppression of functional

J. ENTOMOL. Soc. BRIT. COLUMBIA 105, DECEMBER 2008

CaPoly+Carbaryl+Phosmet

CaPoly+Phosmet

Phosmet

CaPoly+Carbaryl+Novaluron

CaPoly+Novaluron

Novaluron

CaPoly+Carbaryl+Thiacloprid

CaPoly+Thiacloprid

Thiacloprid

CaPoly+Carbaryl+Acetamiprid

CaPoly+Acetamiprid

Acetamiprid

0 1000

105

2000 3000 4000 5000

Cumulative Mite Days (Apple Rust Mites)

Figure 4. Additive effect of thinning materials and codling moth insecticides on seasonal apple

rust mite populations. F = 3.03, P = 0.0067, df= 11, 47.

Check

Amm. Thiosulfate (3x)

Amm. Thiosulfate (1x)

Sulfur (dry flowable) (3x)

Suflur (dry flowable) (1x)

Ca Polysulfide (3x)

Ca Polysulfide (1x)

0 2 4

6 8 10 12 14 16

Cumulative Mite Days (Predatory Mites)

Figure 5. Effect of sulfur-containing products on predatory mites. F' = 2.68, P=0.0.049, df= 6,

2d.

response (Poletti et al. 2007) may also play

a role in the disruption of biological con-

trol.

It is evident from previous studies

(Beers et al. 2005) that while neonicoti-

noids can cause mite outbreaks, they would

not do so in every case. This makes the role

of other disruptive materials more impor-

tant on a relative scale. In an organophos-

phate-based pest management program,

calcium polysulfide and carbaryl had been

used with few apparent deleterious effects;

under this program, only about 7% of

Washington’s apple orchards were treated

with acaricides (NASS 1992). The low mite

levels documented by the survey are likely

typical of acaricide use from the early

1970s, when integrated mite control was

first established, until the early 2000s when

shifts in codling moth insecticides began.

However, there has been a substantial in-

crease in the percentage of Washington

apple acreage treated with sulfur fungicides

(7.8x) and calcium polysulfide (11) since

106 J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

Check

Amm. Thiosulfate (3x)

Amm. Thiosulfate (1x)

Sulfur (dry flowable) (3x)

Suflur (dry flowable) (1x)

Ca Polysulfide (3x)

Ca Polysulfide (1x)

0 200

400 600 800

MOLD A hy LL LOG AOE,

CLI M MLD ky LG LA

1000

Cumulative Mite Days (Apple Rust Mites)

Figure 6. Effect of sulfur-containing products on apple rust mite. F = 22.3, P = <0.0001, df=

6, 27.

1991 (NASS 1992, 2006). These com-

pounds, which are typically applied early in

the season, may predispose the orchard to

later disruption by codling moth insecti-

cides.

The toxic effect of carbaryl and sulfur-

containing products on mites is well known

(McMurtry ef al. 1970). In the case of car-

baryl, moderate levels of resistance in G.

occidentalis (Babcock and Tanigoshi 1988)

may have mitigated the disruptive effect to

some extent. Sulfur products have a long

history of disruption of integrated mite con-

trol, and although resistance in G. occiden-

talis populations had been found in Califor-

nia vineyards, this had apparently not oc-

curred in Washington orchards (Hoy and

Standow 1982). Thus it is reasonable to

expect that increased use of these materials

could contribute to mite outbreaks.

The organophosphate—based programs

of the past few decades have provided one

of the most stable periods in integrated mite

control in Washington orchards. The insec-

ticides that replaced the organophosphates

were initially thought to be more selective,

but a number have shown nontarget effects

on beneficial arthropods. Because of this

destabilization, acaricide use has increased

in recent years (NASS 1992, 2006), leading

to increased production costs and increasing

the probability for resistance development

in pest mite species. It could be argued that

because of widespread resistance to organo-

phosphates in populations of both pests and

natural enemies, that many of the organo-

phosphates are now fairly selective from a

pest management perspective. While human

and environmental health concerns out-

weigh pest management issues, it will re-

quire further study and manipulation to re-

establish the highly successful integrated

mite control program as the primary means

of mite control in Washington apple or-

chards.

ACKNOWLEDGEMENTS

We thank the cooperating growers and

fieldmen who generously allowed us to use

their orchards for this experiment. Thanks

also go to Randy Talley, Amy Blauman,

Michelle Blauman, Aaron Galbraith, Tim

Houghland, Kelly Green and Brandon

Talley for assistance with mite counts. We

also thank Bayer CropScience, Cerexagri,

Chemtura, Dow Agrosciences, and Gowan

for donation of materials. This work was

funded in part by grants from the Washing-

ton Tree Fruit Research Commission and

the Washington State Commission on Pesti-

cide Registration.

J. ENTOMOL. SOc. BRIT. COLUMBIA 105, DECEMBER 2008 107

REFERENCES

Ako, M., C. Borgemeister, H.-M. Poehling, A. Elbert and R. Nauen. 2004. Effects of neonicotinoid insecti-

cides on the bionomics of twospotted spider mite (Acari: Tetranychidae). Journal of Economic Entomol-

ogy 97: 1587-1594.

Ako, M., H.-M. Poehling, C. Borgemeister and R. Nauen. 2006. Effect of imidacloprid on the reproduction

of acaricide-resistant and susceptible strains of Tetranychus urticae Koch (Acari: Tetranychidae). Pest

Management Science 62: 419-424.

Babcock, J.M. and L.K. Tanigoshi. 1988. Resistance levels of Typhlodromus occidentalis (Acari: Phytoseii-

dae) from Washington apple orchards to ten pesticides. Experimental and Applied Acarology 4: 151-157.

Baker, H. 1952. Spider mites, insects, and DDT. Yearbook of Agriculture, 1952: 562-566.

Beers, E.H., J.F. Brunner, J.E. Dunley, M. Doerr and K. Granger. 2005. Role of neonicotinyl insecticides in

Washington apple integrated pest management. Part II. Nontarget effects on integrated mite control.

Journal of Insect Science 5: 16.

Clancy, D.W. and H.J. McAlister. 1956. Selective pesticides as aids to biological control of apple pests.

Journal of Economic Entomology 49: 196-202.

