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Columbia

COVER: Aeshna interrupta Walker (Odonata: Aeshnidae)

Aeshna interrupta (Variable Darner) is one of the most common large dragonflies

in British Columbia. It is a boreal species, ranging across North America from Newfound-

land to Alaska and south in the western mountains to California and New Mexico. It lives

in marshes and peatlands and is the typical Aeshna of grassland ponds and lakes in the

Interior. On the Coast it is one of the predominant dragonflies in peat bogs.

The scientific name “interrupta” refers to the shape of the stripes on the sides of

the thorax. In eastern North America and on the Pacific coast, these are “interrupted”, that

is, each is broken into two spots. On the Great Plains and in the BC Interior, the stripes are

unbroken but thinner than in any other species. The common name “variable” describes

these stripes. The stripes and spots of the male are blue; those of the female are blue or,

more commonly, yellow.

Most dragonflies spend the majority of their lives in the aquatic larval (nymphal)

stage. After about 10 to 14 moults, depending on the species and environmental condi-

tions, the fully grown larva metamorphoses into an adult inside its last larval skin, then

crawls out of the water. Now exposed to air, the dragonfly begins its final moult — the top

of the thorax splits open and the adult squeezes out. It pumps blood into its wings and

abdomen, which expand slowly, and gradually the body hardens. After an hour or two the

dragonfly can fly, but only weakly at first. It leaves the empty larval skin, the exuvia,

clinging to the support.

Photograph details:

Male Aeshna interrupta photographed during emergence at a grassland pond near

Riske Creek, Chilcotin region, BC, 15 June 1978. Pentax Spotmatic IT with 50 mm/1.4

Macro Takumar lens, handheld and with available light. Kodachrome 64 film. Robert A.

Cannings.

The Journal of the Entomological Society of British Columbia is

published annually in December by the Society

Copyright© 2010 by the Entomological Society of British Columbia

Designed and typeset by Hugh Barclay and Jen Perry.

Printed by FotoPrint Ltd., Victoria, B.C.

Printed on Recycled Paper.

Pug

J. ENTOMOL. SOC. BRIT. COLUMBIA 107, DECEMBER 2010 l

Journal of the

Entomological Society of British Columbia

Volume 107 Issued December 2010 ISSN #007 1-0733

Directors of the Entomological Society of British Columbia, 2010-2011 «2.00.0... 2

G.G.E. Scudder. Melacoryphus admirabilis (Uhler) (Hemiptera: Lygaeidae) new

to Canada, with additional Canadian provincial records for other Heteroptera

K.G.A. Hamilton and Yong Jung Kwon. Taxonomic changes in Dicraneura

Hardy, Colladonus Ball and Macrosteles Fieber (Homoptera-

Auchenorrhyncha) in the Montane Cordilleran Ecozone ...................0000000000 Ld

L.M. Humble, E. John, J. Smith, G.M.G. Zilahi-Balogh, T. Kimoto and M.K.

Noseworthy. First records of the banded elm bark beetle, Scolytus schevyrewi

Semenov (Coleoptera: Curculionidae: Scolytinae), in British Columbia ....... 21

Jeremy R. deWaard, Leland M. Humble and B. Christian Schmidt. DNA barcod-

ing identifies the first North American records of the Eurasian moth, Eupithe-

cia pusillata (Lepidoptera: Geometridae)

Robert A. Cannings, Marc A. Branham and Robert H. McVickar. The fireflies

(Coleoptera: Lampyridae) of British Columbia, with special emphasis on the

light-flashing species and their distribution, status and biology 33

Richard S. Winder, Donna E. Macey and Joe Cortese. Dominant bacteria associ-

ated with broods of mountain pine beetle, Dendroctonus ponderosae

(Coleoptera: Curculiomidae, SCOlVtMAG). .c:...c00ccsssssssecnssssesvnecesesssrenessesessseoness 43

George D. Hoffman and Sujaya Rao. Cereal leaf beetle, Oulema melanopus (L.)

(Coleoptera: Chrysomelidae), attraction to oat plantings of different ages ....57

Alan L. Knight. Effect of sex pheromone and kairomone lures on catches of cod-

ling moth (Lepidoptera: Tortricidae) on clear interception traps ................... 67

Alberto Pantoja, Aaron M. Hagerty, Susan Y. Emmert, Joseph C. Kuhl, Keith

Pike, Juan M. Alvarezand Andrew Jensen. Aphids (Homoptera: Aphididae)

associated with rhubarb in the Matanuska Valley, Alaska: species composition,

Seasonal abundance, and potential Virus VECtOLS. ..........0.essresscccesssacerccsesetene 75

NOTES

G.G.E. Scudder. New distribution records for United States Lygaeoidea

(PAC UHM ticle LC TOPE A) ca 2h cle ct su sohe asked has odeduesehsidiesuassuodesssaeenntsnaseecdersnonodes 83

G.G.E. Scudder. The Schizopteridae (Hemiptera), a family new to Canada ...... 85

G.E. Haas, N. Wilson, J.R. Kucera, T. O. Osborne, J.S. Whitman and W.N. John-

son. Range expansion and hosts of Ctenophthalmus pseudagyrtes Baker

(Siphonaptera: Ctenophthalmidae) in central Alaska

NOPIGE FO'CONTRIBUTORS ic ioe occ edevelesscce Inside Back Cover

bho

J. ENTOMOL. SOC. BRIT. COLUMBIA 107, DECEMBER 2010

DIRECTORS OF THE ENTOMOLOGICAL SOCIETY

OF BRITISH COLUMBIA FOR 2010-11

President:

Rob McGregor

Douglas College

President-Elect:

Ward Strong

B.C. Ministry of Forests and Range

Past-President:

Tom Lowery

Agriculture and Agri-Food Canada

Treasurer:

Lorraine Maclauchlan

B.C. Ministry of Forests and Range

Secretary:

Leo Rankin

B.C. Ministry of Forests and Range

Directors, first term:

Art Stock, Tracey Hueppelsheuser, Dezene Huber

Directors, second term:

Naomi Delury, Jim Corrigan,

Graduate Student Representative (1° year)

Chandra Moffatt

Regional Director of National Society:

Bill Riel

Canadian Forest Service, Victoria

Editor, Boreus:

Jennifer Heron

Jennifer. Heron@gov.be.ca

Jeremy DeWaard

Editor of Web Site:

Bill Riel

briel@pfc.forestry.ca

Honorary Auditor:

Sheila Fitzpatrick

Agriculture and Agri-Food Canada

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

Editorial Committee, Journal:

Editor-in-Chief (Re tiring): Subject Editors:

Hugh Barclay Sheila Fitzpatrick (Agriculture)

Victoria, B.C. Lorraine MacLauchlan (Forestry)

hbarclay@shaw.ca Rob Cannings (Systematics/Morphology)

Technical Editor: Jen Perry Editor Emeritus: Peter Belton

J. ENTOMOL. SOC. BRIT. COLUMBIA 107, DECEMBER 2010

Melacoryphus admirabilis (Uhler) (Hemiptera: Lygaeidae)

new to Canada, with additional Canadian

provincial records for other Heteroptera

G.G.E. SCUDDER!

ABSTRACT

The lygaeid Melacoryphus admirabilis (Uhler) is recorded from Saskatchewan and new

to Canada. New provincial records are given for 16 other species of Heteroptera, be-

longing to the families Alydidae, Artheneidae, Cymidae, Geocoridae, Lygaeidae, Miri-

dae, Oxycarenidae, Rhyparochromidae and Tingidae.

INTRODUCTION

Further research on collections of Cana-

dian Heteroptera has resulted in the discov-

ery of another species new to Canada. In

addition, new provincial records have been

established for 16 other species. Some of

these significantly change the known distri-

bution of species in Canada.

Data cited are those on specimen labels.

The order follows Maw et al. (2000). Col-

lection and Museum abbreviations used in

the text are as follows:

CNC: Canadian National Collection of

Insects, Agriculture and Agri-Food Canada,

Ottawa, ON (R.G. Foottit).

DBUC: Department of Biological Sci-

ences, University of Calgary, Calgary, AB

(J.E. Swann).

LC: D.J. Larson Private collection, Ma-

ple Creek, SK.

LM: Lyman Entomological Museum,

Macdonald College, McGill University,

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

UCCB: Department of Biology, Univer-

sity College of Cape Breton, Sydney, NS

(D. McCorquodale).

NEW RECORDS

Family MIRIDAE

Phytocoris eureka Bliven

In Canada, previously only reported

from British Columbia (Stonedahl 1988;

Maw et al. 2000), but widely distributed in

the western United States (Stonedahl 1988).

New provincial record. AB: 19,

Kananaskis, U. of C. Field Station, 51°

O'49"N 114°12'01"W, 11-19.vii.2004 (L.

Wooldridge) [DBUC].

Family TINGIDAE

Corythucha distincta Osborn & Drake

In Canada, previously only reported

from British Columbia (Parshley 1919;

Downes 1925, 1927; MacNay 1952; Maw

et al. 2000), but in the United States re-

corded south to California and in South

Dakota (Froeschner 1988b).

New provincial record. AB: 1¢, Fish

Creek Provincial Park, 50°54.406'N 114°

01.260'W, sweep of field near ranch house,

17.vii.2009 (J.E. Swann & G. Hull)

[DBUC]; 14, Fish Creek Provincial Park,

50°54.594'N 114°01.698'W, sweep,

17.vii.2009 (J.E. Swann & G. Hull)

[DBUC]; 14, Waterton Lakes National Pk.,

Cardston Entrance, Malaise, 13.vii1.1989

(R. Longair) [DBUC].

Corythucha salicata Gibson

In Canada, reported from British Co-

lumbia, Manitoba, Northwest Territories,

Ontario and Saskatchewan (Maw et al.

' Beaty Biodiversity Centre and Department of Zoology, University of British Columbia, 6270 University

Boulevard, Vancouver, BC V6T 1Z4

4 J. ENTOMOL. Soc. BRIT. COLUMBIA 107, DECEMBER 2010

2000). This species is confined to the West-

ern Cordilleran region in the United States

(Froeschner 1988b).

New provincial record. AB: 17¢ 69,

Fish Creek Provincial Park, 50°54.594'N

114°01.698'W, sweep, 17.viti.2009 (J.E.

Swann & G. Hull) [DBUC]; 2¢ Fish Creek

Provincial Park, 50°55.717'N 114°

07.307'W, sweep, goldenrod, 19.vi1i.2009

(G. Hull) [DBUC].

Family ALYDIDAE

Protenor belfragei Haglund

In Canada, previously reported from

Saskatchewan east to Prince Edward Island

(Maw et al. 2000), and widely distributed in

the United States (Froeschner 1988a).

New provincial record. NS: 1¢, Cape

Breton Co., Sydney, UCCB, 1.x1.1999 (MLI.

Kerr) [UCCB]; 19, id., 1.ix.1999 (B.H.W.

MacIntosh) [UCCB].

Family ARTHENEIDAE

Chilacis typhae (Perris)

An alien species in Canada previously

reported from British Columbia and Ontario

(Scudder 2000; Maw et al. 2000; Scudder

and Foottit 2006), as well as New Bruns-

wick, Nova Scotia, Prince Edward Island

(Wheeler 2002) and Quebec (Roch 2008).

In the United States it is recorded from both

the east (Wheeler and Fetter 1987) and the

west (Wheeler and Stoops 1999). Wheeler

(2002) also added 21 new U:S. state records

for C. typhae.

New provincial record. AB: 12, Cal-

gary, U. of Calgary, 11.vi.2009 (Tim Loh)

[DBUC]. SK: 19, Battle Creek, near Merry

Flats, 30.ix.2006 (D. Larson) [LC]; 2,

Larson Ranch, Hwy. 21, 16 km S. Maple

Creek, 24.1x.2008 (D. Larson) [CNC; LC];

2¢ 19, id., 2.viii.2009 (D. Larson) [CNC;

LC].

Family CYMIDAE

Cymus coriacipennis (Stal)

In Canada, previously only reported

from British Columbia (Scudder 1961;

Maw et al. 2000). It is a Western Cordil-

leran species in the United States (Hamid

1975; Ashlock and Slater 1988).

New provincial record. SK: 14, Maple

Ck., Hwy. 21, 16 km S, 8.vii.2003 (D. Lar-

son) [LC]; 1¢, id., 8.vi.2004 (D. Larson)

[CNC].

Family GEOCORIDAE

Geocoris atricolor Montandon

In Canada, previously only reported

from British Columbia (Parshley 1919;

Walley 1934; Maw et al.2000) and Alberta

(Walley 1934, Strickland 1953; Maw et al.

2000). A Western Cordilleran species in the

United States (Ashlock and Slater 1988).

New provincial record. SK: 14, Cypress

Hills Pk., Center Block, Highland Trail,

2.x.2008. .(D.. Larson) [LC]; 193;larson

Ranch, Hwy. 21, 16 km S. Maple Creek,

12.vii.2002 (D:. Larson) {LC Tee

17.ix.2008 (D. Larson) [LC]; 23, 19, id.,

24.ix.2008 (D. Larson) [CNC; LC]; 19,

Maple Creek, 6 km N, 9 km E, sandy road

allowance, 12.1x.2008 (D. Larson) [LC].

Geocoris howardi Montandon

In Canada, previously reported from

Alberta, British Columbia, Northwest Terri-

tories and Yukon (Maw et al. 2000). Also

known from Alaska, the species is distrib-

uted across boreal North America (Readio

and Sweet 1982; Ashlock and Slater 1988).

New provincial records. MB: 19, Car-

berry, 29.vi1.1953 (Brooks-Kelton) [CNC];

12, Churchill, 31.vii.1937 (W.J. Brown)

[CNC]; 16 39, id.,. 10.vin.1937 ye:

Brown) [CNC]; 19, id., 12.vii.1952 (J.G.

Chillcott) [CNC]; 2¢ 12 1 immature,

Churchill, 4 km W, Akudik marsh, 58°

44'47"N 94°06'47"W, gen’l._ coll.,

16.v11.2006 (Boreal & Arctic Entomol)

[UM]; 1 immature, Churchill, 6 km E No.

Stud. Ctr., 58°46'14"N 93°54'46"W, krum-

holz tundra, gen’l. coll. (39), 11.vi11.2006

(Boreal & Arctic Entomol) [UM]; 1 1mma-

ture, Churchill, 12 km W, Launch Rd., 58°

45'18"N 93°59'04"W, bluffs + fen near A-

frame, gen’l. coll. (84), 17.vi11.2006 (Boreal

& Arctic Entomol) [UM]; 1 immature,

Churchill, 15 km S, Stud. Ctr., 58°37'00"N

93°49'15"W, gen’l. coll. near pond, Sample

16, 9.vi1.2006 (Boreal & Arctic Entomol)

[UM]; 1¢ 22 1 immature, Churchill, 15 km

S No. Stud. Ctr., 58°37'00"N 93°49'15"W,

burned area-boreal for., gen’l. coll. (62),

14.vi1i.2006 (Bor. & Arc. Entomol.) [UM];

12, Turtle Mt., 28.vii.1953 (Brooks-

Kelton) [CNC]. NS: 19, S. Berwick,

J. ENTOMOL. Soc. BRIT. COLUMBIA 107, DECEMBER 2010

22.viii.1963 (V.R. Vickery) [LM]. ON: 1d

32, Black Hawk, 3.viii.1960 (Kelton &

Whitney) fTENC]; 19; ‘Eagle Raver,

11.viii.1960 (Kelton & Whitney) [CNC];

14, Hastings Co., 29.viii.1954 (J.F. Brim-

ley) [CNC]; 39, Kapuskasing, 18.vii.1961

(G. Brumpton) [CNC]; 5¢, Little Current,

9.vii.1961 (G. Brumpton) [CNC]; 1, Nes-

torville, 24.vi.1965 (K.P. Butler) [LM]; 29,

One Sided Lake, 1.vili.1960 (Kelton &

Whitney) [CNC]; 39, id., 2.viii.1960

(Kelton & Whitney) [CNC]; 19,Sioux Nar-

rows, 6.vili.1960 (Kelton & Whitney)

HONG). °2OC 2-24) IG: “Kazubazua,

18.viii.1931 (G.S. Walley)[CNC]; 19, id.,

25:vii: 1933" (G.S. Walley) [CNC]; 1¢,

Laniel, 21.vii.1932 (W.J. Brown) [CNC].

SK: 1¢ 19, Christopher Lake, 13.vii.1959

Gi&” J Brooks): = (CNC "22,- -id.,

15.vii.1959) (A. & J. Brooks) [CNC]; 10,

Cypress Hills, E. Block, Ambrose Place, 25

km SE Maple Creek, 10.v11.2006 (D. Lar-

son) [LC]; 2¢ 19, Cypress Hills Pk., Cen-

ter Block, Highland Trail, 2.x.2008 (D.

Larson) [LC]; 19, Saskatoon, 22.vii.1949

(L. Konotopetz) [CNC]; 192, Torch River,

23.vill. 1950 (L.A. Konotopetz) [CNC].

Geocoris pallens Stal

In Canada, previously only reported

from Alberta, British Columbia (Forbes

1900; Torre-Bueno 1946; Slater 1964; Ash-

lock and Slater 1988; Maw et al. 2000). G.

pallens has been collected from most of the

western United States, and has a range ex-

tending eastward to Indiana, Illinois, Mis-

sour! and Arkansas (Readio and Sweet

1982), but also occurs from Mexico to Cen-

tral America and in Hawaii (Ashlock and

Slater 1988).

New provincial record. SK: 14, Larson

Rch., 16 km S Maple Creek, Hwy. 21,

5.vi1i.2002 (D. Larson) [CNC].