Croft, B.A. 1983. Introduction, pp. 1-18. Jn B. A. Croft and S. C. Hoyt (eds.), Integrated management of

insect pests of pome and stone fruits. Wiley Interscience, New York.

FRAC. 2008. Fungicide Resistance Action Committee. FRAC code list. http://www.frac.info/frac/

publication/anhang/FRAC_Code_List_2007_web.pdf.

Glass, E.H. and S.E. Lienk. 1971. Apple insect and mite populations developing after discontinuance of

insecticides: 10-year record. Journal of Economic Entomology 64: 23-26.

Hoy, M.A. and K.A. Standow. 1982. Inheritance of resistance to sulfur in the spider mite predator Metaseiu-

lus occidentalis. Entomologia Experimentalis et Applicata 31: 316-323.

Hoyt, S.C. 1969. Integrated chemical control of insects and biological control of mites on apple in Washing-

ton. Journal of Economic Entomology 62: 74-86.

Hoyt, S.C. and L.E. Caltagirone. 1971. The developing programs of integrated control of pests of apples in

Washington and peaches in California, pp. 395-421. Jn C. B. Huffaker (ed.), Biological control. Plenum

Publishing Corporation, New York, NY.

James, D.G. 1997. Imidacloprid increases egg production in Amblyseius victoriensis (Acari: Phytoseiidae).

Experimental and Applied Acarology 21: 75-82.

James, D.G. and T.S. Price. 2002. Fecundity in twospotted spider mite (Acari: Tetranychidae) is increased

by direct and systemic exposure to imidacloprid. Journal of Economic Entomology 95: 729-732.

Levene, H. 1960. Robust tests for equality of variances. /m I. Olkin, S. G. Ghurye, W. Hoeffding, W. G.

Madow, and H. B. Mann (ed.), Contributions to probability and statistics. Stanford University Press,

Stanford, CA.

McMurtry, J.A., C.B. Huffaker and M. Van de Vrie. 1970. Ecology of the tetranychid mites and their natu-

ral enemies: A review. I. Tetranychid enemies: their biological characters and the impact of spray prac-

tices. Hilgardia 40: 331-458.

NASS. 1992. Agricultural chemical usage, 1991 fruit crops, NASS. National Agricultural Statistics Service,

Washington, DC.

NASS. 2006. Agricultural chemical usage, 2005 fruit summary, National Agricultural Statistics Service.

NASS, Washington, DC.

Newcomer, E.J. and F.P. Dean. 1946. Effect of Xanthone, DDT, and other insecticides on the Pacific mite.

Journal of Economic Entomology 39: 783-786.

Poletti, M., A.H.N. Maia and C. Omoto. 2007. Toxicity of neonicotinoid insecticides to Neoseiulus califor-

nicus and Phytoseiulus macropilis (Acari: Phytoseiidae) and their impact on functional response to

Tetranychus urticae (Acari: Tetranychidae). Biological Control 40: 30-36.

SAS. 1988. Statistical Analysis Institute. User's guide, Cary, NC.

Smith, T.J., J.E. Dunley, E.H. Beers, J.F. Brunner, G.G. Grove, C.-l. Xiao, D. Elfving, F.J. Peryea, R.

Parker, M. Bush, C. Daniels, T. Maxwell, S. Foss and S. Martin. 2006. 2006 Crop protection guide for

tree fruits in Washington. EB0419, Washington State University Cooperative Extension, Pullman, WA.

Whalon, M.E. and B.A. Croft. 1984. Apple IPM implementation in North America. Annual Review of En-

tomology 29: 435-470.

108 J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

J. ENTOMOL. Soc. BRIT. COLUMBIA 105, DECEMBER 2008

109

SCIENTIFIC NOTE

Additional records for semiaquatic Hemiptera

in southwestern British Columbia

R.D. KENNER! and K.M. NEEDHAM?

Scudder (1977) published an annotated

checklist of the aquatic and semiaquatic

Hemiptera of British Columbia (BC); sub-

sequently, several more species were added

(Kenner and Needham 2004, Maw ef al.

2000, Scudder 1986). Scudder (1977) lists

only a single record for Hydrometra martini

Kirkaldy: Lytton, 26.vii.1931, W. Downes.

For Mesovelia mulsanti White, three re-

cords are given for southwestern BC, all

from Vancouver Island: Elk Lake (Saanich

district), 1x.1924, W. Downes; Duncan,

4.1x.1926, W. Downes; Malahat, 4.1x.1929,

W. Downes; as well as several records for

the interior of BC. We report additional

records for these two species from south-

western BC. Voucher specimens for all new

records were deposited in the Spencer Ento-

mological Museum (SEM) at the University

of British Columbia.

Hydrometridae, water measurers or

marsh treaders, are small bugs resembling

tiny walkingstick insects (Phasmida) and

have long, slender heads and bulging eyes

at about mid-length on the head. They are

often found walking slowly over the water

surface searching for prey. There are nine

species in North America, all in the genus

Hydrometra (Polhemus 2008); only one of

these, H. martini, is known from Canada

(and BC) (Maw et al. 2000). We collected a

single specimen of H. martini from a 10 m

wide roadside ditch in Pitt Meadows (ditch

between Neave Road and the Pitt River

dike, 49°20’N 122°38’W, 12.iv.2008, R.D.

Kenner and K.M. Needham). The specimen

is apterous, strongly suggesting breeding at

this site. This appears to be only the second

time this species has been collected in this

province. The 77 year gap between records,

even though knowledgeable collectors have

been looking for this species (Scudder pers.

comm.), suggests that H. martini is truly

rare in BC.

Mesoveliidae, water treaders, are small

predatory bugs usually found crawling or

running over the water surface. There are

three species in North America, all in the

genus Mesovelia (Polhemus 2008), two of

which are known from Canada, with only

one species, M. mulsanti known from BC

(Maw ef al. 2000). We report the first re-

cords for M. mulsanti from the Lower Fra-

ser Valley: Bowen Island, Killarney Lake,

49°23°N_ = 123°21’W, 9 20.vili.1998, R.D.