Geocoris uliginosus (Say)

In Canada, reported from Newfoundland

(Lindberg1958), Ontario (Walley 1934) and

Quebec (Walley 1934; Moore 1944, 1950;

Béique and Robert 1963; Larochelle 1984;

Roch 2008). G. uliginosus var. speculator

Montandon was synonymized with G. u/igi-

nosus by Readio and Sweet (1982).

Early records for British Columbia

(Downes 1927), repeated by Walley (1934)

and noted by Lindberg (1958), are incor-

rect. Downes (1927) reported the species

from Merritt on August 11, 1923 by R.

Hopping, and also at Victoria. | have been

unable to locate the Merritt material taken

by Hopping on that date, but other speci-

mens now in the CNC collected in 1923 by

R. Hopping have been found to be G. bulla-

tus (Say). Also, specimens in the CNC from

Victoria, 5.1x.1923 (K.F. Auden) also prove

to be G. bullatus.

In North America, in general G. uligino-

sus has a range that extends from the Gulf

Coast north to southern Canada, and from

the east coast west to the foothills of the

Rocky Mountains in Colorado (Readio and

Sweet 1982; Ashlock and Slater 1988).

Also known from Cuba (Alayo 1973) and

the West Indies (Baranowski and Slater

2005).

New provincial record. NB: 1¢, Fundy

Nat. Pk., 8.vii1.1954 (J.F. Brimley) [CNC].

Family LYGAEIDAE

Subfamily LYGAEINAE

Melacoryphus admirabilis (Uhler)

This species and genus is keyed by Sla-

ter (1992) and is macropterous with a black

membrane with a narrow white margin, and

with clavus and corium black for most part,

with costal margin and apical third of co-

rium red. The species is widely distributed

in the United States and occurs in Mexico

(Ashlock & Slater 1988).

New Canadian record. SK: 19°, Grass-

lands Natl. Pk., Tp. 1 & 2, Rge. 6 & 7, W3,

EBGE sweep samples, 29-30.vi1.2008

LGNC]}.

Subfamily ORSILLINAE

Nysius angustatus Uhler

Widely distributed across Canada, and

previously reported from the Northwest

Territories east to New Brunswick (Maw et

al. 2000), and known from Mexico and

most of the United States (Ashlock and

Slater 1988).

New provincial record. NS: 23 29,

Coldbrook, 22.viii.1963 (V.R. Vickery)

[LM].

Nysius niger Baker

In Canada, previously reported from

6 J. ENTOMOL. SOc. BRIT. COLUMBIA 107, DECEMBER 2010

Yukon and the Northwest Territories to

Newfoundland (Maw et al. 2000), but not

previously recorded form Nova Scotia and

Prince Edward Island. Recorded from most

of the United States, Bermuda, Mexico to

Central America, and the West Indies

(Ashlock and Slater 1988), although not

noted from the latter by Baranowski and

Slater (2005).

New provincial record. NS: 39, Kent-

ville, 15-17.vii.1966 (L.A. Kelton) [CNC].

Nysius tenellus Baker

In Canada, previously only reported

from British Columbia (Barber 1947; Maw

et al. 2000). However, the species is re-

corded from Florida, most of the western

United States, Mexico, Central America

and the West Indies (Barber 1947; Ashlock

and Slater 1988; Baranowski and Slater

2005).

New provincial record. SK: 14, Jones

Peak, 9 km W, Eastend, 49°30'N 108°57'W,

14.vi1.2005 (Larson) [CNC].

Family OXYCARENIDAE

Crophius bohemani (Stal)

In Canada, previously only reported

from British Columbia (Downes 1927;

Walley 1934; Barber 1938; Maw et al.

2000). The record for the “North West Ter-

ritories” (Gibson 1911) is obviously an er-

ror, and was not included by Barber (1938).

In the United States C. bohemani is a West-

ern Cordilleran species.

New provincial record. SK: 1419, Cy-

press Hills Park, Center Bloc, Highland

Trail, 25.1%.2008: (D.* Larson): [CNC LC];

12, Cypress Hills, Center Block, War

Lodge Coulee, 29.1x.2008 (D. Larson)

[LC].

Family RHYPAROCHROMIDAE

Subfamily RHYPAROCHROMINAE

Tribe DRYMINI

Eremocoris ferus (Say)

In Canada, previously reported from

British Columbia, Nova Scotia, Ontario and

Quebec (Maw et al. 2000). Sweet (1977)

elevated Eremocoris borealis (Dallas) from

synonymy with E. ferus and stated that this

species has a Carolinian and Austroriparian

Zone distribution extending from the Gulf

of Mexico in the eastern United States north

to lowland locations in New England, with

the northern records being from Massachu-

setts, New Hampshire, Connecticut and

southern New York. He also noted that

specimens from Illinois, lowa and Indiana

were all referable to E. ferus, and appeared

to mark the northern limit of the distribu-

tion of the species in the Midwest. Further-

more, Sweet (1977) stated that he had not

seen FE. ferus specimens from west of the

100° meridian, although relictual popula-

tions in Texas might indicate that the spe-

cies extends west of this meridian.

Over the past few years I have studied

collections of Eremocoris from the west,

and have specimens that I consider to be E.

ferus from not only British Columbia and

Saskatchewan, but also from Arizona, Cali-

fornia, Idaho, Montana, Oregon, Utah and

Washington state.

New provincial record. SK: 19, Cypress

Hills Pk., C. Block, Ski Lodge, 25.vi.2004

(D. Larson) [CNC]; 19, Larson Ranch,

Hwy. 21, 16 km S. Maple Creek, 16.v.2009

(D. Larson) [LC].

Tribe MEGALONOTINI

Megalonotus sabulicola (Thomson)

An alien species, in Canada previously

reported from British Columbia (Scudder

1960, 1961; Asquith and Lattin 1991), On-

tario (Maw et al. 2000) and Quebec

(Scudder and Foottit 2006). Asquith and

Lattin (1991) discussed the occurrence of

this species in the Pacific Northwest, with

records in the United States shown for Cali-

fornia, Idaho, Oregon, Utah and Washing-

ton. Wheeler (1989) also discussed the oc-

currence of M. sabulicola in the eastern

United States, with new records for Dela-

ware, New Jersey, Pennsylvania, Rhode

Island, Virginia and West Virginia, and

additional localities in Maryland and New

York. I have also collected M. sabulicola in

Montana (Scudder 2010). Wheeler (1989)

found that in the mid-Atlantic region of the

United States, this bug feeds mainly on the

fallen seeds of the spotted knapweed, Cen-

taurea biebersteinii DC (= C. maculosa

auct. non Lam).

New provincial record. SK: 1, Larson

Ranch, Hwy. 21, 16 km S Maple Creek,

J. ENTOMOL. SOc. BRIT. COLUMBIA 107, DECEMBER 2010

10'v: 2008.” Larson) [LC]240" 29> z.,

flooded grass, 12.vi.208 [CNC; LC].

Tribe MYODOCHINI

Neopamera albocincta (Barber)

To date in Canada, only reported from

Ontario (Scudder 1985; Maw et al. 2000).

Recorded from most of the eastern United

States to Texas in the south, as well as the

West Indies, and Mexico to South America

(Ashlock and Slater 1988).

New provincial record. QC: 19, Riv. Du

sud'Co., Iberville; ‘CH322, 31.vi. 1975 (N.

Dorion) “fCNC]; 164; - id., ‘CH500,

26.viii.1974 (N. Dorion) [CNC]; 19, id.,

CH503, 26.viii.1975 (N. Dorion) [CNC].

Sisamnes claviger (Uhler)

So far only recorded from British Co-

lumbia in Canada (Scudder 1985, 1992,

1993, 1994; Maw et al. 2000), but widely

distributed in the United States (Ashlock

and Slater 1988).

New provincial record. SK: 22, Cypress

Hills, Center Block, War Lodge Coulee,

2971x2008 °(D: Larson) [CNC; LC]; 29,

Cypress Hills Pk., Center Block, Highland

Trail, 14.1v.2009 (D. Larson) [CNC; LC];

32, Larson Ranch, Hwy. 21, 16 k S Maple

Creek, 10.v:2008 (D. Larson) [CNC; LC];

1GlQ, id) 24.ix:2008 (D:. Larson) [LC];

14, Sand Hills, 7 km W Piapot, 4.vii.2009

(DD: Karson) [EC].

Tribe STYGNOCORINI

Stvgnocoris rusticus (Fallén)

In Canada, this alien is recorded from

British Columbia east to Newfoundland

(Maw et al. 2000), but until now there have

been no records for either Manitoba or Sas-

katchewan.

Asquith and Lattin (1991) mapped the

restricted distribution in the Pacific North-

west, with records shown for Oregon and

Washington. Wheeler (1983) reviewed the

more extensive distribution in the eastern

United States.

New provincial record. SK: 1, Cypress

Hills, Center Bock, War Lodge Coulee,

29.ix.2009 (D. Larson) [LC]; 14, Cypress

Hills Park, Center Block, Highland Trail,

25.ix.2008, (D. Larson) [CNC]; 1d,

Frenchman R. Valley, Cypress L.,

20.viii.2009, (D. Larson) [LC]; 14, Larson

Ranch, Hwy. 21, 16 km S Maple Creek,

23.ix.2008, (D. Larson) [LC]; 23 19, id.,

24.1x.2008 (D. Larson) [CNC; LC].

Tribe UDEOCORINI

Neosuris castanea (Baker)

In Canada, to date reported only from

British Columbia (Scudder 1993, 1994;

Maw et al. 2000). Recorded from the West-

ern Cordilleran states, to the south and

Mexico (Ashlock and Slater 1988).

New provincial record. SK: 19, Maple

Ck., Hwy. 21, 16 km S, 30.v11.2003 (D.

Larson) [LC]; 14, Old-Man-on-his-Back

Ridge, 49°11'N 109°16'W, 5.1x.2009 (D.

Larson) [CNC].

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 col-

lections for permission to examine the ma-

terial in their care and/or the loan of speci-

mens.

I am especially indebted to Dr. D. Lar-

son (Maple Creek, SK) and Dr. J.E. Swann

(DBUC) for their cooperation in this study

and for allowing me to include records from

their respective provinces. I thank Launi

Lucas for the final preparation of the manu-

script.

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J. ENTOMOL. Soc. BRIT. COLUMBIA 107, DECEMBER 2010

J. ENTOMOL. SOC. BRIT. COLUMBIA 107, DECEMBER 2010

Taxonomic changes in Dicraneura Hardy, Colladonus Ball

and Macrosteles Fieber (Homoptera-Auchenorrhyncha)

in the Montane Cordilleran Ecozone

K.G.A. HAMILTON! and YONG JUNG KWON’

ABSTRACT

A neotype is proposed for Typhlocyba carneola Stal, 1858 (=Dikraneura carneola var.

shoshone DeLong & Caldwell, 1937, syn.nov.), and Dikraneura sitkana Ball & DeLong

is elevated to specific rank for the taxon previously known as Dikraneura carneola. Five

other species are described from the Montane Cordilleran Ecozone: Colladonus keltoni

Hamilton, Colladonus okanaganus Hamilton, Macrosteles frigidus Kwon, Macrosteles

similis Kwon and Macrosteles vulgaris Kwon.

INTRODUCTION

Eleven apparently undescribed species

were discovered during the preparation of a

faunal synopsis of the Montane Cordillera

Ecozone (MCE) of British Columbia. Six of

these species belong to well studied genera.

Colladonus Ball was revised by Nielson

(1957) for the species north of Mexico and

6 species have been added subsequently

(Nielson 1962, Hamilton and Langor 1987);

this contribution adds two more. Dikra-

neura Hardy was revised by Knight (1968)

and a former synonym is elevated to spe-

cific status following designation of a neo-

type for the species with which it has been

confused. Macrosteles Fieber is a large

genus discussed by Kwon (1988, unpub-

lished); his dissection techniques and five

of his new species from the Atlantic Mari-

time Ecozone will appear elsewhere

(Hamilton and Kwon 2010) and three oth-

ers, occurring in the MCE, are formally

described below.

Four other species belong to the typical

subgenus Empoasca Walsh, which is a

taxon with more than 600 species; it is in

need of revision before any additional spe-

cies are described. A fifth species is known

only from a single female. Since it belongs

to the Delphacid genus Delphacodes Fie-

ber, in which only males can be recognized

with certainty, no formal description can be

presented.

Specimens and notes examined in this

study are deposited in the following institu-

tions:

AMNH: American Museum of Natural

History, New York.

CNC: Canadian National Collection of

Insects, Arachnids and Nematodes, Ottawa,

ON

GL: Grassland Leafhopper survey; field

notes from H.H. Ross in CNC.

MLBM: Monte L. Bean Life Science

Museum, Brigham Young University,

Provo, UT

NCSU: North Carolina State University,

Raleigh

OrSU: Oregon State University, Corval-

lis

Colladonus keltoni Hamilton, sp.n.

Diagnosis. Crown slightly less than half

as long as wide, apically rounded (Fig. 1B-

E); colour uniform brown, paler on teg-

mina, unmarked except for crown of head:

apex with 2 black spots and usually also a

mustache-shaped mark between eyes; teg-

mina hyaline with dark hind wing veins

visible (Fig. 1A). Male genitalic characters

'Biodiversity Theme, Agriculture and Agri-food Canada, Central Experimental Farm, Ottawa, Ontario,

K1A 0C6, Canada

"Dept. of Agricultural Biology, College of Agriculture & Life Sciences, Kyungpook National University,

Daegu 702-701, Korea

12 J. ENTOMOL. Soc. BRIT. COLUMBIA 107, DECEMBER 2010

TSA

- Sitinn ae

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PP OE ARE Se

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Figures 1-2. Colladonus spp. 1, C. keltoni; 2, C. okanaganus: A, habitus, dorsolateral aspect;

B-C, dorsum of male; D-E, dorsum of female. Habitus to larger scale (1 mm), dorsum to

smaller scale (0.5 mm).

as in C. ponderosus Ball (Nielson 1957,

figs. 4A, C): male pygofer produced on

caudoventral margin and bearing a short

spine at tip; style tips curved outwards, but

with gonopore at midlength of shaft, and

terminal processes extending at least half

length of shaft, as in C. fahotus Ball

(Nielson 1957, fig. 47B). Length: male, 4.5-

4.9 mm; female, 4.8-5.2 mm.

Types. Holotype male, B.C.- Cran-

brook, 23 July 1959 (L.A. Kelton). Para-

types: 3 females, same data as holotype; 3

females, Yahk, 22 July 1959 (L.A. Kelton),

lodgepole [pine]; 1 female, Penticton, 27

June 1974 (M.W. Nielson); 1 male,

Okanagan Mission, 18 June-2 July 1971,

sticky board on Prunus emarginata; | male,

same data, 1-16 July 1971; 1 male, Bear

Creek (Okanagan Valley), 8 Aug. 1970

(K.G.A. Hamilton); 1 female, Okanagan

Falls, 4 July 1976 (K.G.A. Hamilton); 1

female, Little Fort, 3 July 1976 (K.G.A.

Hamilton); 2 males, Elko, 14 Aug. 1985

(K.G.A. Hamilton); MT- 1 male, 5 km E

Grantsdale, 3 June 1992 (K.G.A. Hamil-

ton). All types (7 males, 8 females) No.

21835 in CNC.

Remarks. All females and the darkest

males may be distinguished by the head

markings. This is one of three species of

Colladonus that are associated with pines,

all of which have strongly produced male

pygofers. The combination of genitalic

characters distinguishes males from the

other two pine species, C. ponderosus and

C. tahotus.

Colladonus okanaganus Hamilton, sp.n.

Diagnosis. Crown slightly less than half

as long as wide, apically rounded (Figs.

2B); head and venter yellow, strongly con-

trasting with blackish brown notum and

tegmina; costa and small spots on preapical

cells hyaline (Fig. 2A). Male genitalic char-

acters as in C. flavocapitatus (Van Duzee):

pygofer spine arising from midlength of

truncate pygofer tip, styles tips long and

curved outwards, and terminal processes of

aedeagus extending to midlength of shaft

(Nielson 1957, figs. 45 A-C), but with proc-

ess of pygofer slightly longer, as long as

terminal processes of aedeagus, and

gonopore basad of midlength of shaft (as in

Nielson 1957, fig. 19B). Length: male, 4.6-

5.0 (holotype); female unknown.

Types. Holotype male, B.C.- Okanagan

Mission site 12 (Okanagan Valley), 30 July

J. ENTOMOL. SOc. BRIT. COLUMBIA 107, DECEMBER 2010

1974 (J.E.H.). Paratypes: 1 male, Arm-

strong, 15-29 July 1971, sticky board on

Prunus emarginata; | male, Penticton, site

1 SB, 18 July 1974 (J.E.H.). All types No.

21835 in CNC.

Remarks. The boldly contrasting colour

is similar only to that of an undescribed

species of Colladonus from the west coast

of Vancouver Island.

Dikraneura carneola (Stal)

Typhlocyba carneola Stal, 1858: 196

(Sitka Island, Alaska).

Dikraneura carneola_ var. shoshone

DeLong & Caldwell, 1937: 27, syn.nov.

(Idaho).

Diagnosis. This species is closely allied

to a sister species, formerly considered as a

“variety” sitkana (name elevated to specific

rank, below). The latter differs in minor

details of male genitalia and has a more

southerly range (Figs. 3-4), overlapping that

of D. carneola in southern B.C. to Idaho.

Remarks. This taxon was described

from unusually pinkish specimens. Ball and

DeLong (1925) considered the species to be

very widespread in western North America,

with variable colour. Specimens from Utah

were especially yellower and were named

as “var. sitkana,” possibly as geographic

variants, but more probably these are eco-

phenotypes. Later, DeLong & Caldwell

(1937) figured the male genitalia for the

first time. They considered the genitalic

characters to be variable and associated the

widespread “D. carneola” with a different

aedeagal type than that of “var. shoshone”

from Idaho. Knight (1968) concluded that

these two aedeagal types represent separate

species. Lack of material from Alaska

prompted him to retain the name “D.

carneola” for the widespread species. How-

ever, 78 specimens were later taken close to

Sitka at Haines (1 nymph, 12 males, 16

females, GL 1142) and Potter (14 males, 35

females, GL 1127). Five males dissected

from each series are referable to D. sho-

shone. A female from much farther north-

west on coastal Alaska (King Salmon) is

probably conspecific. Three males from

inland sites in northwestern Canada (Banff,

AB; Atlin and Chicotin, BC) indicate that

the coastal populations have a montane

connection to the southern BC and ID

populations. The two names are therefore

synonyms and the southern “variety” is

indeed a valid species.

Types. Stal’s types have been sought,

but have not been found (Knight 1968).

Neotype of carneola, here designated:

male, AK- Haines, 5 August 1968 (Ross,

Ross & Miller) GL 1142.

Dikraneura sitkana (Ball & DeLong),

stat.nov.

Typhlocyba carneola var. sitkana Ball &

DeLong, 1925: 330.

Typhlocyba_ carneola:

(misidentification).

Diagnosis. Characters and distribution

as in Knight (1968) for D. carneola [nec

Stal], but the single female of “carneola”

from Edmonton, AB cannot be positively

identified. This female probably belongs to

the common prairie-inhabiting D. variata

Hardy, which sometimes has pinkish fe-

males.

Material examined: 52 males, 66 fe-

males from B.C.- Baldy Mtn. 7000-

7550' [2300-2500 m ASL], 13 Aug. 1970

(K.G.A. Hamilton); 48 males, 48 females,

MT- Bozeman, 9 Sept 1971 (H.H. Ross)

[GL] 1271. All types No. 21835 in CNC.

Ten males from each of these series were

dissected to verify that there is no admix-

ture of other species.

Macrosteles frigidus Kwon, sp.nov.

Diagnosis. Yellow to yellowish green,

often with faint smoky tint on tegmina (Fig.

SA-B). Crown with anterior margin

rounded in male, more or less pointed in

female; anterior black spots isolated, sub-

equal in size to posterior spots, which are

always prominent and isolated; median and

lateral spots absent; frons without any

prominent dark streaks (Fig. 5C-E). Teg-

mina more or less mottled with large, dis-

coid pale areas. Male abdomen with 1°

acrotergite only as broad as long, 2™ acro-

tergite with “neck” shorter than half of

acrotergite width; 1° tergal apodeme reach-

ing posteriorly to two-thirds of tergite

length (Fig. 5F); 1° sternal apodemes with

posterior lobes slightly longer (1.1-1.2 *)

Knight, 1968:

14 J. ENTOMOL. SOC. BRIT. COLUMBIA 107, DECEMBER 2010

Figures 3-4. Ranges of Dikraneura spp. 3, D. carneola; 4, D. sitkana.

than wide (Fig. 5G, J-L); 2" sternal apode-

mes with posterior lobes slightly exceeding

twice basal width (Fig. SN-Q); apophysis

processes narrowly produced (Fig. 5M).

Aedeagal shaft in posterior aspect (Fig.

S5W-Z) with apical processes reflexed ba-

sally, crossed, extending nearly to middle

of shaft, and subapex narrowed, bearing

paired lateral teeth slightly above gonopore;

in lateral aspect (Fig. 5S-V) more or less

convex subapically on dorsal side, spiculate

ventrobasally; subgenital plates (Fig. 5R)

each with mesal margin about as long as

basal margin. Length: male, 2.9-3.2 mm;

female, 3.2-3.4 mm.

Types. Holotype male, B.C.- Quesnel,

31 Aug. 1948 (G.J. Spencer). Paratypes: 5

females, same data as holotype; 2 males, 4

females, Minnie Lake, 27 July 1925 (H.G.

Crawford); 3 males, 4 females, Stanley, 3

Aug. 1949 (R. Stace-Smith); 2 males, |

female, 7 mi S of Nelson, 6 Aug. 1969 (P.

Oman); Alta.- 3 males, 4 females, Wain-

wright, 27 July 1957 (A.R. & J.E. Brooks);

! male, High Prairie, 17 July 1961 (AR:

Brooks); 1 male, Grande Prairie, 26 July

1961 (A.R. Brooks); Sask.- 1 male, Torch

R.; 20. July 1950: (L.A... Konotopets); 4

male, 2 females, Prince Albert, 23 July

1959 (A.R. & J.E. Brooks); 1 male, Saska-

toon, 1 Aug. 1960 (A.R. Brooks); Man.- 2

males, 2 females; Virden, 9 July 1953

(Brooks & Kelton); Y.T.- 2 males, Snag, 24

July 1948 (Mason & Hughes); AK.- 3

males, 1 female, Big Delta, 13 July 1951

(J.R. McGillis). Holotype and 41 paratypes

J. ENTOMOL. Soc. BRIT. COLUMBIA 107, DECEMBER 2010 15

FER iN ars Pape oar KISS Riga? cb

| ee Nd! anh i tg he

‘ ns ents

SPs tae

or,

Figure 5. Macrosteles frigidus, based on specimens from B.C., except variants (from Yukon,

Alta. and AK respectively): A, habitus of male, dorsal aspect; B, same, of female; C, male head

in facial aspect; D, same, in anterior aspect; E, same, of female; F, male abdominal tergites 1-2

in dorsal aspect; G, male abdominal sternite | in dorso-anterior aspect; H, same, in anterior

aspect; I, same, in lateral aspect; J-L, same, variation of outlines of sternal lobes; M, male ab-

dominal sternite 2 in anterior aspect; N, same, in dorsal aspect; O-Q, same, variation of out-

lines of sternal lobes; R, male subgenital plate, ventral aspect; S-V, variation in aedeagus, lat-

eral aspect; W-Z, same, posterior aspect. Habitus (A-B) to largest scale; face (C-E) to second

largest scale; abdominal plates (F-R) to second smallest scale; aedeagus (S-Z) to smallest scale.

16 J. ENTOMOL. Soc. BRIT. COLUMBIA 107, DECEMBER 2010

No. 22874 in CNC; 3 paratypes in OrSU

and | paratype in NCSU.

Additional specimens, excluded from

the type series, were examined from Texas

(36 specimens), Alberta and Saskatchewan

(3 specimens). The Texan morph has some-

what shorter head and apodemes, while the

others have more strongly pointed heads,

especially in the female.

Remarks. This species is similar to MV.

galeae Hamilton (in Hamilton and Langor

1987), but is readily distinguished by the

very bold coronal spots between the eyes.

From all other species in the genus with

crossed aedeagal processes it may be distin-

guished by its convex subapex of the

aedeagus in lateral aspect.

Macrosteles similis Kwon, sp.nov.

Diagnosis. Relatively elongate; yellow

to yellowish green, often with very faint

smoky tint on fore wings (Fig. 6A-B).

Crown somewhat rounded in male, more or

less pointed in female; black anterior spots

often fused together with lateral spots to

form a transverse band along coronal mar-

gin, and with median and lateral spots often

confluent; frons with prominent, dark trans-

verse bands on either side of median stripe

(Fig. 6C-E). Tegmina with dark stripe along

claval suture. Male abdomen with 1" acro-

tergite broad, 2"° acrotergite nearly trans-

verse, triangularly produced ventrally,

“neck” slender, less than half as long as

width of acrotergite; 2"’ tergal apodeme

often reaching to middle of tergite (Fig.

6F); 1° sternal apodemes with posterior

lobes usually as long as wide (Fig. 6G, J-L),

slightly inclined in lateral aspect (Fig. 61);

2™ sternal apodemes with posterior lobes

about twice as long as basal width (Fig.6N-

Q); apophysis processes narrowly produced

(Fig. 6M). Aedeagal shaft smooth ventrally,

with serrate lateral flanges lying along ante-

rior edge of shaft in lateral aspect, in poste-

rior aspect (Fig. 6V-X) narrowly developed,

apical processes slender, convergent, or

crossed apically in posterior aspect, in lat-

eral aspect (Fig. 6S-U) turned anteriad, then

hooked dorsad; subgenital plates each with

mesal margin longer than basal margin

(Fig. 6R). Length: male, 3.6-4.0 mm; fe-

male, 4.0-4.4 mm.

Types. Holotype male, AK- Big Delta,

13 July 1951 (J.R. McGillis). Paratypes: 1

female, same data as holotype; | male, |

female, same locality, 30 June-26 July 1951

(W. Mason); | male, Fairbanks, 4 Aug.

1951 (H.C. Severin); 2 males;Circle) Hot

Spgs., 4 Aug. 1951 (H.C. Severin); B.C.- 3

males, Atlin 2200' [700m], 29-30 July 1955

(B.A. Gibbard); 1 male, 10 mi S of Rev-

elstoke, 22 Aug. 1978 (K.G.A. Hamilton)

on Juncus spp.; 1 male, Orchard Pt. bog,

Brooks Pen., Vancouver Is., 4 Aug. 1981

(R.A. & S.G. Cannings); N.W.T.- 2 males,

4 females, Rocknest Lake 65°39'N

114°20'W, 26 Aug. 1966 (G.E. Shewell);

Qué.- 5 males, Natashquan, 7 Aug. 1929

(W.J. Brown); CO- 1 male, 2 mi S of

Gould, 9000' [2400m], 13 Aug. 1968 (P.

Oman); UT- 16 males and 38 females,

Uinta Mts. 10,000' [3000m], Uintah Co., 21

Aug. 1983 (M.W. Nielson) on Carex sp.

Holotype and 22 paratypes No. 22876 in

CNC; 54 paratypes in MLBM; | paratype

in OrSU.

Remarks. Similar to MM fieberi

(Edwards), but differing in the much shorter

posterior lobes on the male 2” sternal apo-

demes, and by the serrate lateral flanges of

the aedeagus lying along the anterior edge

of the shaft instead of along the sides

(Beirne 1952, figs. 91-92). This aedeagal

character appears to be variable in Euro-

pean populations of M. fieberi, but not in

North American populations.

Macrosteles vulgaris Kwon, sp.nov.

Diagnosis. Yellow to yellowish green,

frequently with smoky markings on body

and tegmina (Fig. 7A-C). Crown broad,

slightly more than twice as wide as long,

often rounded in male, more or less pointed

in female; spot pattern confluent or isolated,

as in M. quadrilineatus (Forbes). Frons

with black transverse bands usually conflu-

ent (Fig. 7D-G). Tegmina unmarked or

faintly crossbanded. Male abdomen with 1°

acrotergite very broad, 2™ acrotergite

nearly transverse, triangularly produced

ventrally, “neck” slender and elongate, only

slightly shorter than acrotergite; 2" tergal

apodeme reaching middle of tergite (Fig.

J. ENTOMOL. Soc. BRIT. COLUMBIA 107, DECEMBER 2010 17

S ae:

Figure 6. Macrosteles similis, based on males from Qué. and female from N.W.T.: A, habitus of

male, dorsal aspect; B, same, of female; C, male head in facial aspect; D, same, in anterior aspect; E,

same, of female; F, male abdominal tergites 1-2 in dorsal aspect; G, male abdominal sternite | in

dorso-anterior aspect; H, same, in anterior aspect; I, same, in lateral aspect; J-L, same, variation of

outlines of sternal lobes; M, male abdominal sternite 2 in anterior aspect; N, same, in dorsal aspect;

O-Q, same, variation of outlines of sternal lobes; R, male subgenital plate, ventral aspect; S-U, varia-

tion in aedeagus, lateral aspect; V-X, same, posterior aspect. Habitus (A-B) to largest scale; face (C-

E) to second largest scale; abdominal plates (F-R) to second smallest scale; aedeagus (S-X) to small-

est scale.

18 J. ENTOMOL. SOC. BRIT. COLUMBIA 107, DECEMBER 2010

ho wt

Figure 7. Macrosteles vulgaris, based on specimens from B.C.: A, B, habitus of male, dorsal aspect;

C, same, of female; D, male head in facial aspect; E, same, of female in anterior aspect; F, G, same, of

male; H, male abdominal tergites 1-2 in dorsal aspect; I, male abdominal sternite 1 in dorso-anterior

aspect; J, same, in anterior aspect; K, same, in lateral aspect; L, M, same, variation of outlines of ster-

nal lobes; N, male abdominal sternite 2 in anterior aspect; O, same, in dorsal aspect; P, male subgeni-

tal plate, ventral aspect; Q-S, variation in aedeagus, lateral aspect; T-V, same, posterior aspect. Habi-

tus (A-C) to largest scale; face (D-G) to second largest scale; abdominal plates (H-P) to second small-

est scale; aedeagus (Q-V) to smallest scale.

J. ENTOMOL. Soc. BRIT. COLUMBIA 107, DECEMBER 2010

7H); 1° sternal apodemes with posterior

lobes about as long as basal width (Fig. 71,

L-M); apophyses abruptly bent at middle in

anterior aspect (Fig. 7J), dorsal aspect with

lower part narrower than median part be-

tween posterior lobes; 2" sternal apodemes

apically truncate (Fig. 7N) with posterior

lobes reduced, triangular (Fig. 7O).

Aedeagus (Fig. 7Q-V) as in M. quadrilinea-

tus; Subgenital plates each with mesal mar-

gin longer than basal margin (Fig. 7P).

Length: male, 3.2-3.7 mm; female, 3.5-4.2

mm.

Types. Holotype male, B.C.- Cowichan

Lake, 6 June 1955 (R. Coyles). Paratypes:

65 males, 30 females, same data as holo-

type; 8 males, Kootenay Bay, 23 June-29

Aug. 1948-49 (D.B. Waddell); 1 male,

Soda Creek, 21 July 1950 (G.J. Spencer); 1

male, 3 females, Malahat, 20 Sept. 1950

(W. Downes); 22 males, 11 females, Oliver

2500' [800m], 2 July 1953 (J.R. McGillis);

1 male, Diamond Head Trail

3200' [1000m], Squamish, 7 Aug. 1953

(G.J. Spencer); 1 male, 6 females, Duncan,

9 June 1955 (R. Coyles); 1 male, 5 females,

Miracle Is. Park, 11 June 1955 (R. Coyles);

3 males, 3 females, same locality, 29 May

1959 (R. Madge & R.E. Leech); | male,

Spectacle Lake, Oliver, 10 June 1959 (L.A.

Kelton); 11 males, 14 females, Terrace, 15

July-3 Aug. 1960 (W.R. Richards); | male,

Shames, 18 mi SW of Terrace, 17 July

1960 (C.H. Mann); 8 males, 11 females, 5

mi E of Sidney, 23 Aug. 1971 (J. Saw-

bridge); Man.- | male, Shoal Lk., 28 June

1976 (K.G.A. Hamilton) on Distichlis

stricta; Sask.- 1 male, Pipestone Creek, 7

June 1958 (A.R. Brooks); CA- 3 males,

Truckee, Nevada Co., 29 Aug. 1967 (L.

Kelton); 1 male, | female, 38 mi SE Mt.

Shasta, 10 July 1972 (P. Oman); | male, 4

females, 18 mi W_ of Susanville

5400' [1530m], 10 July 1972 (P. Oman); 1

male, 7.6 mi N of Bridgeport, 19 June 1982

(P. Oman); 1 male, 2 females, Squaw Val-

ley. Placer -Co..5 Oct. 1983 (DG.

Denning); ID- | male, Paris, 8 July 1920 (F

4741); MT- 1 male, 24 females, Bozeman,

9 Sept. 1971 (H.H. Ross); OR- 6 males, 5

females, Corvallis, 14 Aug. 1928 (O.A.

Lg

Hills); 1 female, same data, but 4 July

1927; 1 male, Lostine, 12 Aug. 1929 (O.A.

Hills); 10 males, Gresham, 8 July 1949 (R.

Rosenstiel) on strawberry; 1 male, Astoria,

13 June 1951 (E.A. Dickason); 1 male, Ore-

gon City, 28 Aug. 1962 (Koontz) on potato;

6 males, 3 females, Forest Grove, 2 July

1965 (F.P. Larson) black light trap; 3

males, Woodburn at Pudding R., 26 July

1965 (F.P. Larson) black light trap; 3

males, same data, but 11 Sept. 1966; |

male, Hillsboro, 28 July 1965 (F.P. Larson)

black light trap; 2 males, Troutdale, 18 July

1966 (F.P. Larson) black light trap; 2

males, 2 females, Canby, 29 Aug. 1966

(F.P. Larson) black light trap in corn; 3

males, Lily Lake 13 mi E of French Glen

7200' [2000m], 10 July 1968 (P. Oman); 3

males, MacDonald Forest at Corvallis, 17

July 1968 (P. Oman); 2 males, 6 females,

Seal Rock, 31 Aug. 1968 (P. Oman); 8

males, 13 females, same data, but 1 May

1970; 4 males, 5 females, same data, but I-

27 May; | male, 2 mi NW of Banks, 18

June 1969 (P. Oman); 1 male, 25 mi SE of

Joseph, 10 Aug. 1969 (P. Oman); 7 males,

5 females, Tou Velle Park, Jackson Co., 2

May 1970 (P. Oman); 1 male, 3 females,

Agate Desert, Jackson Co., 2 May 1970 (P.