Kenner, 1 specimen, apterous; Vancouver,

Jericho Park, Main Pond, west end, 49°

16’N 123°11’W, 08.viii.2008, 8 specimens,

1 macropterous, 7 apterous; 22.1x.2008, 4

specimens, all apterous, R.D. Kenner. In

addition, we collected this species on Van-

couver Island: Hamilton Marsh, 49°24’N

124°38’W, 11.viii.2004, R.D. Kenner and

K.M. Needham, 1 specimen, apterous.

Searches through the collections of the

SEM and Royal British Columbia Museum

(Victoria) add the following unpublished

records: Saanich, 48°33’N 123°22’W,

15.ix.1924, 30.ix.1925, W. Downes; Gali-

ano Island, north end, 49°0’N 123°34’W, 1

& 2.1x.1984, G.G.E. Scudder. Although the

records for M. mulsanti from the interior of

BC span the range from April to Septem-

ber, those from the coast are only from Au-

‘Cowan Vertebrate Museum, Department of Zoology, University of British Columbia, Vancouver, BC

V6T 1Z4

? Spencer Entomological Museum, Department of Zoology, University of British Columbia, Vancouver,

BC V6T 124

110 J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

gust and September. This species may not

be as rare as these few records suggest

since most specimens are apterous and may

easily be dismissed as ‘unidentifiable’ im-

mature bugs.

We thank G.G.E. Scudder for confirma-

tion of identifications and useful discus-

sions, and R.A. Cannings and C.R. Copley

for checking the Royal BC Museum collec-

tion for additional records.

REFERENCES

Kenner, R.D. and K.M. Needham. 2004. New waterboatman records for Western Canada (Hemiptera:

Corixidae). Journal of the Entomological Society of British Columbia 101: 57-58.

Maw, H.E.L., R.G. Foottit, K.G.A. Hamilton and G.G.E. Scudder. 2000. Checklist of the Hemiptera of

Canada and Alaska. NRC Research Press, Ottawa, Ontario, Canada.

Polhemus, J.T. 2008. Aquatic and semiaquatic Hemiptera. Chapter 15 in: An Introduction to the Aquatic

Insects of North America, 4" Edition (R.W. Merritt, K.W. Cummins and M.B. Berg, editors) pp. 385—

423.

Scudder, G.G.E. 1977. An annotated checklist of the aquatic and semiaquatic Hemiptera (Insecta) of Brit-

ish Columbia. Syesis 10: 31-38.

Scudder, G.G.E. 1986. Additional Heteroptera new to British Columbia. Journal of the Entomological

Society of British Columbia 83: 63-65.

J. ENTOMOL. Soc. BRIT. COLUMBIA 105, DECEMBER 2008

111

SCIENTIFIC NOTE

Podosesia syringae (Lepidoptera: Sesiidae):

a new clearwing moth record for British Columbia

VIRGILIU M. AURELIAN!, MARIO LANTHIER?

and GARY J.R. JUDD®

Podosesia syringae (Harris), commonly

known as the ash borer, is a clearwing moth

(Sesiidae) whose larvae are borers within

the trunks of lilac, Syringa vulgaris L., and

various ash species, Fraxinus spp. This

native North American insect (Eichlin and

Duckworth 1988) is considered a major

pest of wild, cultivated and ornamental ash

trees in eastern provinces and_ states

(Appleby 1973, Solomon 1983). In their

review of North American Sesiidae, Eichlin

and Duckworth (1988) reported collections

of P. syringae from Washington State and

eastern Alberta, but British Columbia (BC)

was excluded from their description of its

western range. Adults of this species occur

in two distinct colour morphs, a black

morph that has a dark brown abdomen, and

a yellow morph that has a light brown ab-

domen surrounded by yellow bands. The

two morphs are geographically distinct, the

yellow morph fraxini being restricted to

Western North America and the nominate

morph being restricted to Eastern North

America (Eichlin and Duckworth 1988). A

wide hybridization zone exists in the mid-

West and the prairies (Eichlin and

Duckworth 1988), suggesting the two ex-

treme morphs may be two different subspe-

cies. Future molecular research will be im-

portant in elucidating whether a subspecies

level is warranted for the ash borer.

In winter of 2006 one of us (ML) dis-

covered many unknown lepidopteran larvae

infesting various three-year-old Fraxinus

spp. nursery stock collected from commer-

cial tree nurseries located in Westbank and

Armstrong, BC. In early 2007 cut sections

of these infested ashes were brought into

the laboratory and the emerging adults were

identified by one of us (VMA) as the ash

borer, Podosesia syringae (Harris).

Voucher specimens from BC

(CNCLEP00041170 & CNCLEP0041171)

were deposited in the Canadian National

Collection of Insects, Arachnids and Nema-

todes, Ottawa, ON and confirmed as the ash

borer by J-F. Landry (personal communica-

tion).

In spring 2007 we surveyed wild and

ornamental lilacs surrounding nursery

plantings in Westbank, and found evidence

of larval feeding and pupal exuviae charac-

teristic of the ash borer. In June 2007,

Pherocon 1-CP style wing traps (PheroTech

Int. Inc., Delta, BC) baited with Clearwing

Borer lures SC L103 (Scentry Biologicals,

Billings, Montana,USA) containing (Z,Z)-3,

13 octadecadienyl acetate, a known clear-

wing male sex attractant and possibly the

major component of the female pheromone

for this species (Nielsen and Purrington

1978), were deployed in and around six ash

nursery plantings in Westbank and Arm-

strong, BC. In a total of 102 traps deployed

in both regions we captured 325 male P.

syringae adults. No black morphs were

captured in BC. Superficially, the yellow

morph resembles a paper wasp (Polistes

spp.) but on a sticky trap it could also be

confused with the Western poplar clear-

wing, Paranthrene robinae Hy. Edwards;

which is native to BC. However, on closer

examination P. syringae can be distin-

' Department of Biological Sciences, University of Alberta, CW315 Biological Sciences Building, Ed-

monton, AB, T6G 2E9

: CropHealth Advising and Research, P.O. Box 28098, Kelowna, BC, V1W 4A6

> Corresponding Author: Agriculture and Agri-Food Canada, Pacific Agri-Food Research Centre, Box

5000, 4200 Highway 97, Summerland, BC, VOH 1Z0. Email: Gary.Judd@agr.gc.ca

$ 175

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J. ENTOMOL. Soc. BRIT. COLUMBIA 105, DECEMBER 2008

Westbank

Armstrong — — —

100

2/7 9/7 16/7 23/7 30/7 6/8 13/8

Trap Checks (Day / Month)

Figure 1. Temporal pattern of seasonal catches of male P. syringae adults in sex-attractant

baited sticky traps at two locations within the Okanagan Valley, BC, in 2007.

guished from P. robinae by the presence of

a very long first metatarsal segment and

distinctive forewing venation.