Oman); 1 male, 2 females, same data, but

19 May 1971; 1 male, 2 females, Joseph, 6

June 1970 (P. Oman); 1 male, 10 mi ESE of

Ruch, 14 May 1971 (P. Oman); 10 males,

11 females, 19 mi W of Klamath Falls, 24

June 1971 (P. Oman); 1 male, 20 mi E of

Seneca, 14 Aug. 1971 (P. Oman); 3 males,

1 female, 12 mi W _ of Silver Lake

5000' [1500m], Lake Co., 12 July 1978 (P.

Oman); 11 males, 4 females, Johnson

Meadow, Klamath Co., 17 July 1979 (P.

Oman); | male, 2 females, north edge of

Big Lake 4650' [1350m], Linn Co., | Oct.

1979 (P. Oman); 2 males, 2 females, Saun-

ders Lake 2 mi N of Hauser, 8 Oct. 1979 (P.

Oman); WA- 2 mi S of Humptulips, 22

Aug. 1971 (Viraktamath). Holotype and

240 paratypes No. 22877 in CNC; 190

paratypes in OrSU, 13 paratypes in MLBM,

3 in University of Kentucky, Lexington and

1 in AMNH.

Additional records, based on unassoci-

20 J. ENTOMOL. SOC. BRIT. COLUMBIA 107, DECEMBER 2010

ated females and therefore excluded from

the type series, are: CA- Fort Ord., 2 mi E

of Gasquet; OR- 2 mi E of Carlton, Merlin,

Mt. Vernon, Odell, Sams Valley (N_ of

Medford), Skookum Meadow (Klamath

(05):

Remarks. Distinguished from M. quad-

rilineatus and related species by the angu-

late apophyses of the male 2™ sternal apo-

deme, and by the unusual length of the 2"

acrotergite “neck.” This species appears to

replace M. quadrilineatus as the most com-

mon temperate-zone Macrosteles west of

the Rocky Mountains.

REFERENCES

Ball, E.D. and D.M. DeLong. 1925. The genus Dikraneura and its allies in North America. Annals of the

Entomological Society of America 18: 324-340.

Beirne, B.P. 1952. The Nearctic species of Macrosteles (Homoptera: Cicadellidae). The Canadian Ento-

mologist 84: 208-232.

DeLong, D.M. and J.S. Caldwell. 1937. The genus Dikraneura — a study of the male genitalia — with de-

scriptions of new species. Proceedings of the Entomological Society of Washington 39: 17-33.

Hamilton, K.G.A and Y.J. Kwon. 2010. Taxonomic supplement to “short-horned” bugs (Homoptera-

Auchenorrhyncha). Pp. 420-431 in D. MacAlpine (Ed.), Assessment of Species Diversity in the Atlantic

Maritime Ecozone. NRC Press Biodiversity Monograph Series.

Hamilton, K.G.A. and D.W. Langor. 1987. Leafhopper fauna of Newfoundland and Cape Breton Islands

(Rhynchota: Homoptera: Cicadellidae). The Canadian Entomologist 119: 663-695.

Knight, W.J. 1968. A revision of the Holarctic genus Dikraneura (Homoptera: Cicadellidae). Bulletin of the

British Museum of Natural History (Entomology) 21: 102-201.

Kwon, Y.J. 1988. Taxonomic revision of the leafhopper genus Macrosteles Fieber of the world

(Homoptera: Cicadellidae). Dissertation, University of Wales College of Cardiff, U.K. 557 pp.

Nielson, M.W. 1957. A revision of the genus Co//adonus (Homoptera, Cicadellidae). U.S. Department of

Agriculture Technical Bulletin 1156. 52 pp. + 11 plates.

Nielson, M.W. 1962. New species of leafhoppers in the genus Co//adonus (Homoptera, Cicadellidae). An-

nals of the Entomological Society of America 55: 143-147.

J. ENTOMOL. SOc. BRIT. COLUMBIA 107, DECEMBER 2010

First records of the banded elm bark beetle,

Scolytus schevyrewi Semenov (Coleoptera:

Curculionidae: Scolytinae), in British Columbia

L.M. HUMBLE!, E. JOHN!, J. SMITH’, G.M.G. ZILAHI-BALOGH’,

T. KIMOTO?’ and M. K. NOSEWORTHY'!

ABSTRACT

The banded elm bark beetle, Scolytus schevyrewi Semenov was detected for the first

time in British Columbia near Kelowna during 2010. Fifty-eight S. schevyrewi were

captured in an experiment that targeted the European elm bark beetle, Scolytus

multistriatus (Marsham). It was a test of the efficacy of a new trap design relative to the

multiple funnel trap currently used in surveillance programs for invasive bark- and

wood-boring Coleoptera. Data on the seasonal occurrence of the banded elm bark beetle

are presented.

Key Words: banded elm bark beetle, Scolytus schevyrewi, forest pest, invasive species

INTRODUCTION

The banded elm bark beetle, Scolytus

schevyrewi Semenov (Coleoptera: Curculi-

onidae: Scolytinae), is an invasive bark

beetle native to central and eastern Asia

(Negron et al. 2005; CABI/EPPO 2009; Lee

et al. 2009). It was first reported from North

America in Colorado and Utah in 2003

(Negron et al. 2005) and was soon found to

be more widely distributed (Negron et al.

2005). In the U.S.A., S. schevyrewi is now

reported from 28 states including all states

west of the Mississippi River (except Ar-

kansas, lowa, Louisiana and North Dakota)

as well as from Connecticut, Delaware,

Illinois, Indiana, Maryland, Michigan, Min-

nestoa, Missouri, New Jersey, Ohio, Penn-

sylvania and Virginia, (Lee et al. 2009;

NAPIS 2010). Specimens in reference col-

lections indicate that S. schevyrewi was

present in the U.S.A. (Colorado) as early as

1994 (Lee et al. 2009). In Canada, banded

elm bark beetle was first detected in Alberta

in 2006 (Langor et al. 2009) and has subse-

quently been reported from locations in

Saskatchewan, Manitoba and Ontario

(CABI/EPPO 2009). While species of elms

(Ulmus) are the only reported hosts for S.

schevyrewi 1n North America and are its

primary hosts in Asia, it has also been re-

corded to attack Caragana, Elaeagnus,

Malus, Prunus, Pyrus and Salix across its

native central and eastern Asian range

(Wood and Bright 1992; Bright and

Skidmore 1997, 2002; Negron et al. 2005).

S. schevyrewi is of immediate concern

as a potential vector of Dutch elm disease

(DED), caused by the fungal pathogens

Ophiostoma himal-ulmi Brasier & M.D.

Mehrota, Ophiostoma novo-ulmi_ Brasier

and Ophiostoma ulmi (Buisman) Nannf.

(Harrington et al. 2001). Jacobi et al. (2007)

isolated DED from adult S. schevyrewi

emerging from infected Ul/mus americana

L. at levels similar to those from co-

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

Victoria, BC, V8Z 1M5

> Canadian Food Inspection Agency, 506 West Burnside Road, Victoria, BC, Canada, V8Z 1M5

*Canadian Food Inspection Agency, Forestry Division, 1853 Bredin Road, Kelowna, British Columbia,

V1Y 7S9

*Canadian Food Inspection Agency, Plant Health Surveillance Unit, 4321 Still Creek Drive, Burnaby,

British Columbia, V5C 6S7

22 J. ENTOMOL. SOC. BRIT. COLUMBIA 107, DECEMBER 2010

emergent European elm bark beetle, Scoly-

tus multistriatus (Marsham), the primary

vector of DED in North America. Concur-

rently, Koski and Jacobi (2007) demon-

strated that DED was transmitted to feeding

wounds during maturation feeding by S.

schevyrewi adults artificially inoculated

with the disease. The efficiency of S.

schevyrewi as a disease vector remains un-

known as neither its efficiency as a DED

vector (Lee et al. 2009) nor field transmis-

sion of DED from infected to uninfected

elms have been determined (Negron et al.

2005). However, the evidence suggests that

the risk of S. schevyrewi serving as an addi-

tional vector of DED is very high. We re-

port the first records for S. schevyrewi in

British Columbia (BC) and document its

seasonal occurrence.

MATERIALS AND METHODS

S. schevyrewi was detected in traps

near Kelowna, BC during ongoing trials to

test the efficacy of bottle traps relative to

12-unit multiple funnel traps (ConTech

Inc., Delta, BC). Each bottle trap was con-

structed from an inverted clear 2-liter pop

bottle with half of the circumference of the

side wall removed. The threaded portion of

the bottle’s neck was inserted into a 2.54

cm diameter hole drilled in the lid of a 16

oz white plastic cosmetic jar (Industrial

Plastics, Victoria, BC). The trap and the

lures were hung from the pivoting triangu-

lar loops of picture frame hangers riveted

(2.7 cm below the base) to the outside and

inside walls of the bottle, respectively.

Ten replicates of a bottle trap paired

with a funnel trap were established on a

berm on the west margin of the Glenmore

Landfill (49.9556°, -119.4235°). The land-

fill is situated in the Okanagan Very Dry

Hot Ponderosa Pine Variant (PPxh1) of the

Ponderosa Pine biogeoclimatic zone (Hope

et al. 1991); ridges dominated by Ponderosa

pine (Pinus ponderosa P. & C. Lawson) are

present 70 m west of and 800 m east of the

berm. The berm is landscaped with both

ornamental and native trees including Colo-

rado blue spruce (Picea pungens Engelm.),

corkscrew willow (Salix sp.), Douglas fir

(Pseudotsuga menziesii (Mirb.) Franco),

Lombardy poplar (Populus nigra L. cv.

'Italica'), London plane (Platanus x acerifo-

lia (Air.) Willd.), maple (Acer spp.), moun-

tain ash (Sorbus sp.), Ponderosa pine, Rus-

sian olive (Elaeagnus angustifolia L.),

Scots pine (Pinus sylvestris L.), and sumac

(Rhus sp.).

All traps were baited with a proprietary

release system releasing multistriatin, 4-

methyl-3-heptanol and alpha-cubebene at

sub-milligram rates per day at 20 °C

(ConTech Inc., Delta, B.C.) and half of the

replicates were also baited with a half-size

ultra-high release ethanol lure (270 mg/day

at 20° °C, Confech Inc:, Delta, .B:Ganwiie

primary lure for the experiment was se-

lected to target S. multistriatus, which is

widely distributed in the study area (van

Sickle and Fiddick 1982). Traps within a

pair were separated by 4-5 m and pairs

were separated by at least 30 m. Collecting

cups contained 125 ml of propylene glycol

to retain any captured insects. Traps were

deployed on 13 April 2010 and serviced

approximately every two weeks through 5

August 2010 when all lures were replaced.

The screening aid of LaBonte et al. (2003)

was used to separate S. schevyrewi from

other species of Sco/ytus present in the sam-

ples. While the experiment is still ongoing

and identifications of all insects captured

are not complete, all S. mu/tistriatus and S.

schevyrewi recovered to 28 September 2010

have been’ determined. Because S.

schevyrewi is new to the fauna of British

Columbia, we feel it is important to docu-

ment its occurrence in the province prior to

the completion of the study.

J. ENTOMOL. Soc. BRIT. COLUMBIA 107, DECEMBER 2010

RESULTS

In total 27, 551 S. multistriatus and 58

S. schevyrewi were captured between 13

Apr. and 28 Sep. 2010. Collection dates

[number of males and females] for S.

schevyrewi are: 12-28.v.2010 [1d, 49];

28.v-11.vi.2010 [33]; 11-28.vi.2010 [156,

49)» 28.vi=15.vii.2010: [11¢, 109]; 15-

28.vii.2010 [24, 492]; 5-20.viii.2010 [13];

and 2-28.ix.2010 [1¢, 29] No S.

schevyrewi were recovered from the 13-

29.iv.2010, 29.iv-12.v.2010, 28.vu-

5.viil.2010 and the 20.viti-2.1x.2010 sample

periods. In contrast, S. multistriatus was

recovered throughout the complete sam-

pling period. Voucher specimens of 5S.

schevyrewi have been deposited in the Ca-

nadian National Collection (CNC), Ottawa,

ON; the Royal British Columbia Museum

(RBCM), Victoria, B.C., Natural Resources

Canada, Canadian Forest Service, Pacific

Forestry Centre (PFCA), Victoria, BC; and

the Spencer Entomological Collection

(UBCZ), Beaty Biodiversity Museum, Uni-

versity of British Columbia, Vancouver,

BC.

DISCUSSION

The high numbers of S. multistriatus

and the detection of S. schevyrewi at the

Kelowna landfill are surprising as no U/mus

are planted on the site. Sixteen elms were

located around a parking lot 400 m to the

north and scattered mature trees were also

noted on a rural property 725 m south of the

trap line. Deciduous hosts growing along

the berm at the landfill and dead and dying

limbs of the elms to the north of the landfill

were examined by LMH, EJ and MN on 5

August 2010 for signs of attack by bark

beetles. None of the hosts exhibited signs of

attack, thus the source of the S. multistria-

tus and S. schevyrewi populations remains

unknown. The traps were also well re-

moved from two other potential sources of

the Scolytus spp., yard waste and solid

wood packaging. The collection site for

urban yard waste is 600-800 m to the north-

east, while that for wood waste is 600-700

m to the east.

Scolytus schevyrewi has replaced S.

multistriatus as the predominant bark beetle

attacking elms in Colorado, Utah and New

Mexico (Lee et al 2009) and has been im-

plicated as the causal agent of Siberian elm,

Ulmus pumila L., mortality in Colorado

(Negron et al 2005). Siberian elm is widely

planted in the arid interior of BC and has

naturalized in the Okanagan, Similkameen

and Kettle valleys (Brayshaw 1996), and

may be impacted by S. schevyrewi popula-

tions.

ACKNOWLEDGEMENTS

We thank the City of Kelowna for al-

lowing access to the Glenmnore Landfill

for the trapping experiment. We are grateful

to Vince Nealis, Les Safranyik and two

anonymous reviewers for their helpful com-

ments on the manuscript.

REFERENCES

Brayshaw, T.C. 1996. Trees and shrubs of British Columbia. Royal British Columbia Museum Handbook.

UBC Press, Vancouver.

Bright, D.E. and R.E. Skidmore. 1997. A catalog of Scolytidae and Platypodidae (Coleoptera), Supplement

1 (1990-1994), NRC Research Press, Ottawa.

Bright, D.E. and R.E. Skidmore. 2002. A catalog of Scolytidae and Platypodidae (Coleoptera), Supplement

2 (1995-1999), NRC Research Press, Ottawa.

CABI/EPPO 2009. Scolytus schevyrewi. Distribution Maps of Plant Pests No. 729. CABI Head Office,

Wallingford, UK _ [online].

Available

from http://www.cabi.org/cabdirect/

24 J. ENTOMOL. SOC. BRIT. COLUMBIA 107, DECEMBER 2010

FullTextPDF/2009/20093321026.PDF [accessed 11 Aug. 2009].

Harrington, T.C., D. McNew, J. Steimal, D. Hofstra and R. Farrell 2001. Phylogeny and taxonomy of the

Ophiostoma piceae complex and the Dutch elm disease fungi. Mycologia 93: 111-136.

Hope, G.D., D.A. Lloyd, W.R. Mitchell, W.R. Erickson, W.L. Harper and B.M. Wikeem. 1991. Chapter 9:

Ponderosa Pine Zone /n Ecosystems of British Columbia. British Columbia. Edited by D.V. Meidinger,

and J. Pojar, Ministry of Forests Special Report Series No. 6. pp. 139-151 [online]. Available from http://

www. for.gov.be.ca/hfd/pubs/Docs/Srs/Srs06.pdf [accessed 25 Aug. 2010].

Jacobi, W.R., R.D. Koski, T.C. Harrington and J.J. Witcosky. 2007. Association of Ophiostoma novo-ulmi

with Scolytus schevyrewi (Scolytidae) in Colorado. Plant Disease 91: 245-247.

Koski, R.D. and W.R. Jacobi. 2007. Vectoring capabilities of the banded elm bark beetle (Scolytus

schevyrewi Semenov) in relation to the Dutch elm disease fungus (Ophiostoma novo-ulmi Brasier) in

Colorado. Phytopathology 97: S59.

LaBonte, J.R., R.J. Rabaglia and E.R. Hoebeke. 2003. A screening aid for the identification of the banded

elm bark beetle, Sco/vtus schevyrewi Semenov [online] http://ceris.purdue.edu/napis/pests/barkb/schevy/

schevyrewilDnew1 A.pdf [accessed 19 Oct. 2003].

Langor, D.W., L.J. DeHaas and R.G. Foottit. 2009. Diversity of non-native terrestrial arthropods on woody

plants in Canada. Biological Invasions 11: 5-19.

Lee, J.C., I. Aguayo, R. Aslin, G. Durham, S.M. Hamud, B. Moltzan, A.S. Munson, J.F. Negron, T. Peter-

son, I.R. Ragenovich, J.J Witcosky and S.J. Seybold. 2009. Co-occurence of the invasive banded and

European elm bark beetles (Coleoptera: Scolytidae) in North America. Annals of the Entomological

Society of America 102: 426-436.

NAPIS (National Agriculture Pest Information System). 2010. Pest Tracker: Reported Status of Banded

Elm Bark Beetle - Scolytus schevyrewi [online]. Available from http://pest.ceris.purdue.edu/

searchmap.php?selectName=INBQSGA [accessed 11 Aug. 2010].

Negron, J.F., J.J. Witcosky, R.J. Cain, J.R. LaBonte, D.A. Duerr, S.J. McElwey, J.C. Lee and S.J. Seybold.

2005. The banded elm bark beetle: a new threat to elms in North America. American Entomologist 51:

84-94,

Van Sickle, G.A. and R.L. Fiddick. 1982. Forest Insect and Disease Survey 1979, Pacific region. /n Annual

reports of the Forest Insect and Disease Survey 1978 and 1979. Canadian Forestry Service, Department

of the Environment, Ottawa, Ont. pp. 71—79.

Wood, S.L. and D.E. Bright. 1992. A catalog of Scolytidae and Platypodidae (Coleoptera), Part 2: Taxo-

nomic index, volume A. Great Basin Naturalist Memoirs 13: 1-833.

J. ENTOMOL. Soc. BRIT. COLUMBIA 107, DECEMBER 2010

DNA barcoding identifies the first

North American records of the Eurasian moth,

Eupithecia pusillata (Lepidoptera: Geometridae)

JEREMY R. deWAARD!”, LELAND M. HUMBLE'™

and B. CHRISTIAN SCHMIDT‘

ABSTRACT

The first North American records of the juniper pug moth, Eupithecia pusillata (Denis

& Schiffermiiller, 1775) (Lepidoptera: Geometridae), brought to our attention using

DNA barcoding, are presented. Documentation and collection localities suggest it was

introduced, established, and likely has persisted, at least in the Greater Vancouver area

of British Columbia since the mid-1970s. We discuss the integration of DNA barcoding

into routine biosurveillance and forest insect surveys to prevent such delay in recogni-

tion of non-indigenous species—in this case, 34 years.

Key Words: Eupithecia pusillata, Eupithecia interruptofasciata, Eupithecia niphado-

philata, juniper pug moth, Juniperus, non-indigenous species, invasive species, DNA

barcoding

INTRODUCTION

DNA barcoding of biological specimens

has demonstrated repeatedly its utility as a

molecular diagnostic technique that merits

integration into biosurveillance programs.

In contrast to other molecular tools com-

monly employed for species identification

of intercepted organisms, DNA barcoding

is a generic and standardized approach that

meets international standards of data quality

and transparency (Floyd et al. 2010). Sev-

eral studies have demonstrated the efficacy

of this technique for detecting non-

indigenous species and determining native

provenance, for example in leeches (Siddall

and Budinoff 2005), agromyzid leafminers

(Scheffer et al. 2006), tephritid fruit flies

(Armstrong and Ball 2005; Barr 2009),

siricid wasps (Wilson and Schiff 2010), true

bugs (Nadel et al. 2010), and numerous taxa

of moths (Ball and Armstrong 2006; Si-

monsen et al. 2008; Humble et al. 2009;

deWaard et al. 2009; Gilligan and Epstein

2009; Armstrong 2010). Here we report the

first North American records of the juniper

pug moth, Eupithecia pusillata (Denis &

Schiffermiiller, 1775) revealed by DNA

barcoding.

MATERIALS AND METHODS

While compiling a DNA barcode library

for the Geometridae of British Columbia

(deWaard et al., submitted), the cytochrome

c oxidase subunit I (COI) sequences de-

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

BC, Canada V6T 1724

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

(email: jdewaard@interchange.ubc.ca)

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

Victoria, BC, Canada V8Z 1M5 (email: Leland. Humble@nrcan-rncan.ge.ca)

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

K.W. Neatby Building,

Chris.Schmidt@inspection.ge.ca)

960 Carling Avenue,

Ottawa, ON, Canada KIA 0C6 = (email:

26 J. ENTOMOL. Soc. BRIT. COLUMBIA 107, DECEMBER 2010

rived from two Eupithecia specimens were

found to be divergent from known native

Eupthecia. The two sequences were com-

pared to a reference barcode database of

Lepidoptera barcodes using the identifica-

tion engine (BOLD-ID) of the Barcode of

Lite Data Systems (BOLD) (Ratnasingham

and Hebert 2007), and tentatively identified

as Eupithecia pusillata, a Eurasian species

not known to occur in North America. The

reference barcode database for Geometridae

used by BOLD-ID 1s continually validated

by specialists to ensure accurate identifica-

tions, and is particularly well parameterized

due to a global campaign to barcode the

nearly 23,000 species of the family (see

http:// www.lepbarcoding.org/ geometridae

index.php). The nine sequences with identi-

cal and near-identical matches from Europe

were obtained from Axel Hausmann

(Zoological State Collection, Munich, Ger-

many) and Marko Mutanen (University of

Oulu, Oulu, Finland) and combined with

related North American specimens (sensu

Bolte 1990). A neighbour-joining tree was

constructed on BOLD using the Kimura-2-

parameter distance method (Fig. 1).

To pursue confirmation of the identity

of the specimens, the two putative FE. pusil-

lata specimens obtained from the RBCM

(Royal British Columbia Museum, Victoria,

BC) and PFCA (Arthropod reference col-

lection, Pacific Forestry Centre (PFC).

Natural Resources Canada, Canadian Forest

Service, Pacific Forestry Centre, Victoria,

BC), were dissected to examine the genita-

lia following the methods given by Lafon-

taine (2004). Images of genitalia were taken

using a Leica M205C microscope equipped

with a Leica DFC490 camera kit and Leica

LAS Montage system that assembles multi-

ple images in successive planes of focus

into a single image with a large depth of

field. The specimens were verified by com-

parison of the structure of genitalia with

specimens held in the CNC (Canadian Na-

tional Collection of Insects, Arachnids and

Nematodes, Ottawa, ON), and figures of E.

pusillata in Skou (1986) and Mironov

(2003). Related species in the EF. niphado-

philata Dyar, 1904 group (Bolte 1990) were

ruled out by genitalic comparison to speci-

mens in the CNC, as were other North

American species.

Historical data associated with the

specimens were compiled from specimen

labels and Forest Insect and Disease Survey

(FIDS) records (Van Sickle et al. 2001).

The single specimen from PFCA, collected

by FIDS, is uniquely identified by a regis-

tration number (e.g. 76-9-0019-01) that

links the specimen to a FIDS sampling

form, completed at the time of sample col-

lection, as well as a rearing record docu-

menting the status of laboratory rearings.

These records are held on file at PFC.

RESULTS

Specimens examined: |< — /abe/l data

(handwritten information in italics, individ-

ual lines separated by comma, multiple

labels separated by *|°):

No. 76-9-0019-01, Date 19 vii, F.1.[D.]

S.1976 | c. juniper, Port, Coquitlam BC

Ac. No. PFC, 2007-0271.

The specimen was initially identified as

Eupithecia unicolor (Hulst). The FIDS re-

cords document that this specimen was one

of two adults reared from five larvae and

five pupae (10 individuals in total) col-

lected by the B.C. Forest Service on Mt.

Burke, Port Coquitlam (UTM 10 53 546

[49.3, -122.7], Elevation 900 ft), on 15 May

1976. The host recorded was common juni-

per (Juniperus communis L.); Remarks &

Symptoms state “Attacking several orna-

mentals with moderate damage”. The date

recorded on the specimen label is the date

of adult eclosion. While the Rearing Record

indicates a second adult eclosed on 8.vii.76

and was subsequently spread, the specimen

could not be found in the PFCA reference

collection.

1° — label data:

BC, N. Vancouver, 5 AUG 1986, CS.

Guppy | ROYAL BRITISH, COLUMBIA

MUSEUM, ENT991-12573 |.

This specimen was identified as

J. ENTOMOL. Soc. BRIT. COLUMBIA 107, DECEMBER 2010 27

BC ZSM Lep 22832

BC ZSM Lep 22831

MMo0781 |

MMO1785 | FI

MMo08340 |

PFC-2007-0271

ENT991-012573

BC ZSM Lep 03633 DE

BC ZSM Lep 21945 IT

MMO1786 |

mmos143_—

PFC-2007-0259 |

Dun-08-073 | CA: BC

CNCLEP00035444 |

uasms9396. | CAVAB

Dun-08-081 _

CGWC-1141 |

CGWC-1155 |

CGWC-1202 _

CGWC-1205 |

Eupithecia pusillata

CA: BC

| E. niphadophilata

CA: BC E. interruptofasciata

0.01

Figure 1. Neighbour-joining tree of Eupithecia pusillata and two closely related species, F.

niphadophilata and E. interruptofasciata. Tree was reconstructed with the barcode fragment of

the cytochrome oxidase I (COI) gene. Sequences shaded in grey are from two individuals col-

lected in Vancouver, Canada. Abbreviations: DE — Germany, FI — Finland, IT — Italy, CA —

Canada, BC — British Columbia, AB — Alberta.

Eupithecia sp. in the collection before ten-

tative assignment to Eupithecia intricata

taylorata Swett by JRD.

Diagnosis: Eupithecia pusillata is most

similar to E. niphadophilata and particu-

larly E. interruptofasciata, but a number of

Eupithecia species are superficially very

similar and identification should be based

on examination of genitalia. Compared to

E. interruptofasciata, which is structurally

most similar, the male 8" sternite apical

prongs are narrower, more blunt and the

apical cleft is shallower; the base of the

sternite 1s also narrower overall with a shal-

lower medial invagination. The basal half

of the male vesica is armed with one spine,

not two as in E. interruptofasciata. In the

female genitalia, the large spines on the left

side of the ductus bursae do not extend be-

yond the mid-point of the ductus, but ex-

tend beyond the midpoint in both E. inter-

ruptofasciata and E. niphadophilata.

Description: A small moth with a wing-

span of 16-22 mm (Mironov 2003) (Figs.

2a, 2e). Forewing narrow, mostly shades of

light brown with black transverse lines and

oblong discal spot. Hindwing pale grey-

brown with weakly marked transverse lines

and variable discal spot. Abdomen pale

grayish brown with narrow black lateral

stripes. Male genitalia (Fig. 2d) composed

of broad valva with small ventral process,

heavily sclerotized sacculus, vesica with

three horn-like cornuti, simple aedeagus

(Fig. 2c) and elongated 8" sternite with two

narrow apical processes (Fig. 2b). Female

genitalia composed of elongate and scle-

rotized bursa copulatrix (Fig. 2h) with small

spines at base and larger spines at margin.

Ovipositor is simple with long setae (Fig.

2f). Terminal segment of pupal case 1s stout

with prominent lateral lobes and cremaster

bearing four pairs of hook-like setae (Figs.

Din wal).

28 J. ENTOMOL. SOc. BRIT. COLUMBIA 107, DECEMBER 2010

Figure 2. Morphology of Eupithecia pusillata. a) male, dorsal view, b) male, 8" sternite, c)

male, aedeagus, d) male, genital capsule e) female, dorsal view, f) female, ovipositor, g) fe-

male, bursa copulatrix, h) pupa, terminal segment, dorsal view, 1) pupa, terminal segment, lat-

eral view. Scale bars: a, e = 5 mm; b—d, f-i = 0.5 mm. A colour version of this figure is

available from Dr. Lee Humble.

Distribution and Habitat: In its native

European range, the nominate subspecies is

widely distributed from southern Europe,

its range extends to the Mediterranean from

eastern Spain to mainland Greece and Ro-

mania, then extends north and west across

northern Ukraine into western Russia. With

the exception of Corsica, it has not been

recorded from the islands of the Mediterra-

nean. To the north it is present in the British

Isles, through central Europe, north to

northern Scandinavia, and into western

Russia across the southern Kola Peninsula

(Skou 1986; Mironov 2003; Karsholt & van

Nieukerken 2010). A disjunct population of

E. pusillata is present in the Caucasus

Mountains (Mironov 2003). In Asia, its

range extends across Russia from Sahkalin

through Siberia, the Altai and Caucasus

regions (Skou 1986). The subspecies E.

pusillata scoriata Staudinger, 1857 has

been recorded only from Iceland and south-

western Greenland (Mironov 2003). Mi-

ronov et al. (2008) recently described a

third subspecies, E. pusillata kashmirica

Mironov and Ratzel from the Himalayas. In

natural settings, E. pusil/lata can be found in

heaths, forest edges, rocky cliffs, and simi-

lar habitats where the primary host grows.

In urban areas, it can be common in gar-

J. ENTOMOL. Soc. BRIT. COLUMBIA 107, DECEMBER 2010

dens. It is known from sea level up to ap-

proximately 2,500 m elevation in the Sierra

Nevada (Spain) and the Alps (Switzerland)

(Weigt 1993; Mironov 2003).

Life History and Notes: The following

data are based on European populations,

and it 1s expected that flight times, voltin-

ism and larval hosts will be similar in North

America, should extant populations be dis-

covered. Univoltine, with larval stage from

late April to mid-June and adult flight pe-

riod from mid-July to late September (Skou

1986; Mironov 2003). As its common name

implies, the primary host of E. pusillata is

common juniper, Juniperus communis L.

(Cupressaceae) (Skou 1986), of which it

feeds on young needles and flowers. It is

generally regarded as monophagous

(Mironov 2003), although it has also been

recorded feeding on Douglas-fir, Pseu-

dotsuga menziesii (Mirb.) Franco

(Pinaceae) in France (Roques et al. 2006),

where this North American tree is culti-

vated. The host of the subspecies scoriata

and kashmirica 1s not known, but is pre-

sumed to also be Juniperus. Eupithecia

pusillata overwinters in the egg stage and

pupates in a loose web in the ground (Skou

1986). It is attacked by a variety of ichneu-

monid and braconid species listed in Mi-

ronov (2003). It is not known if other native

or ornamental species of Juniperus are suit-

able hosts in British Columbia.

DISCUSSION

Eupithecia Curtis is a large genus with

1529 described species and_ subspecies

(Scoble 1999; Scoble & Hausmann 2007),

and about 160 species in North America

(Powell and Opler 2009). The North Ameri-

can species were revised by McDunnough

(1949), and the Canadian fauna was revised

by Bolte (1990). Eupithecia pusillata is part

of the niphadophilata species group, which

includes two Nearctic and one Palearctic

species (Bolte 1990), all feeding primarily

on junipers (Skou 1986; Bolte 1990).

Although we currently have only two

specimens of Eupithecia pusillata from

North America, we can extract a great deal

of information from the associated data

documentation. First of all, the collections

were made in urbanized Vancouver, BC,

suggesting the species was introduced. The

lack of records, particularly from inland BC

(which is well-surveyed for macro-

Lepidoptera), the Yukon Territory and

Alaska, lead us to conclude that the species

is not naturally Holarctic like some

Eupithecia (see Skou 1986, Bolte 1990).

Furthermore, the six Eupithecia species

considered Holarctic all show at least 1%

COI sequence divergence (data not shown)

indicative of separation in the Pleistocene.

The absence of additional records also sug-

gests that there has not been substantial

spread beyond the point of introduction.

Secondly, the locality of the first collection

(Mt. Burke), the number of individuals re-

corded (ten), and the damage observations

in the FIDS record, all indicate that there

was an established E. pusillata population

in BC in 1976 (but note this is the only

FIDS record of a Eupithecia on juniper

from greater Vancouver). And lastly, the

1986 collection from North Vancouver sug-

gests that the population has persisted, or it

did so for at least a decade. Subsequent

surveys, initially in the Vancouver area, are

required to determine the contemporary

status of this species.

The excellent documentation of FIDS

that enabled inferences about the status of

E. pusillata is unfortunately a relict of the

past; the program ceased in 1996 after al-

most 50 years of operation due to budgetary

cut-backs (Van Sickle et al. 2001). Pro-

grams such as this, based on surveying or

inventorying diversity, are simultaneously

a) a tremendous resource for managers,

foresters and scientists, and b) reliant on

tremendous resources themselves particu-

larly in terms of highly qualified personnel

(e.g. Marshall et al. 1994). The present case

illustrates the value of these long-term,

well-documented biological surveys, but

these programs are often hindered by the

30 J. ENTOMOL. Soc. BRIT. COLUMBIA 107, DECEMBER 2010

necessity to rear immatures to allow the

diagnosis of species. Just as DNA barcod-

ing makes an invaluable tool for biosurveil-

lance (Floyd et al. 2010), it could likewise

assist any regional or national biomonitor-

ing program of similar scope to FIDS. Bar-

coding could not only identify immature

stages (Ahrens et al. 2007) making rearing

nonobligatory, it could also identify the

plant meal of gut contents (Miller et al.

2007), identify parasitoids (Rougerie et al.

in press), and trace complex food webs

(Sheppard et al. 2004; Smith et al. in press).

Decreasing costs and increasing capabilities

of sequencing (e.g. Shokralla et al. 2010)

are certain to make species diagnosis in this

form time- and cost-effective. Furthermore,

most years of the FIDS program predated

electronic databases, so it would also be

better served by modern and online rela-

tional databases such as BOLD

(Ratnasingham and Hebert 2007). With

DNA barcoding in place, a resource similar

to FIDS could once again be realized, and

without having to expend substantial re-

sources as a cost. It would also, without

question, speed the time of non-indigenous

species detection—from years (34 in the

case of E. pusillata) to days.

ACKNOWLEDGEMENTS

We are indebted to Axel Hausmann

(Zoological State Collection Munich) and

Marko Mutanen (University of Oulu) for

allowing us access to the Eurasian se-

quences. We also wish to thank Stephanie

Kirk for the specimen imaging, the staff at

the Canadian Centre for DNA Barcoding

for their assistance with genetic analysis,

and Sava Barudzija (Natural Resources

Canada, Canadian Forest Service, Frederic-

ton, NB) for extraction of historical FIDS

records. Rene Alfaro, Isabel Leal and two

anonymous reviewers provided valuable

comments on the manuscript. 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.