Weekly trap checks revealed a male

flight period that lasted six weeks in West-

bank with peak flight occurring in the first

half of June (Fig. 1). Further north in Arm-

strong, the flight period was more extended,

ending in the first half of August, but the

smaller peak flight occurred about the same

time as it did in Westbank. In eastern North

America adult flight begins in April and

ends in July (Neal and Eichlin 1983). In

summer 2008 we redeployed sex-attractant

the fall of 2007. Our catches of P. syringae

confirmed the presence of this species in

the absence of ash, supporting the view that

local plantings of lilac on residential prop-

erties outside the nursery of initial discov-

ery are now a potential source of this insect.

From these results we conclude that P. sy-

ringae is established within parts of the

Okanagan Valley, BC. However, the origi-

nal source of infestation remains unknown.

We thank Larisa Aurelian for help col-

lecting biological specimens. This research

was partially supported with funds from a

BC Ministry of Agriculture and Lands Plant

traps around a fallow nursery field from Health Grant held by GJ.

which all ash trees had been harvested in

REFERENCES

Appleby, J.E. 1973. Observations on the life history and control of the lilac borer. Journal of Economic

Entomology 66: 248-249.

Eichlin, T.D. and W.D. Duckworth. 1988. Sesioidea: Sesiidae. Jn. R.B. Dominick. The moths of America

north of Mexico, fascicle 5.1. Wedge Entomological Research Foundation, Washington, D.C. USA.

Neal, J.W. and T.D. Eichlin 1983. Seasonal response of six male Sesiidae of woody ornamentals to clear-

wing borer (Lepidoptera: Sesiidae) lure. Environmental Entomology 12: 206-209.

Nielsen, D.G. and F.F. Purrington. 1978. Field attraction of male Podosesia syringae and P. aureocincta

to blends of synthetic compounds. Environmental Entomology 7: 708-710.

Solomon, J.D. 1983. Culturing the lilac borer Podosesia syringae, a serious pest of green ash (Fraxinus

pennsylvanica). Journal of the Georgia Entomological Society 18: 300-313.

J. ENTOMOL. Soc. BRIT. COLUMBIA 105, DECEMBER 2008

113

SCIENTIFIC NOTE

Survival of male click beetles, Agriotes obscurus L.,

(Coleoptera: Elateridae) during and after storage

at different temperatures

WILLEM G. VAN HERK', J. TODD KABALUK',

VIOLA W.M. LAM! and ROBERT S. VERNON!

The dusky wireworm, Agriotes obscurus

L. (Coleoptera: Elateridae) is a significant

pest of vegetable and field crops in the Fra-

ser Valley of BC (Vernon ef al. 2001).

Adults emerge from the soil from late

March through May and die soon after mat-

ing (males) or egg laying (Brian 1947),

restricting their availability for research and

necessitating storage methods that prolong

survival. Here we compare beetle survival

during and after storage at various tempera-

tures for various durations.

Male A. obscurus beetles were col-

lected at the Pacific Agri-Food Research

Centre (Agassiz, BC) during their peak

emergence (1° wk of May, 2005) and

placed in groups of 10 beetles into 150 ml

plastic containers with a freshly-cut apple

piece (approx. 2 x 2 x 0.5cm) placed on 70

g of moist sandy clay-loam soil. Containers

were put in growth chambers (15 per cham-

ber) (Controlled Environments Ltd., Winni-

peg, MB) set at 5, 8.5, 12, and 20 °C (4 0.1

°C), and dead beetles removed, and apple

pieces replaced, biweekly. Three containers

were removed from each chamber after 2,

4, 6, 8, and 10 wks to determine subsequent

survival at room temperature (RT) (21 +1 °

C), and beetles transferred to 10cm Petri

dishes (one per container) placed on 5cm

high racks inside Styrofoam boxes. Each

box (36 x 26 x 9cm deep) contained 2.5ml

water to prevent desiccation; a lcm gap

between the box and its lid permitted air

exchange. Beetle feeding and observation

continued (as above) for up to 12 wks.

Beetle mortality during storage was

highest at 20 °C, and similar at 5, 8.5, and

12° C for the first 6 wks but considerably

lower at 8.5 °C thereafter. Beetle mortality

was rapid within the first 2 wks of storage,

and increased with duration for all tempera-

tures except 8.5 °C (Table 1). Regression

(stepwise, backward; Proc REG, SAS 9.1;

SAS Institute 2002) of the proportion dead

per container during storage (m) to tem-

perature (t) and duration (d) yielded the

following: m = 0.003t? — 0.022t + 0.050d

(SE slopes: 0.0005, 0.012, 0.009 respec-

tively; P < 0.0001, 0.06, <0.0001 respec-

tively, d.f. = 3,57, adj. R’ = 0.88), indicat-

ing that survival increases as storage tem-

perature decreases.

For post-storage survival analysis, each

beetle was considered an experimental unit,

and storage temperatures were compared

with Kaplan-Meier survival analysis (Proc

LIFETEST, SAS 9.1), with strata duration

(Cox and Oakes 1984). The survival time of

50% of beetles (ST50) was subsequently

estimated by modelling survivorship for

each storage temperature-duration combina-

tion. Survivorship curves were compared

with log-rank tests; ST50 values were com-

pared using 95% confidence intervals.

Post-storage beetle mortality was rapid

regardless of previous storage temperature.