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J. ENTOMOL. SOC. BRIT. COLUMBIA 107, DECEMBER 2010

J. ENTOMOL. SOc. BRIT. COLUMBIA 107, DECEMBER 2010

The Fireflies (Coleoptera: Lampyridae) of British Columbia,

with special emphasis on the light-flashing species

and their distribution, status and biology

ROBERT A. CANNINGS!, MARC A. BRANHAM’

and ROBERT H. McVICKAR°

ABSTRACT

In British Columbia the family Lampyridae (fireflies) is poorly known. Ten species in

five genera are recorded but adults of only two species produce flashed bioluminescent

signals. Before 1997, museum collections that we examined held specimens of only one

flashing species from three BC localities, despite recent and widespread reports of flash-

ing fireflies in the province. A solicitation of specimens and sight records from ento-

mologists and naturalists resulted in the discovery of 14 additional collection localities

for the two species. Sight records are summarized but are not recorded in detail. As far

as is known, Photuris pennsylvanica (DeGeer) (sensu lato), is restricted in BC to the

southern Rocky Mountain Trench (East Kootenay region). Photinus obscurellus Le-

Conte, herein recorded in BC for the first time, is widespread in the northeast, central

and southern Interior of the province. This paper briefly summarizes the BC lampyrid

fauna, examining the status, distribution and biology of the two flashing species in more

detail.

Key Words: Lampyridae, fireflies, flashing species, Photuris, Photinus, British Colum-

bia, distribution, identification

INTRODUCTION

33

Beetles of the family Lampyridae

(fireflies) are poorly known in British Co-

lumbia (BC). McNamara (1991) recorded

nine species in four genera, but adults of

only one of these species produce flashing

signals. This paper briefly summarizes the

BC lampyrid fauna but examines, in more

detail, the status of the two flashing species,

one in each of the genera Photuris LeConte

and Photinus Laporte, now known to occur

in BC. One of these species is recorded in

the province for the first time.

Adults of nocturnal bioluminescent spe-

cies use their flashed signals in courtship.

Characteristics of these signals, such as

flash number, flash duration and the inter-

val between flashes, are important in spe-

cies recognition but, in some cases, such as

in Photuris, are still not completely reliable

for identification (Lloyd and Branham in

press, Branham and Greenfield 1996, J.E.

Lloyd pers. comm.). These beetles are

chemically defended and, when handled or

attacked, often exude defensive compounds

from the body, especially the elytra (Eisner

et al. 1978). As larvae, many lampyrid spe-

cies are soil dwelling predators that eat in-

sects, snails, worms and other invertebrates,

while others are arboreal or aquatic and

feed on snails (LaBella and Lloyd 1991).

Photinus larvae are subterranean, seldom

observed, and may specialize on eating

Royal British Columbia Museum, 675 Belleville Street, Victoria, BC V8W 9W2, (250) 356-8242, rcan-

nings@royalbcmuseum.be.ca.

> Department of Entomology and Nematology, University of Florida, P.O. Box 110620, Gainesville, FL

32611-0620, marcbran@ufl.edu.

* Died 13 May 2008. We dedicate this paper to the memory of Bob McVickar, whose enthusiasm for the

project got it off the ground.

34 J. ENTOMOL. Soc. BRIT. COLUMBIA 107, DECEMBER 2010

earthworms; Photuris larvae are surface

dwelling, omnivorous predators and scav-

engers of dead insects and fallen berries

(Buschman 1984, Lloyd 2002). All lam-

pyrid larvae have a luminous organ on ab-

dominal segment 8 (Lloyd 2002); light pro-

duction warns predators that these larvae

are distasteful (Sivinski 1981, De Cock and

Matthysen 2003).

Adults of the best known BC fireflies

are diurnal, non-flashing species. These

taxa use pheromonal communication for

pair formation (Lloyd 2002, Branham and

Wenzel 2003). Ellychnia Blanchard is the

most diverse genus in BC with five species:

E. corrusca (Linnaeus) (widespread), FE.

facula LeConte (Okanagan, central coast),

E. greeni Fender (southern Interior), £.

hatchi Fender (widespread on coast), and E.

lacustris LeConte (Terrace). All but E. cor-

rusca, Which is transcontinental, are re-

stricted in Canada to BC (McNamara

1991). Phausis nigra Hopping (southern

Interior) is unknown outside BC but Phau-

sis rhombica Fender (southern Interior) is

also known from Alberta (McNamara

1991), Washington and Oregon (Fender

1961) and Montana (M.A. Branham pers.

obs.) -- both are Cordilleran species. Pyro-

pyga nigricans (Say) (southern Interior)

ranges from BC east to the Atlantic Ocean

(McNamara 1991).

Before this study, collections of flashing

lampyrid adults in BC were rare. The only

literature records for BC are a reference to

Photuris pennsylvanica (DeGeer) in south-

eastern BC (Fender 1961) and a subsequent

inclusion of the species in the BC fauna by

McNamara (1991). These references evi-

dently refer to collections in 1928 and 1958

(in RBCM and UBC collections, respec-

tively [see Specimens Examined]). Sight-

ings of flashing fireflies, beginning in 1996

in the central and south-central Interior of

the province, where no such species had

been collected before, stimulated two of us,

Cannings and McVickar, to solicit speci-

mens and sight records of flashing beetles

from the BC naturalist and entomological

communities by word of mouth and through

various newsletters (Cannings 1999).

In 2010, an additional extensive compi-

lation of observations made by naturalists,

ranchers and others in the East Kootenay

region was organized by the Columbia

Wetlands Stewardship Partners (Jamieson

2010). This study, motivated by plans to

promote fireflies as iconic wetland inhabi-

tants and as a focus for conservation and

wetland education, improved our knowl-

edge of the distribution of Photuris in the

area. No specimens were collected. The

survey covered the East Kootenay Trench

from the US border to Donald, (north of

Golden), a distance of about 345 kilome-

tres.

RESULTS AND DISCUSSION

Specimens examined

Specimens received by the RBCM were

identified by Marc Branham and James

Lloyd and are listed below along with those

examined from museum collections (see

also Fig. 1).

Museum collection abbreviations: CNC

— Canadian National Collection of Insects,

Arachnids and Nematodes, Ottawa, ON;

PFC -- Pacific Forestry Centre, Victoria,

BC; RBCM -- Royal British Columbia Mu-

seum, Victoria, BC; UBC -- Spencer Ento-

mological Collection, Beaty Biodiversity

Museum, University of BC, Vancouver,

BC. There are no BC specimens of Photuris

or Photinus in the E.H. Strickland Entomo-

logical Museum, University of Alberta,

Edmonton AB, the Oregon State Arthropod

collection, Oregon State University, Cor-

vallis, OR or the California Academy of

Sciences, San Francisco, CA, the museum

that contains much of the Ralph Hopping

and Hugh Leech beetle collections

(including considerable BC material) and

thus the US collection most likely to house

significant numbers of BC beetles.

Photuris pennsylvanica (DeGeer 1774)

CANADA, BC, Fort Steele, 16.vi.1958,

R.J. Andrews (8%, UBC); Fort Steele,

25.vi.1959, Forest Insect Survey (41¢

J. ENTOMOL. SOC. BRIT. COLUMBIA 107, DECEMBER 2010

%

yr a

. mt

. of

sa ;

} i

»

7

ft

ad

*

ae

te

refs ed

x

a

“

aie

:

at

Set |

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te

35

Figure 1. Distribution of Photuris pennsylvanica (@) and Photinus obscurellus (@) in British

Columbia as represented by specimens.

caught in flight, UBC; 4 caught in flight,

RBCM); Fort Steele, CP railway tracks,

south yard switch, 30.vi.1998, Greg Ross

(34, RBCM); Haha Creek, between Haha

Creek Rd. and Haha Creek, 11.vii.1998,

Greg Ross’ (24, RBCM);’Ta Ta’ Lake,

18.vi.1958, E.K. White (3¢, UBC); Win-

dermere, 27.vi.1928, W.B. Anderson (1.3,

RBCM).

Photinus obscurellus LeConte 1852

CANADA, BC, SBednesti Lake

(93°57 ENG 23°22" 07" W), 30.41.2007.

R.V. Rea (14, RBCM); Bonaparte Lake, 17

km W at Moose Lake (51°18’14"°N x

120°55’44”W), 15.vii.1997, Joe Cortese

(12, PFC, with photos, Fig. 2). Fort St.

John, 20 km SW (56 067°37”N_ x

121°04°59”°W), 4.vi1.2009, Mark Phinney

(62, 19, RBCM); Fort St. John, 20 km SW

(56° 06’47”"N x 121°05’19"W), 4.vii.2009,

Mark Phinney (3¢, RBCM); Horsefly,

swamp 6 km S of Bells Lake, 3.vii.2002,

Marcus Charles (14, RBCM); 100 Mile

House, Horse Lake, Fawn Creek Rd.,

10.vii.2002, Pat Griffin (4¢, RBCM);

Prince George, Ness Lake, 9690 Anne Rd.,

10.vii.1999, Marie Pearson (14, RBCM);

OQuesnely Beryl” Rd? 162°" 57°22" Ne *x

122°26734”W), 10.viii.2010, Clint Tibideau

(14, RBCM); Quesnel, Cottonwood House

(3°05 20° Nex 12291245" W), 13.vi 2005;

John Massier (12, RBCM); Sheridan Lake,

36 J. ENTOMOL. SOc. BRIT. COLUMBIA 107, DECEMBER 2010

Figure 2. Photinus obscurellus (&): left, dorsal view; right, ventral view. Scale lines are | mm

apart. Specimen photographed at Pacific Forestry Centre by Bob Duncan. Collected by Joe

Cortese at Moose Lake, 17 km west of Bonaparte Lake (51°18’14”°N x 120°55’44”’W) on 15

July1997 (Males look similar but have light-producing organs covering the 5" and 6" visible

ventral segments (true abdominal segments VI and VII).

vii.2003, Frank McFadden (4¢, RBCM);

Shuswap Lake, Eagle Bay, Herman Lake, N

end (50°55’08"N x 119°09°45”W),

24.vi.1997, Dawn Kellie (34, RBCM);

same locality and date, Dennis St. John

(44, RBCM); same locality and date, R.H.

McVickar (2¢, RBCM; 1, CNC); same

locality, 10.vii.1996, R.H. McVickar (2¢,

RBCM); Skmana Lake, 7.vu.1998, R.H.

MceVickar (14, RBCM).

Sight records

In addition to the collections and associ-

ated observations listed above, numerous

sightings of flashing fireflies were reported

by 40 respondents across the province. Six-

teen people participated in the 2010 survey

in the East Kootenays; observations in-

cluded direct sightings as well as hearsay,

past and present. Some recent sightings

included exact coordinates and dates or

even photographs; others were less precise.

Unless an observer is experienced, sight-

ings of flashing fireflies are difficult to as-

sign to species, and most of the sight re-

cords gathered were not verified by voucher

specimens. For these reasons and because,

for the most part, collected specimens fall

within the known distributions of the two

flashing species in BC (Fig. 1), sight re-

cords are not listed in detail. They do, how-

ever, support the notion that these beetles

are not rare and that their populations, espe-

cially those of Photinus, are widespread.

We have reports from two areas outside the

generally known ranges of Photuris and

Photinus in BC. There are three from the

West Kootenay region (two in the Nelson

area, one near Trout Lake) (Jakob Dulisse,

pers. comm.). Nelson is not far from the

Cranbrook populations of Photuris, but the

two areas are separated by extensive moun-

tainous terrain. Trout Lake is closer to the

Shuswap region where Photinus is found.

Three other unsubstantiated reports come

from Vancouver Island (Campbell River,

Parksville, Sidney) (R.A. Cannings, unpubl.

data) but there are no others from the BC

coast and no specimens have been seen or

reported from west of the Coast Mountains.

Photuris pennsylvanica

Before the 2010 survey, sightings were

reported from nine localities in the East

Kootenay region (some the same as speci-

men localities), all presumably for Photuris.

Dates range from 1966 to 2008 and locali-

ties range from Canal Flats south to Haha

Creek and the Bull River Fish Hatchery

near Wardner. Photographs of specimens

(no vouchers collected) accompany the

record documented by Barb Houston at

J. ENTOMOL. Soc. BRIT. COLUMBIA 107, DECEMBER 2010

37

Figure 3. Photuris pennsylvanica (<4): left, dorsal view; right, ventral view. Live male speci-

men photographed in vial by Barb Houston at Bummers Flats, north of Cranbrook

(49°39°20"N x 115°40’29”W) on 19 June 2008.

Bummers Flats (49°39’20”"N~ x

115°40’29"W) on 19.vi1.2008 (Fig. 3). In

the 2010 information survey, 16 additional

localities were reported, ranging from

Brisco in the north to Newgate near the US

border in the south. Most of these observa-

tions were actually made in earlier years but

seven were of fireflies seen in 2010. How-

ever, only three of these records represent

sites not recorded before 2010 (Jamieson

2010).

Photinus obscurellus

About 20 of the sight records reported

came from within the known range of Phot-

inus obscurellus; they are assumed to be of

this species. Even as long ago as the 1920s,

anecdotal records from localities as far

apart as Enderby, Prince George and the

Kiskatinaw River in the Peace River region

embraced the present known latitudinal

range of the species. Other general areas

represented are Prince George north to

McLeod Lake, Quesnel and the Shuswap

region. The most westerly sightings range

from Vanderhoof in the north, to Nazko

Lakes Provincial Park (Pynn 1996) on the

Chilcotin Plateau, to the Carpenter Lake

area in the south.

The identification of Photuris and Phot-

inus in BC

Adults of the two known species of

flashing fireflies in BC can be distinguished

from the other lampyrids in the province by

the presence of pale, light-producing organs

on the underside of the abdomen (Figs. 2,

3). They completely cover the 5th and 6th

visible ventral segments in males but form a

more restricted band in females.

These species can be separated from

each other by the structure of the fore and

mid tarsal claws -- in Photuris these claws

are bifid (one of the two claws is “split’)

while those of Photinus are simple (Lloyd

2002). In Photuris, the legs are long and

Slender (Fig. 3); in Photinus they are

shorter and more flattened (Fig. 2).

With few exceptions, North American

Photuris cannot be accurately identified to

species using either morphological or flash

pattern characters. Although the morphol-

ogy and scarce behavioural observations

that are available for Photuris specimens

collected in BC are consistent with charac-

teristics of Photuris pennsylvanica (J.E.

Lloyd, pers. comm.), this identification

should not be considered more than a con-

venient and useful referent and working

designation. Nevertheless, we are using the

name here because it is already in use for

the BC records (Fender 1961, McNamara

38 J. ENTOMOL. Soc. BRIT. COLUMBIA 107, DECEMBER 2010

1991) of Photuris.

An understanding of the complex taxo-

nomic history of Photinus obscurellus is

useful for those attempting to identify

Nearctic Photinus specimens. LeConte

(1852) described the species but in his syn-

opsis (LeConte 1881) he considered it a

synonym of Photinus ardens LeConte.

Lloyd (1966), describing the behaviour of

Photinus ardens (p. 47-49), is actually re-

ferring to Photinus obscurellus; at the time

he was following LeConte (1881) and

Green (1956). But his field studies later

revealed that Photinus obscurellus deserved

formal recognition, and in 1969 he de-

scribed the flashing behavior of the true

Photinus ardens and restored Photinus ob-

scurellus to species status (Lloyd 1969).

Because Photinus obscurellus was treated

as a synonym of Photinus ardens in

Green’s (1956) widely used identification

key, specimens of Photinus obscurellus in

museum collections are almost always mis-

identified as Photinus ardens.

Behaviour and habitat

Photuris pennsylvanica

We have little information on details of

the behaviour of Photuris pennsylvanica in

southeastern BC. Six collection records in

the East Kootenay region range from 16

June to 11 July; sight records extend this

period only one day earlier. At Haha Creek

on 11 July 1998 Greg Ross collected two

specimens and noted that individuals ir-

regularly produced a 1-second flash every 8

or 9 seconds.

Jamieson (2010) notes that in the East

Kootenays, the species inhabits pothole

wetlands on the benches of the main valley

(Hahas, Ta Ta, Butts and Cub Lakes) as

well as the wetlands along the major rivers

-- the Kootenay (Fort Steele, Bummers

Flats, and so on) and the Columbia (Canal

Flats, Luxor Creek, Brisco areas). These

habitats are typically associated with

springs or small creeks that flow year-

round. The springs sometimes emerge in

the bottom of wetlands or ponds.

Photinus obscurellus

In BC, 14 collection records range from

13 June to 10 August; the latter date is a

month later than any others. At Herman

Lake near Shuswap Lake, McVickar re-

corded flashing in May 1996 and it contin-

ued for six weeks. In 1997 the onset was

later but continued into July and was not

finished by 8 July. The site is a cattail

(Typha latifolia Linnaeus) marsh bordering

a small lake about 250 m long; the fireflies

mostly flash in and above the cattails but

are active up to 250 m inland from the

marsh. The first flashes appear when the

last light is fading but the full performance

commences during complete darkness,

about midnight in June. Before the onset of

the full flashing display there appears to be

a warm-up period, which begins before the

onset of full darkness. Females emit as

many as 7 or 8 flashes in quick succession.

Males settle into a pattern of two or three

flashes produced while flying in a straight

line followed by another flash given on a

curved flight path. The most common pat-

tern is: flash, flash, curving flash. Females

settle into a pattern of single flashes given

at considerable intervals.

On warm nights between | and 21 July

2002 near Horse Lake in the Cariboo, about

15 insects at a time flew and flashed. Speci-

mens were collected on 10 July about 22:15

PDT, just after dark (Pat Griffin, pers.

comm.). Marie Pearson (pers. comm.) ob-

served at least 100 fireflies (and collected

one) flashing at 23:30 PDT near Ness Lake

northwest of Prince George on 10 July

1999. The habitat consisted of a small,

spring-fed marsh, flooded in spring but only

moist in summer; typical ground cover is

moss, willows (Salix spp.), Buckbean

(Menyanthes trifoliata Linnaeus) and Bog

laurel (Kalmia microphylla (Hook.) Heller).

In 1997, Photinus obscurellus individuals

were active between mid-July and 5 Au-

gust; the Pearsons have seen them there

since 1983.