Beetles stored at 12 °C died more quickly at

RT than those stored at 8.5 or 5 °C (y” =

12.64, P = 0.0004; 7? = 19.27, P < 0.0001,

respectively). Beetles stored at 5 °C sur-

vived longest, but not significantly longer

than those stored at 8.5 °C (y° = 2.91, P =

0.09). Comparison of ST50 values and sur-

vival curves indicated that beetles survived

longer if stored at 5 °C than at 8.5 °C, if

Agriculture and Agri-Food Canada, Pacific Agri-Food Research Centre, P.O. Box 1000, Agassiz, British

Columbia, Canada VOM 1A0

114 J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

Table 1.

Mortality of Agriotes obscurus beetles after storage at various temperature for different dura-

tions, and subsequent survivorship at room temperature (RT). N = 30 per storage duration-

temperature combination. Post-storage survivorship at RT modelled with Kaplan-Meier sur-

vival curves; ST50 = time required for 50% survival.

S S Mean (SE) number of

torage torage heetles: dead (out of Survival of beetles at RT after storage (d)

temperature duration

(CC) (wk) 10) at the end of the STS50 Upper Lower

storage period 95% CI 95% CI

5 2 2.33 (0.67) 68.0 54.0 89.0

5 4 2.0 (0.58) 20.0 12.0 33.0

D 6 3.0 (1-53) 19-0 5.0 33.0

S 8 4.0 (2.08) 30.5 19.0 47.0

= 10 5.67 (0.33) 12.0 5.0 26.0

8.5 Z 2.0 (0.58) 27.0 26.0 47.0

8.5 + 2.0 (1.0) 26.0 14.0 47.0

8.5 6 3.67 (0.33) 19.0 14.0 33.0

8.5 8 1.33 (0.67) 19.0 12.0 21.0

8.5 10 2.67 (1.67) 12:0 7.0 33.0

12 Z 1.33 (0.88) 33.0 12.0 47.0

12 4 2.67 (0.33) 12.0 12.0 14.0

12 6 3.00 (0.58) 12.0 5.0 21.0

12 8 4.67 (2.19) 95 5.0 19.0

12 10 8.67 (0.88) n/a’ n/a’ n/a!

20 2 5.0 (1.15) 5.0 5.0 7.0

20 4 9.67 (0.33) n/a! n/a! n/a!

20 6,810 10.0 (0) n/a’ n/a’ n/a!

' Not enough beetles survived storage to permit analysis.

stored for 2 wks (y° = 15.48, P<0.0001) or 8

wks (7? = 6.15, P=0.013; Table 1), but not

when stored for 4 or 6 wks (P>0.05, Table

1). Surprisingly, beetles stored for 2 wks at

5, 8.5, or 12 °C survived longer at RT than

those stored at 20 °C (Table 1).

These results indicate that storage at

lower temperatures prolongs male click

beetle survival, and that storage at 8.5 °C

caused highest overall survival. Future re-

search should investigate how cold storage

conditions affects beetle physiology.

We thank Drs. D. Gray, C. Stevenson,

D. Raworth and R. Bennett, and sundry

reviewers for valuable advice.

REFERENCES

Brian, M.V. 1947. On the ecology of beetles of the genus Agriotes with special reference to A. obscurus.

Journal of Animal Ecology 16: 210-224.

Cox, D.R., and D. Oakes. 1984. Analysis of Survival Data. London: Chapman and Hall.

SAS Institute Inc. 2002. SAS/STAT user’s guide, Version 9.1. SAS Institute Inc. Cary, North Carolina.

Vernon, R.S., E. LaGasa, and H. Philip. 2001. Geographic and temporal distribution of Agriotes obscurus

and A. lineatus (Coleoptera: Elateridae) in British Columbia and Washington as determined by phero-

mone trap surveys. Journal of the Entomological Society of British Columbia 98: 257-265.

J. ENTOMOL. Soc. BRIT. COLUMBIA 105, DECEMBER 2008

115

SCIENTIFIC NOTE

Effect of handling and morbidity induction on weight,

recovery, and survival of the Pacific Coast wireworm,

Limonius canus (Coleoptera: Elateridae)

WILLEM G. VAN HERK’” and ROBERT S. VERNON’

Wireworms can recover from prolonged

morbidity induced by exposure to insecti-

cides (Vernon et al. 2008), but it is unclear

to what extent wireworm physiology and

survival are affected by repeated handling

or morbidity inductions, which reduces

larval weight and retards development

(Nicolas and Sillans 1989) in some insects.

It may be that differences in weights or

survival in wireworms repeatedly made

moribund, similarly handled but not made

moribund, or not subjected to handling or

morbidity might occur.

Larvae of the Pacific Coast wireworm,

Limonius canus LeConte, become tempo-

rarily moribund after contact with teflu-

thrin-treated wheat seeds, and recover more

quickly when re-exposed (van Herk and

Vernon 2007), but it is not known if contin-

ued re-exposure further decreases recovery

time. Wireworms repeatedly contact insec-

ticide-treated seeds in the soil, thus a con-

tinued decrease in the morbidity duration

may affect the insecticide’s efficacy in the

field. Here we discuss if wireworms con-

tinue to recover more quickly from teflu-

thrin-induced morbidity, and if this or han-

dling affects their weight and the time to

complete the larval instar.

Wireworms were collected from an or-

ganic farm in Kelowna, BC, in June 2007

and stored in soil at 15 °C. Late-instar, feed-

ing wireworms were randomly allocated to

one of three treatments (24—50 per treat-

ment), the ‘morbidity’, ‘handled’, and

‘control’ treatments. All observations were

made at 21+1 °C. In the ‘morbidity’ treat-

ment, morbidity was induced, and wire-

worm health assessed following van Herk

and Vernon (2007). Individual wireworms

were placed for 2 min in 1.5-ml Eppendorf

microcentrifuge tubes (Fisher Scientific)

with a single, ungerminated wheat seed (cv.

Superb) treated with Tefluthrin 20CS (20%

tefluthrin w/v) at 10 g AI/100 kg seed and

the fungicide Dividend XLRTA (3.21%

difenoconazole, 0.27% mefenoxam) at 13 g

AI/100 kg seed. Seeds were treated in 2007

by Syngenta Crop Protection Canada Inc.