In the Peace River region near Fort St.

John, Mark Phinney collected specimens in

sedge meadow wetlands within a forested

landscape from 00:30 to 01:15 MDT on 4

July 2009. At one site about 25 were flash-

ing, not flying, but perched on the tops of

willows and sedges. The signals usually

J. ENTOMOL. SOc. BRIT. COLUMBIA 107, DECEMBER 2010

consisted of two long flashes, each about 2

seconds in duration, separated by about |

second. The time between these sessions

was variable and seemed affected by the

flashes of neighbours.

Distribution and status

Photuris pennsylvanica

Species of Photuris range from Canada

to Argentina, with 22 known species in

North America and 28 new species descrip-

tions in preparation (J.E. Lloyd, pers.

comm.). They range mostly in the eastern

United States, west to Colorado and south-

west Texas (Lloyd 2002).

As mentioned above, no detailed state-

ment can be made about the geographical

distribution of Photuris pennsylvanica, as

the majority of “determined” specimens in

collections are questionably identified. The

six BC collection records from the

Kootenay and Columbia valleys range from

Windermere in the north to Haha Creek in

the south (Fig. 1). Sight records extend this

almost linear distribution in the Rocky

Mountain Trench from Brisco in the north

to Newgate in the south. In general, accord-

ing to the anecdotal information gathered

from residents in the region, populations

probably have declined in the past several

decades. Specimen and sight records come

from 25 localities between 2001 and 2010,

while at 10 additional sites, beetles have not

been seen since they were reported between

1950 and 2000 (Jamieson 2010).

Because the genus needs revision based

on behavioural, morphological and molecu-

lar data and because the flash patterns of

the BC species have not been studied, the

specific identity of the BC population re-

mains provisional at this time.

Photinus obscurellus

Species of Photinus range from Canada

to Argentina, with 34 described species in

North America and an additional 13 known

but undescribed (J.E. Lloyd, pers. comm.).

The genus is widely distributed on the con-

tinent but there are only scattered popula-

tions west of Texas and Kansas (Lloyd

2002).

Photinus obscurellus ranges from New-

foundland, Nova Scotia, New Brunswick

a9

and Maine through southern Quebec and

Ontario west to North and South Dakota,

Manitoba and Saskatchewan (Lloyd 1969,

unpublished CNC data) with an outlying

population in BC. We are unaware of any

collection records from Alberta, Washing-

ton, Idaho or Montana. In BC the species

appears restricted to the Peace River region

and the central I[nterior from about

Mackenzie south to the Shuswap Lake area

(Fig. 1). A sight record of this species ex-

tends the range west to the central Chilco-

tin.

The dates of all BC collections of Phot-

inus range from 1996 to 2010, the result of

the present study aided by the BC naturalist

community. Why no specimens were col-

lected before this time is a mystery because

many entomologists have collected insects

in central BC over the last century. Can-

nings has never seen a flashing firefly in

BC despite having a strong interest in in-

sects in the province for almost 50 years;

until the present project was begun, he had

never heard of any reports from the central

Interior. Why did Ralph Hopping, Hugh

Leech or James Grant, all avid, professional

coleopterists working at various times be-

tween about 1920 and 1980 within the

southern part of the present range of Phot-

inus obscurellus, apparently never collect

any? On the other hand, as reported above,

the population of Photuris in the East

Kootenays was known from collections as

far back as 1928.

A lack of collections might be attributed

in part to a lack of communication between

residents and entomologists interested in

these beetles. When directly asked in this

study, naturalists and ranchers responded

with numerous memories of flashing fire-

flies as far back as the 1920s in the

Shuswap, Prince George and Peace River

regions, but professional entomologists may

not have heard such accounts.

Although a reason for the significant

separation of Photinus records between

northern Saskatchewan and _ northeastern

BC (approximately 800 km) might be lack

of collecting in this geographical gap, this

hiatus and the historical lack of specimens

40 J. ENTOMOL. Soc. BRIT. COLUMBIA 107, DECEMBER 2010

from BC have suggested to some that Phot-

inus obscurellus might be recently intro-

duced to BC from the East. However, the

widespread distribution of the species in

BC and the newly reported sight records in

the province from many areas over many

decades indicate that this is unlikely. It is

possible that the beetle arrived in BC via

the railways and was somehow helped in its

spread by various railway lines. Even to-

day, most of the sightings and collection

records occur within 30 km of a railway.

Similarly, in the East Kootenay region of

southeastern BC, Photuris pennsylvanica

apparently inhabits wetlands of the Rocky

Mountain Trench, about 220 km from north

to south, with all records close to railway

tracks. Members of this genus exhibit a

very patchy distribution in western North

America; this or other species of Photuris

also occur in Montana, just southeast of the

BC population (M. Ivie, pers. comm.). The

apparent association between these firefly

distributions and railway lines is interesting

and raises several points for consideration.

Wetland habitats are commonly situated in

valleys where many railroads are located. In

addition, the presence of a railroad berm

and the additional weight it applies to the

surrounding soil might be in part responsi-

ble for the creation of new wetland habitat

(M. Ivie, pers. comm.).

We hope this paper stimulates future

systematic, ecological and behavioural re-

search on these firefly populations and that

more targeted studies are able to evaluate

hypotheses concerning these interesting

distribution patterns.

ACKNOWLEDGEMENTS

We thank James Lloyd (University of

Florida, Gainesville) for sharing data from

his files on the distribution of Photinus ob-

scurellus as well as examining several

specimens, particularly those of Photuris,

and checking their identity. Michael Ivie

(Montana State University, Bozeman)

shared his knowledge of lampyrids. Yves

Bousquet (Canadian National Collection of

Insects, Arachnids and Nematodes, Ot-

tawa), Karen Needham (Spencer Entomo-

logical Collection, Beaty Biodiversity Mu-

seum, University of BC, Vancouver), Jane

Seed and Bob Duncan (Pacific Forestry

Centre, Victoria), David Kavanaugh

(California Academy of Sciences, San

Francisco) and Christopher Marshall

(Oregon State University, Corvallis)

checked their collections for BC specimens

of Photuris and Photinus.

We also thank the many dedicated ob-

servers across British Columbia who sup-

plied specimens, sight records and other

insight: Nola Alt, Jack Bowling, Bob

Brawn, Steve Byford, Bob Cale, Syd Can-

nings, Marcus Charles, Rob Cochrane, Joe

Cortese, Neil Dawe, Len Donaldson, Mollie

Donaldson, Jakob Dulisse, Andy Dzilums,

Paul Galbraith, Cam Gillies, Pat Griffin,

Cris Guppy, Larry Halverson, Vicki Han-

sen, Barb Houston, Bob Jamieson, Margie

Jamieson, Gerry Jensen, Phil Jones, Doug

Jury, Dawn Kellie, Mike Kennedy, Burke

Koral, Staffan Lindgren, Joselyn MacGre-

gor, Larry MacKenzie, John Massier, Ann

Matheson, Ron McDougall, Frank McFad-

den, John Morrison, Adam Moss, Ann

Moss, Penny Ohanjanian, Nellie O’Neill,

Dennis Patrician, Marie Pearson, Brian

Pearson, Mark Phinney, Bea Prehara, Larry

Pynn, Leo Rankin, Roy Rea, Greg Ross,

Dennis St. John, Walter Schoen, Lawrence

Simon, Al Singer, Nick Spelay, Joyce

Swanky, M. Tapsen-Jones, Clint Tibideau

and Ellen Zimmerman.

The comments and suggestions of three

anonymous reviewers greatly improved the

manuscript.

J. ENTOMOL. Soc. BRIT. COLUMBIA 107, DECEMBER 2010 4]

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J. ENTOMOL. SOc. BRIT. COLUMBIA 107, DECEMBER 2010 43

Dominant bacteria associated with broods

of mountain pine beetle, Dendroctonus ponderosae

(Coleoptera: Curculionidae, Scolytinae)

RICHARD S. WINDER'”, DONNA E. MACEY'! and JOE CORTESE?

ABSTRACT

Mountain pine beetle (MPB) is the most damaging insect of mature pine forests in west-

ern North America. The current outbreak in British Columbia is the largest ever re-

corded. During a survey of beetle occurrence, well-established infestations were sam-

pled in central B.C. and found to possess larval mortality. Bacteria or other microbes

were among the potential causes of the mortality. Bacteria were isolated from living

larvae and adults, as well as larval and adult beetle cadavers found in bark samples. Bac-

teria were identified by fatty acid methyl ester (FAME) analysis, which indicated 32

species of bacteria present in the MPB larvae. The predominant bacteria (Serratia lique-

faciens, S. plymuthica) were detected in about a third of all sampled larvae, regardless of

mortality. Rahnella aquatilis was found in 11% of all larvae examined and was usually

(93%) associated with larval mortality. Interactions between two bluestaining fungal

symbionts of the MPB (Grosmannia clavigera, Ophiostoma . montium) and two of the

isolated bacteria (S. /iguefaciens and R. aquatilis) were assessed. S. liquefaciens and R.

aquatilis both inhibited the growth of beetle-associated bluestain fungi by 72%. The

bluestain fungi did not impede bacterial growth, and both bacteria grew on autoclaved

bluestain mycelium. Combinations of the two bacterial species formed aggregates on

practical-grade (crab) chitin, but there was no aggregation in pure cultures or on the

autoclaved mycelium of G. clavigera or O. montium. These results indicated that the

two bacteria may be capable of aggregation within the insects, and this may have impli-

cations for their combined effects in the beetle. The role of S. liguefaciens and R.

aquatilis in MPB biology requires further investigation.

Key Words: bark beetle, disease, larva, pathosystem

INTRODUCTION

x10° m?’ of merchantable pine (Walton,

2009). The potential for microbial popula-

As associates, antagonists, and patho-

gens, microbes play important roles in the

life cycle of bark beetles (Barras and Perry,

1975), including the Mountain Pine Beetle

(MPB), Dendroctonus ponderosae Hop-

kins. MPB has been responsible for billions

of board feet of timber losses (Anonymous

2005, Bellows eft al. 1998); a current out-

break has spread over a vast area of British

Columbia (BC), killing at its peak ca.

1.41x10° m?> of merchantable pine; during

2008 the outbreak was still killing 3.6-4.3

tions to constrain the beetle is therefore of

potential interest. In most of BC, MPB has

a l-year life cycle, where incipient or epi-

demic populations attack trees en masse.

Young female adults emerge from host

trees during late summer, and initiate the

attack on new host trees by burrowing into

the phloem and tunnelling vertically to

form egg galleries. Hatched larvae overwin-

ter and feed in the phloem. Pathogenic

——$— sSsSSSsS‘C< ace your message here. For maximum impact, use two or three sentences.

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

Victoria, BC V8Z 1M5 Canada

* Email: rwinder@pfe.cfs.nrean.ge.ca

> Alta Vista Management, P.O. Box 15, Tatla Lake, BC VOL 1V0 Canada

44 J. ENTOMOL. SOc. BRIT. COLUMBIA 107, DECEMBER 2010

bluestain fungi associated with the beetles

spread from the galleries; eventually, trees

succumb to this combined attack.

(Safranyik and Carroll, 2006) During the

spring of 2003, a routine survey of larval

and adult MPB in central BC detected the

presence of large numbers of dead larvae.

The larvae were in lodgepole pine (Pinus

contorta Dougl. ex Loud.) trees containing

well-established broods. Although — the

patchy occurrence of very cold tempera-

tures in the region was suspected as one

possible cause for the mortality, the atypi-

cally darkened and distended hindguts of

the larvae suggested that microbial factors

should not be ruled-out.

Bluestain fungi are microbes that have a

close relationship with the MPB life cycle;

they also figure prominently in interactions

with other beetle-associated microbes. MPB

possesses a specialized mycangium that

carries the inoculum of bluestain fungi from

tree to tree (Whitney and Farris 1970). The

main fungi associated with MPB are Gros-

mannia clavigera (Robinson & Davidson)

Zipfel, deBeer & Wingf. and Ophiostoma

montium (Rumbold) von Arx, and _ their

relative abundances in populations varies

(e.g., Kim et al. 2005, Six and Bentz 2007,

Bleiker and Six 2007, 2009). As_ larval

broods develop, the fungi colonize the

phloem and sapwood (Whitney 1971,

Bleiker and Six 2009). Mining larvae may

ingest phloem colonized by hyphae and

teneral adults consume spores lining pupal

chambers if present prior to emergence

(Whitney 1971, Bleiker and Six 2007,

2009). These fungi are reported to have

mutualistic and antagonistic relationships

with bacteria and yeasts found in the beetle

galleries (Adams et al. 2008).

Natural enemies of Dendroctonus spp.

are diverse (Bellows ef a/. 1998, Bushing

1965, Dahlsten 1982, Moore 1971, 1972a,

1972b). They play a role in holding MPB

populations in check during their endemic

phase (Moeck and Safranyik 1984, Hof-

stetter et al. 2006). Some natural enemies of

MPB have been studied as potential bio-

logical control agents, for example the in-

sect pathogen Beauvaria bassiana

(Balsamo) Vuillemin (Hunt ef al. 1984).

Various options have been suggested for

augmenting natural biological control ef-

fects, including the inundative release of

microbes that may be antagonistic to the

bluestain fungi (Safranyik et al. 2002). The

objective of this research was to determine

the dominant bacteria present in living and

dead larvae and adults of MPB and to char-

acterize interactions between the two prin-

cipal bluestain fungi associated with MPB

and the dominant bacteria associated with

larvae. Our hypothesis was that different

bacteria would predominate in living vs.

dead insects, and that bacteria predominat-

ing in dead insects would be inhibitory to

microbial associates of the living beetles,

i.e. bluestain fungi and predominant bacte-

ra.

MATERIALS AND METHODS

Isolation of Bacteria and Fungi from

Beetles. During the first week of March,

2003, Rectangular 20x30-cm slabs of sap-

wood with the bark and phloem intact were

cut with a chainsaw from five randomly-

selected beetle-attacked lodgepole pine

trees at each of ten sites (Table 1, Fig. 1).

The slabs were taken at points 1.3 m from

the ground at a randomly selected aspect

(north, south, east or west). The geographic

position of each site was recorded with a

GPS unit, and site moisture (humid, mesic,

or xeric) and the percentage of currently

attacked trees in the surrounding stand were

visually estimated. The slabs were stored at

20°C on a laboratory bench for several

days, thus ensuring activity of living larvae.

In the laboratory, the bark was peeled from

the sapwood with a knife and MPB adults

(parental) or larvae were excised with for-

ceps and surface-sterilized according to the

method of Winder and Watson (1994). In-

sect mortality was assessed in each slab as

the percentage of dead larvae and adults

versus the total number of insects present.

Larvae were considered to be dead when

J. ENTOMOL. Soc. BRIT. COLUMBIA 107, DECEMBER 2010

Table 1.

45

Descriptions of sites in British Columbia where mountain pine beetle larvae were collected in

March, 2003.

Site (no.) Location

Elevation (m) Moisture (class) Infested trees (%) Terrain (type)

l Agodak L. 1132 Hydric 50 Flat

2 Quesnel B. 1189 Hydric 60 Flat

3 5100 Rd. 1 1007 Hydric 60 Northern slope

- 5100 Rd. 2 1040 Mesic 70 Southern slope

3 Baker C | 1009 Mesic 80 Western slope

6 Baker C 2 976 Mesic 60 Western slope

7 6600 Rd. 1 979 Mesic 60 Western slope

8 6600 Rd. 2 854 Mesic 50 Eastern slope

9 Redstone | 900 Xeric 10 Flat

10 Redstone 2 859 Xeric 50 Flat

they appeared to have abnormal, distended

hindguts and they failed to move upon firm,

repeated probing with a dissecting needle.

Adults were considered to be dead when

they were overcome with yeasts or fungal

hyphae and non-reactive to probing. Zero to

three live larvae, zero to three dead larvae,

and one to two adults (living or dead) were

selected from each slab, using the number

available to a maximum of three. The lar-

vae and beetles were placed on nutrient

agar (NA) in Petri plates; NA was com-

posed of 8 gL’ Difco Bacto® dehydrated

nutrient broth and 20 gL agar (Sigma).

The insects were tamped downward with

enough force to partially embed them in the

medium. The plates were sealed with paraf-

fin film and incubated at 20+2°C for 7d to

allow microbial colonies to form. A colony

possessing predominant morphology

(colour, size, and growth pattern) was se-

lected from each insect sample and asepti-

cally streaked on NA plates for selection of

pure cultures. The plates were sealed with

paraffin film and incubated at 20+2°C for

1-2 weeks prior to identification. One pure

colony possessing predominant morphology

was selected per insect, for subsequent cul-

turing according to the same method.

Identification of Bacteria. Bacteria

were identified by fatty acid methyl ester

(FAME) analysis using the Sherlock? Mi-

crobial Identification System (MIDI Inc.,

Newark, DE, USA). Pure colonies were

transferred by quadrant-streaking onto

plates of BBLO Trypticase Soy Broth agar

(TSBA) and incubated for 24h at 28+2°C.

This was repeated once again prior to

analysis. Bacterial biomass was harvested

from the third quadrant of the final cultures,

and fatty acids were saponified and methy-

lated according to the protocols provided by

MIDI Inc. for the Sherlock” system. The

resultant fatty acid methyl esters were ex-

tracted into 1:1 (volume-to-volume) hex-

ane: methyl tertiary butyl ether (MTBE)

and analyzed with a gas chromatograph

(S890A Series II with HP-Ultra 2 column,

Hewlett-Packard Co., Mississauga, Ont.,

Canada ) using the TSBA40 method for

aerobic bacteria, also provided by MIDI

Inc. for the Sherlock” system. Extracted

Stenotrophomonas maltophilia (Hugh)

Palleroni & Bradbury (ATCC 13637) and a

hexane:MTBE reagent blank were used as

positive and negative controls, respectively.