(Portage la Prairie, MB). After exposure,

larvae were placed in individual Petri

dishes lined with moistened filter paper to

observe the onset of morbidity, and subse-

quently placed in separate, identical film

canisters filled with finely screened soil

with 20% moisture by weight. Wireworm

‘health’ was assessed every 10 min until 30

min after no further symptoms of morbidity

were observed (i.e. 7-15 times). Morbidity

was induced using this method at 24-h in-

tervals for 4 consecutive days.

Wireworms in the ‘handled’ treatment

were placed for 2 min in Eppendorf tubes

with a single untreated wheat seed and, as

above, observed for 20 min in a Petri dish,

and then transferred to film canisters filled

with soil. To expose these wireworms to

handling comparable to those in_ the

‘morbidity’ treatment, the health of

‘handled’ wireworms was assessed at 10-

min intervals 12 times on the first day and

10 times on the subsequent 3 d. Wireworms

in the ‘control’ treatment were placed in

film canisters and weighed, but were not

placed in Eppendorf tubes or Petri dishes.

Larvae were weighed individually for 5

consecutive d, approx. 6 h before wire-

worms in the ‘handled’ or ‘moribund’ treat-

' Agriculture and Agri-Food Canada, Pacific Agri-Food Research Centre, PO Box 1000, 6947 Highway 7, Agas-

siz, British Columbia, Canada

? Corresponding author: vanherkw@agr.gc.ca

116 J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

ments were placed in Eppendorf tubes the

first 4 d, and at the same time of day on day

five. Larvae were weighed for final time

approx. 2 mo later, at which time the pro-

portion moulted was recorded.

Wireworm recovery durations were

compared among treatments with ANOVA,

and the % decrease in recovery time be-

tween day | and days 2, 3 and 4 was calcu-

lated. Weight loss over time was compared

among treatments with repeated measures

ANOVA; orthogonal contrasts were used to

compare wireworm weight loss between

‘control’ and ‘handled’, and between

‘handled’ and ‘moribund’ treatments. Pro-

portions of larvae that moulted were com-

pared with Chi-square analysis.

In the morbidity treatment, the time to

recovery on day 1 was 88.8 + 2.3 (SEM)

min, similar to the 86.0 to 87.8 min re-

ported by van Herk and Vernon (2007).

Time to recovery on day 1 was significantly

longer than on days 2, 3 and 4 (72.4 + 1.8,

69.4 + 1.8, 71.8 + 1.8 min, respectively;

F=20.70, df=3,196, P<0.0001), but recov-

ery durations on the latter 3 d were not dif-

ferent (P>0.05). Recoveries on days 2, 3

and 4 were 18.5 — 21.8% more rapid than

the initial recovery time. These data suggest

that repeated contact with tefluthrin-treated

seeds in the field will not likely make wire-

worms insensitive to the insecticide.

Analysis of individual wireworm

weights over time using the first five meas-

urements indicated a weight loss over time

(F=8.35, df=4,384, P<0.0001, Table 1) and

an interaction between weight loss and

treatment (F=2.25, df=8, 384, P=0.02).

These remained significant (P<0.05) when

all six weighing days were included in

analyses; orthogonal contrasts indicated

greater weight loss in ‘handled’ than in

‘control’ treatments (F=3.04, df=5,480,

P=0.01), but not between ‘moribund’ and

‘handled’ treatments (F=0.80, df=5,480,

P=0.55). This suggests that weight loss in

the ‘moribund’ treatment was due to han-

dling alone, and that extensive handling of

wireworms may cause stress or damage,

perhaps including elevation of hemolymph

sugar or lipids (Woodring et al. 1989). It is

unlikely the observed weight loss was from

desiccation as it continued from days 5 to

68 when wireworms were continuously in

moist soil. Thus care must be taken to mini-

mize handling events and/or trauma.

All wireworms were alive on day 68

with no difference in moulting in the treat-

ments by day 68 (0.62, 0.52, 0.42, respec-

tively; ea = 2.79, df=2, P=0.25), so that re-

peated morbidity induction and handling

did not affect survival or development.

We thank M. Clodius and C. Harding

for technical assistance, S. Reid for permis-

sion to collect wireworms, and L. Letkeman

for treating wheat seeds.

Table 1.

Mean (SEM) weight (mg) of Limonius canus larvae subjected to one of three treatments:

‘Moribund’: repeated handling plus morbidity induced by insecticide exposure; ‘Handled’: repeated

handling only; ‘Control’: no handling or morbidity induction.

Treatment N day 1 day 2 day 3 day 4 day 5 day 68

Control 24: 25.2(1.2) 25:6(1.2) 2551.2). 255 02) -235.5(12)5 25.712)

Handled 25 26.001.1) 25.80.1) 25.80.41) 25.701.1) 25.6(1.1). -249(1.2)

Moribund 50 25.4(0.8) 25.5(0.8) 25.1(0.8) 25.1(0.8) 25.1(0.8) 24.7 (0.8)

REFERENCES

Nicolas, G., and D. Sillans. 1989. Immediate and latent effects of carbon dioxide on insects. Annual Review of

Entomology 34: 97-116.

van Herk, W.G. and R.S. Vernon. 2007. Morbidity and recovery of the Pacific Coast wireworm, Limonius canus,

following contact with tefluthrin-treated wheat seeds. Entomologia Experimentalis et Applicata 125: 111-

117.

Vernon, R.S., W. van Herk, J. Tolman, H. Ortiz Saavedra, M. Clodius, and B. Gage. 2008. Transitional sublethal

and lethal effects of insecticides after dermal exposures to five economic species of wireworms (Coleoptera:

Elateridae). Journal of Economic Entomology 101: 365-374.

Woodring, J. P., L. A. McBride, and P. Fields. 1989. The role of octopamine in handling and exercise-induced

hyperglycaemia and hyperlipaemia in Acheta domesticus. Journal of Insect Physiology 35: 613-617.

J. ENTOMOL. SOc. BRIT. COLUMBIA 105, DECEMBER 2008

117

Symposium Abstracts: Biodiversity in Stanley Park

Entomological Society of British Columbia

Annual General Meeting,

West End Community Centre Auditorium, Vancouver, BC,

October 4th, 2008

Parks, Protected Areas and Biodiversity

Conservation in B.C.

Geoffrey G.E. Scudder. Zoology Depart-

ment, University of British Columbia, Van-

couver, B.C.

Currently over 12% of the land area of Brit-

ish Columbia is categorized as Protected

Area. This includes national and provincial

parks, ecological reserves, and a few other

areas. However, it does not include regional

and municipal parks and lands owned by

various conservation organizations, al-

though these have definite conservation

value. An assessment of the Protected Ar-

eas network shows that this not only does

not serve as an efficient ecological network,

but it has unequal ecosystem representation,

and evidently does not adequately protect

the biodiversity richness and rarity hotspots

in the province. There are few buffer areas

and most parks and protected areas have not

been adequately assessed for biodiversity.

Impending climate change will undoubtedly

have a dramatic impact on the species and

ecosystems in British Columbia. There is an

urgent need to assess current elements of

biodiversity in all the parks and protected

areas, monitor these, and develop a land-

scape framework with appropriate connec-

tivity and integrity, that will facilitate the

movement of species in the future.

Some first systematic surveys of the in-

sects of Stanley Park in 2007 and 2008

John A. McLean. Department of Forest

Sciences, University of B. C., Vancouver, B.

The winter storms of 2006/07 felled a large

volume of large trees throughout Stanley

Park. Freshly fallen trees are quickly at-

tacked by ambrosia beetles and bark beetles

as part of nature’s first phase of recycling

the dying tissues. The records of some past

outbreaks of insects in Stanley Park will be

reviewed. In 2007, baseline surveys were

made of moths using light traps in undam-

aged forest stands near the Aquarium and

the Hollow Tree. Semiochemical-baited

multiple funnel traps were also set out to

evaluate bark beetle and ambrosia beetle

populations along with pitfall traps to

evaluate ground beetle populations. In

2008, the multiple funnel traps and pitfall

traps were set out in areas that had been

severely affected by the 2006/07 winter

storms after a large portion of the felled

trees had been removed and new seedlings

planted. A brief overview of some of our

results will be given.

A Fascination with Nature or "How big

can small get?"

Peter Woods.

Vancouver, B. C.

Imagine yourself let loose; able to explore

any and every facet of the natural history of

a favourite place. Give yourself ten years or

so. Where would this journey take you? I

have enjoyed just such an opportunity and

the rare privilege to observe, listen, touch,

imagine, and photograph nature in Stanley

Park. This presentation is a sampling of

what a naturalist does. It is a blending of

curlosity, wonder, fascination, observation,

fact, understanding, and shared discovery.

It is also about 'small things' and a world of

parallel universes. It is about what can hap-

pen only when you stop and stand still, in

one, magical place. It is about biodiversity

with inspiration drawn in equal measure

from ‘Winnie the Pooh's hundred acre

wood', and ‘Alice's Adventures in Wonder-

land'. Here are creatures that have taken me

on special journeys captured by a powerful

new tool, the 'eye' that sees through the lens

of a digital camera.

Naturalist-photographer,

118 J. ENTOMOL. SOC. BRIT. COLUMBIA 105, DECEMBER 2008

Challenges of Small Vertebrate Manage-

ment in Stanley Park

Robyn Worcester. Conservation Officer,

Stanley Park Ecology Society, Vancouver,

B.C.

Although Stanley Park is a fragmented

natural area in the heart of a large urban

setting, it is home to a diverse array of resi-

dent and migratory wildlife and presents

unique challenges as one of the oldest and

largest urban parks in North America. Fol-

lowing the massive windstorms in the Park

in December 2006, the small vertebrates of

Stanley Park became recognized as a com-

ponent of ecologically based forest manage-

ment and a renewed interest in their protec-

tion enabled formal research and invento-

ries to be undertaken, in many cases for the

first time. I will discuss the results of the

inventories that took place over the last two

years, the challenges that were faced

throughout the restoration process, and the

ongoing efforts of the Stanley Park Ecology

Society to maintain and enhance the diver-

sity of the small vertebrates in the Park

though data collection and public educa-

tion.

Stanley Park Restoration Project - Re-

spond/Plan/Restore

Jim Lowden. Director of Special Projects —

Vancouver Board of Parks and Recreation,

Vancouver, BC.

A look at managing a multiple objective

program to respond to a natural disaster;

balancing competing agendas while at-

tempting to do good. What we learnt from

the last 22 months in the trenches.

Hurricanes, invasive beetles, and urban

forests: lessons from Point Pleasant

Park.

Jon Sweeney. Natural Resources Canada,

Canadian Forest Service, Fredericton, NB.

Point Pleasant Park, much cherished and

heavily used by people (and their dogs) in

Halifax, Nova Scotia, has been hit by many

disturbances since its inception in 1866, not

least of which were the relatively recent

arrivals of the brown spruce longhorn bee-

tle and Hurricane Juan. Loss of much of the

mature red spruce to wind-throw and the

beetle has obviously altered forest age class

and stand structure in many areas of the

park; arthropod species composition and

diversity has likely also changed as a result,

but unfortunately our baseline knowledge is

limited. I will give a brief history of Point

Pleasant Park, focusing on changes in the

last 2-3 decades, highlight some of our re-

search on the biology and management of

the brown spruce longhorn beetle, and de-

scribe plans for park restoration in the after-

math of the Hurricane.

History of Survey and Control Activities

for Forest Pests in Stanley Park and Ad-

jacent Forest Environs

Leland Humble. Natural Resources Can-

ada, Canadian Forest Service, Victoria, B.

Since its opening in 1887, the 400 ha of

forest lands of Stanley Park have been sub-

jected to multiple abiotic and biotic distur-

bances that have impacted the health of its

forest stands. The extensive blowdown

caused by the 2006/07 winter storms is but

the most recent example of an abiotic dis-

turbance. Ninety-nine species of beetles,

122 species of moths and 11 species of

sawflies have been recorded from Stanley

Park and adjacent forest habitats on the

north shore by the Forest Insect and Disease

Survey during survey activities between

1949 and 1995. Only a few of these species

are damaging. Outbreaks of defoliators

such as the western hemlock looper, the

green-striped forest looper, and the western

blackheaded budworm have caused exten-

sive defoliation or mortality within areas of

the park. Aerial control operations to pro-

tect the forest resources of the park have

been undertaken three times against native

defoliators and once for an introduced defo-

liator. The biology and damage caused by

the major forest pests present in Stanley

Park and the history of the control opera-

tions undertaken in the park are reviewed.

NOTICE TO CONTRIBUTORS

The JESBC is published once per year in December. Manuscripts dealing with all facets

of the study of arthropods will be considered for publication provided the content is of

regional origin, interest, or application. Authors need not be members of the Society.

Manuscripts are peer-reviewed, a process that takes about 6 weeks.

Submissions. The JESBC accepts only electronic submissions. Submit the manuscript in

MS Word or Wordperfect format, along with a cover letter, as an e-mail attachment to the

Editor. Manuscripts should be 12-point font, double-spaced with generous margins and

numbered lines and pages. Tables should be on separate, numbered pages. Figure cap-

tions should be placed together at the end of the manuscript. Each figure should be sub-

mitted as a separate electronic file. Figure lines should be sufficiently thick and lettering

sufficiently large so that they are clear when reduced to fit on the Journal page, 12.7 x

20.5 cm. Preferred graphic formats are GIF and JPG at least 1500 pixels wide. Do not

send raw (uncompressed) files such as TIFF or BMP.

Submission deadline for Vol. 106 is September 1, 2009. Submit contributions to:

Dr. Hugh Barclay, Editor hbarclay@nrcan.gc.ca

Pacific Forestry Centre

506 West Burnside Road Phone (250) 363-3338

Victoria, BC V8Z 1M5 Fax (250) 363-0775

Style and format. Consult the current volume for style and format. Style generally con-

forms to the Entomological Society of America Style Guide, available at http://

www.entsoc.org/pubs/publish/style/index.htm. Pay particular attention to the formats of

References Cited. If there is no precedent, consult Scientific Style and Format: The CBE

Manual for Authors, Editors, and Publishers, 6" Ed., published by the Council of Biology

Editors.

Scientific Notes are an acceptable format for short reports. They must be two Journal

pages maximum, about 4.5 manuscript pages. Scientific Notes do not use traditional sec-

tion headings, and the term "Scientific Note" precedes the title. A short abstract may be

included if desired. Notes are peer-reviewed just as regular submissions are.

Page charges. The Society has no support apart from subscriptions. The page charge for

articles is $35 and includes all tables and figures except coloured illustrations. The page

charge is $40 if none of the authors is a member of the ESBC. A surcharge for coloured

illustrations applies.

Page charge waiver. Authors whose entomological activities are not supported by uni-

versities or official institutions, who are unable to pay page charges from their personal

funds, may apply for assistance when submitting a manuscript.

Electronic reprints. The Society provides authors with an Adobe Acrobat file of an ex-

act copy of the paper as it appears in the Journal. This file will also be posted on the

ESBC Website (http://www.sfu.ca/biology/esbc/) to provide free electronic access to our

Journal for all interested parties.

Back issues. Back issues of many volumes of the Journal are available at $15 each.

Membership in the Society is open to anyone with an interest in entomology. Dues are

$20 per year; $10 for students. Members receive the Journal, Boreus (the Newsletter of

the Society), and when published, Occasional Papers.

Address inquiries to:

Dr. Lorraine Maclauchlan, Secretary — lorraine.maclauchlan@gov.bc.ca

B.C. Ministry of Forests

515 Columbia St Phone (250) 828-4179

Kamloops, BC V2C 2T7 Fax (250) 828-4154

iii

! entomelngiat Manes vs weit Columbia

G.G.E. Scudder. New provincial and state records for Heteroptera (Hemiptera) in Canada and the

United States

Jeremy R. Dewaard, B. Christian Schmidt, Gary G. Anweiler and Leland M. Humble. First Ca-

nadian records of Lampropteryx suffumata ({Denis & Schiffermiller], 1775) (Geometridae:

Larentiinae) 9

D. Thomas Lowery, Michael G. Bernardy, Robyn M. Deyoung and Chris J. French. Identifi-

cation of new aphid vector species of Blueberry scorch virus

John K. Mackenzie, Peter J. Landolt and Richard S. Zack. Sex attraction in Polistes dominulus

(Christ) demonstrated using olfactometers and morphological source extracts

Gary J.R. Judd and Mark G.T. Gardiner. Efficacy of Isomate-CM/LR for management of leafrol-

lers by mating disruption in organic apple orchards of western Canada

D.A. Raworth. Climate change and potential selection for non-diapausing two-spotted spider

mites on strawberry in southwestern British Columbia

Michael D. Doerr, Jay F. Brunner and Timothy J. Smith. Biology and management of bark

beetles (Coleoptera: Curculionidae) in Washington cherry orchards

Kevin Durden, John J. Brown and Maciej A. Pszczolkowski. Extracts of Ginkgo biloba or Ar-

temisia species reduce feeding by neonates of codling moth, Cydia pomonella (Lepidoptera:

Tortricidae), on apple in a laboratory bioassay

S.D. Cockfield and E.H. Beers. Management of dandelion to supplement control of western

flower thrips (Thysanoptera: Thripidae) in apple orchards

Luis Martinez-Rocha, Elizabeth H. Beers and John E. Dunley. Effect of pesticides on inte-

grated mite management in Washington State

SCIENTIFIC NOTES

R.D. Kenner and K.M. Needham. Additional records for semiaquatic

in southwestern British Columbia

Virgiliu M. Aurelian, Mario Lanthier and Gary J.R. Judd. Podosesia syringae (Lepidoptera: Se-

siidae): a new clearwing moth record for British Columbia

Willem G. Van Herk, J. Todd Kabaluk, Viola W.M. Lam and Robert S. Vernon. Survival of male

click beetles, Agriotes obscurus L., (Coleoptera: Elateridae) during and after storage at differ-

ent temperatures

W.G. Van Herk and R.S. Vernon. Effect of handling and morbidity induction on weight,

recovery, and survival of Pacific Coast wireworm, Limonius canus (Coleoptera: Elateridae)

ABSTRACTS

Symposium Abstracts: Biodiversity in Stanley Park Entomological Society of British Columbia

Annual General Meeting, West End Community Centre Auditortum, Vancouver, BC, October

4th, 2008

NOTICE TO CONTRIBUTORS