Bacterial isolates were identified from com-

parisons made against reference strain data

(Anonymous, 2002) in the MIDI TSBA40

database, version 4.10 (MIDI Inc., Newark,

DE, USA). Using the protocol and _ stan-

dards published by Weyant et al. (1996), a

similarity index (SI) => 0.500 was consid-

ered an acceptable identification to the level

46 J. ENTOMOL. Soc. BRIT. COLUMBIA 107, DECEMBER 2010

EE

FragetLake,

Francois Lake

Morice Lake

Nechako Fiver

Tahtsa Lake Cheslatta Lake

Knewstubb Lake

Uotsa Lake

Whitesail Lake

Eutsuk Lake

West Road River AG

Pog

,\Prince George

Fraser River

2

[5]

a

6

a Quesnel Lake

King Island 4

a

Chika Lake

Wakeman River

Knight Inlet

Kingeame Inlet

Gilford Island

Loughborough Inlet

Redonda Island

Quadra Iskand

‘, Powell Lake

Nootka Island

Texada Island

Vancouver Island

ancouver

Alberni Inlet

= eek cll;

\,Nanaimo (Se esiave Lake _.Chilliwac

\Victoria

Skagit Bay Sauk Fiver

Adruralty Inlet

Figure 1. Sites sampled for mountain pine beetles, with numbered locations corresponding to

Table 1.

of species. SI values > 0.300 were consid-

ered an acceptable identification to the level

of genus, while lower similarity index val-

ues were considered inconclusive. When

comparisons resulted in multiple species or

genera exceeding these identity thresholds,

data corresponding to the highest SI value

were used for the identification. Dendro-

gram cluster analysis based on unweighted

pair-matching of fatty acid profiles

(Sherlock” analysis software, MIDI Inc.,

Newark, DE, USA) was used to explore

relatedness among unidentified isolates.

Bacterial inhibition of Bluestain

J. ENTOMOL. SOc. BRIT. COLUMBIA 107, DECEMBER 2010

Fungi. A nested experimental design was

used to assess interactions between the

three most dominant bacteria: Serratia

liquefaciens (Grimes & Hennerty) Bascomb

et al., Serratia plymuthica (Lehmann and

Neumann) Breed ef al., and Rahnella

aquatilis Izard et al.), and two bluestain

fungi commonly associated with D. pon-

derosae, G. clavigera and O. montium

(Solheim and Krokene 1998). Fungal iso-

lates were obtained from C. Breuil

(University of British Columbia, Vancou-

ver, BC) as cultures growing in Petri Dishes

containing malt extract agar. Small pieces

of these cultures were aseptically trans-

ferred to Petri dishes containing 3:7 (v:v)

malt extract agar:NA. This substrate was

optimal for simultaneous fungal and bacte-

rial growth in preliminary trials combining

different ratios of the two agar media (data

not shown). To account for the variable

effects of cultural moisture on microbial

growth, Petri dishes were treated as experi-

mental units, and the difference in fungal

growth in the presence or absence of bacte-

ria was evaluated in each dish. Half of each

dish was inoculated with a ‘lawn’ of either

S. liquefaciens, S. plymuthica, or R.

aquatilis. In each dish, a 5 mm-diam. agar

plug colonized by either G. clavigera or O.

montium was placed on the bacterial side,

and another was placed on the bacteria-free

side. In each plate, the bacterial inoculum

was taken from one of five randomly se-

lected isolates from each of the three spe-

cies of bacteria collected from the beetles.

Each isolate originated from a different

beetle. Each combination of fungus and

bacterial isolate was replicated five times.

The plates were incubated for 7 d at 20°C.

The radial growth of fungal colonies was

measured after incubation and percentage

of growth inhibition was calculated for each

plate by dividing the maximal radius of the

fungal colony on the bacterial side by the

maximal radius of colony in the bacteria-

free area, and multiplying by 100.

Interactions between bacteria and

chitin-containing substrates. A second

completely randomized experiment was

used to assess the interactive effect of bac-

47

teria and chitin-containing substrates on

bacterial growth and aggregation, because it

is possible for bacterial consortia to affect

insect health (Hentzer and Givskov 2003)

or ice nucleation (Pierson ef al. 1998) and

therefore cold tolerance. A sterile transfer

loop was used to aseptically transfer bacte-

ria from cultures of R. aguatilis and S.

liquefaciens to sterile water (100 mL) in

250 mL Erlenmeyer flasks, and the bacte-

rial concentration in each case was diluted

to 1 x 10° bacteria mL’! based on measure-

ments with a Petroff-Hausser counting

chamber (Hausser Scientific Partnership,

Horsham PA) and a microscope (1000 X).

Equal parts of the bacterial suspensions

were combined to make a third combined

bacterial suspension, resulting in a concen-

tration of 0.5 x 10° bacteria mL’! for each

bacterial species. Each of the three suspen-

sions was aseptically transferred with a

sterile transfer loop to an individual set of

nine sterile aqueous cultures in 250 mL

Erlenmeyer flasks containing 0.1 g (d.w.) of

substrate. In each culture set, the substrate

in three flasks consisted of autoclaved hy-

phae of G. clavigera; autoclaved hyphae of

O. montium was included as the substrate in

three more flasks, and the three remaining

flasks contained chitin derived from crab

Shells (poly-N-acetyl-glucosamine, practi-

cal grade, catalogue number C-7170, Sigma

Chem. Co., St. Louis, MO, U.S.A.). The

hyphae of G. clavigera and O. montium

used in this assay were collected from 250

mL liquid (malt extract broth) cultures,

each of which was inoculated with a Smm

diam. agar plug from a stock culture and

incubated on an orbital shaker (100 r.p.m.,

20°C) for 7d. The hyphae were rinsed in

distilled water, autoclaved, rinsed again,

and dried prior to incorporation into the

substrate assay. A set of nine controls con-

taining only water was also inoculated. The

flasks were secured at a random upright

position on the platform of a rotary shaker.

After 24 h incubation on the shaker (100

r.p.m.) at 20°C, bacterial populations were

assessed as before with the Petroff- Hausser

counting chamber and the microscope. The

microscope (1000X) was also used to ob-

48 J. ENTOMOL. SOC. BRIT. COLUMBIA 107, DECEMBER 2010

serve the degree of bacterial adhesion and

aggregation on solids in the liquid cultures.

The growth data were analyzed using

analysis of variance; no statistical analysis

of aggregation or adhesion was performed

because there was no variance in the re-

sults.

Statistical analysis. Aside from FAME

analysis, all statistical analyses were per-

formed with Statistica 6.1 (Statsoft Inc.,

Tulsa, OK, U.S.A.). For data corresponding

to larval sampling versus larval mortality,

separate one-way analyses of variance

(ANOVAs) were performed. Trunk aspect

(degrees from North) was used as a cate-

gorical factor in one-way ANOVA, while

site was nested as a random factor within

moisture type for a two-factor ANOVA. In

these and subsequent similar analyses,

Levene’s test was used to check for homo-

geneity of variance, and the Newman-Keuls

multiple range test was used to compare

means. Where Levene’s test indicated sig-

nificant (P < 0.05) heterogeneity of vari-

ance, ANOVA was performed after appro-

priate data transformation (arcsinVY).

Where transformations were unable to re-

move significant (P < 0.05) heterogeneity

of variance, non-parametric comparisons

were employed. Larval mortality was sub-

jected to regression analysis, using percent

mortality as the dependant variable and

either elevation or percentage of green trees

attacked as independent variables. Student’s

t test for dependent samples was also used

to compare hypothetical (expected) mortal-

ity values with mortality associated with the

five most dominant bacteria. The expected

values, derived from observed overall mor-

tality rates, were generated by apportioning

50% of expected larval observations as

dead, and 97% of expected adult observa-

tions as dead. For the experiment assessing

fungal growth vs. bacterial inoculations,

data were subjected to an ANOVA for

nested factors. Bacterial isolates were

treated as a random categorical variable

nested within bacterial species, with five

isolates per bacterial species, each repli-

cated five-fold. Bacterial species was

treated as a categorical variable nested

within fungal species (3 per fungal species).

A series of chi-square tests were utilized to

compare the effect of chitinous substrates

on the growth of S. liquefaciens and R.

aquatilis, wherein the expected values were

the mean growth in controls or mean

growth in each of the substrates.

RESULTS

Isolation of bacteria from_ beetles.

Characteristics of the sample sites are pro-

vided in Table 1. The slabs provided a total

of 67 dead MPB adults, 2 living adults, 110

living larvae, and 112 dead larvae. All

adults were mature (parental), and there

were no exit holes apparent in the bark. A

typical living larva is shown in Figure 2. Of

the forty-four slabs containing insects, |

lacked larvae and seven lacked adults. Lar-

val mortality in the slabs ranged from 0 to

100%. Larval mortality at hydric sites

(54.3%) was not significantly different

from mortality in mesic sites (60.4), but

mortality in the both of these site types was

significantly greater (p =0.018) than mortal-

ity in the xeric sites (24.1%). There was

also a significant (p = .039) effect of site

location on mortality (Figure 3). There were

no significant differences in mortality at-

tributable to trunk aspect (p = 0.63). There

were no significant regression trends for

mortality versus elevation (p = 0.32) or

mortality versus the percentage of green

trees attacked (p = 0.686).

Identification of bacteria. Bacterial

colonies were produced in 55% of the bee-

tles sampled. This resulted in 161 pure cul-

tures, of which 130 provided sufficient

growth on TSBA medium for FAME analy-

sis. Twenty-seven species of bacteria were

identified in the beetles based on matches to

reference strains in the MIDI TSBA40 li-

brary (Table 2). Eleven isolates had SI val-

ues less than 0.300, and were categorized as

unidentified. Dendrogram cluster analysis

separated the unidentified isolates into 5

species-related unknown groups based on

J. ENTOMOL. SOc. BRIT. COLUMBIA 107, DECEMBER 2010

49

Figure 2. A typical living mountain pine beetle larva included in the cultural isolations.

100

RY

= f

se \

2 \

is 60 NS

3

E

3

>

S40

20

0

Hydric

Site

Agodak L.

Quesnel B.

5100 Rd. 1

5100 Rd..2

Baker C-1 ]

Baker C-2 :

i Mo

6600 Rd. 1

6600 Rd. 2

Redstone 1 |

. Redstone 2

{0

OONOAR Wh =

i)

a:

Mesic Xeric

Site Moisture

Figure 3. The effect of site location and moisture on the mortality of 232 mountain pine beetle

larvae in the Southern Interior of B.C., March, 2003. Site numbers correspond to the sites listed

in Table 1.

their FAME profiles. Five of six isolates in

one of the unidentified species groups were

found in dead larvae. Serratia spp. were

frequently isolated from larvae (living and

dead), possibly indicating a role in the in-

sect’s normal gut microflora. R. aquatilis

was also frequently isolated (3 adults, 9

larvae). The incidence of R. aquatilis was

widespread, occurring at eight of the ten

sites and in 11% of the insects, but it was

mainly (93%) found in larval cadavers.

Other bacteria occurring principally in dead

insects were less frequently isolated, al-

though mortality associated with Pseudo-

monas syringae vat. tabaci (Wolf & Foster)

Young et al. was significant (p = 0.01) in

comparison to expected mortality (Table 2).

Twelve bacterial species were isolated from

50 J. ENTOMOL. Soc. BRIT. COLUMBIA 107, DECEMBER 2010

Table 2.

The incidence of bacterial species isolated from mountain pine beetle, and the mortality of associated

adults and larvae.

Isolates Isolates Total Beetle

in living in living sites mortality

(and dead) (anddead) (no.) (%)"

larvae (no.)° adults (no.)°

Similarity Incidence

index (%)' (%)

Species

Serratia liquefaciens i228 22 11 (10) 0 (7) 8 61

Serratia plymuthica 16:8 17 7 (8) 1 (6) 7 62

Rahnella aquatilis 89 + 5 11 ELOY OG) 8 0335

Pseudomonas syringae Vv. tabaci 91 +2 5 0 (4) 0 (2) 3 86 *

Hafnia alvei 79 +6 5 1 (3) 0 (2) 4 83

Pantoea citrea 7644 - 1 (2) 0 (2) 4 60

Enterobacter pyrinus Tas 3 2 (2) 0 (0) 3 50

Erwinia chrysanthemi 159 3 0 (0) 0 (4) 3 100

biotype [V/VI

Proteus vulgaris GC supgp. A 7447 1 (0) 0 (1) 2 60

Kluyvera ascorbata 62+ 8 2 1 (0) 0 (2) 2 67

Brevibacillus agri 86 + 0 2 0 (3) 0 (0) 2 100

Pseudomonas syringae V. 9142 2 21) 0 (0) l oe

phaseolicola

Paenibacillus lentimorbus 64+ | 2 L(t) 0 (0) l 50

Salmonella typhimurium GC t2d13 2 2(0) 0 (0) 2 0

subgp. B

Sphingobacterium multivorum 8446 2 0(1) 0 (1) l 100

Kluyvera intermedia (e: l 0 (0) 0 (1) | 100

Pseudomonas fluorescens 92 l 0(1) 0 (0) l 100

biotype G/C

Serratia grimesii Grimont 81 l 0 (1) 0 (0) l 50

Bacillus megaterium subgp. A 92 l 0 (1) 0 (0) l 100

Cedecea davisae 65 l 0(1) 0 (0) l 100

Chromobacterium violaceum 89 l 0 (0) 0 (1) l 100

Pantoea agglomerans GC 1 l 0 (1) 0 (0) l 100

subgp. |

Klebsiella pneumoniae vat. 85 l 0 (1) 0 (0) l 100

ozaenae

Raoultella terrigena 59 l 0 (0) 0 (1) l 100

Pseudomonas putida biotype B 85 l 0(1) 0 (0) l 100

Salmonella cholerasuis v. 67 l 1 (0) 0 (0) | 0

arizonae

Sphingomonas paucimobilis o7 l 0 (1) 0 (0) l 100

Xenorhabadus nematophilus a l 0 (0) 0 (1) l 100

' Where there was more than one observation, indices in this column are shown as the mean + s.d.

* The incidence percentage is calculated as (number of beetle hosts / number of beetles tested) x 100.

* In this column, the number of isolates found in living insects is followed by the number of isolates

found in dead insects, in parentheses.

* Percentages followed by an asterisk are significantly different (p < 0.01, Student’s t test for depend-

ent samples) than expected mortality (larvae = 51%; adults = 100%).

J. ENTOMOL. Soc. BRIT. COLUMBIA 107, DECEMBER 2010

bark samples also producing insects with R.

aquatilis; these included three isolates of P.

syringae vat. tabaci (Table 3).

Bacterial inhibition of bluestain fungi.

There were significant differences in

growth inhibition for parameters corre-

sponding to fungal species (F'; = 107.4, p =

0.000000) and bacterial species (F’> = 3.2, p

= 0.045270). Bacterial isolates within spe-

cies had no significant impacts (F')2= 1.5, p

=0.125199). Bacterial inhibition of O. mon-

tium was nearly twice the inhibition experi-

enced by G. clavigera. Bacterial species

produced similar levels of inhibition against

the growth of O. montium. S. liquefaciens

was significantly (p < 0.05) less inhibitory

towards the growth of G. clavigera (Table

4). Caution is warranted in quantitative

interpretation of these results, in that vola-

tile compounds from the microbes could

permeate throughout the plates, affecting

the scale of the response in both controls

and treatments. The relative responses,

however, indicate a potential for inhibition.

51

Interactions between bacteria and

chitin-containing substrates. Chi-square

comparisons indicated that S. liquefaciens

and R. aquatilis thrived on autoclaved my-

celium and, to a greater degree, on practical

grade chitin (Table 5). All cultures with

combined inocula appeared to have ap-

proximately equal proportions of each bac-

terial species when cell morphology was

observed under a compound microscope.

Combined cultures generally performed as

well as individual cultures, with no clear

indication of a competitive advantage for

either of the two bacteria. The bacteria did

not adhere to the smooth surfaces of the

autoclaved mycelia, but R. aquatilis would

occasionally adhere to minute rough areas

on particles of the practical grade chitin.

These rough areas appeared to serve as loci

for adhesion and aggregation of the bacte-

ria. When R. aquatilis was combined with

S. liquefaciens, bacteria adhered to many

more sites on the chitin particles, and nu-

merous small bacterial aggregates formed

(Table 5).

DISCUSSION

Although it was not isolated from every

dead larva, R. aquatilis appears to be asso-

ciated with mortality in MPB. Parental

adults had an expected mortality of 100%

during the winter; R. aquatilils is therefore

not expected to have caused any increased

mortality in adults. R. aquatilis is an enteric

bacterial species that occurs widely in water

and soil environments (Berge ef a/. 1991,

Heulin et al. 1994, Horie et al. 1985), and it

has been detected from a variety of insects,

including bark beetles (Vasanthakumar ef

al. 2006; Delalibera et al. 2005). There are

reports of this species linked to opportunis-

tic bacterial infections in humans (Caroff e¢

al. 1998, Lebessi et al. 1990, Maraki ef al.

1994, Matsukura et al. 1996, Oh and Tay

1995), and some strains are reported to pos-

sess antagonistic properties against bacte-

rial plant diseases (Laux ef al. 2002, 2003).

Regarding quantification, the frequencies

reported here are for surface-sterilized in-

sects receiving a moderate amount of com-

pression during placement on agar. This

was sufficient to express hindgut contents

onto the isolation medium, but a specific

method for extraction of gut contents could

have generated higher incidence statistics

for bacteria tending to aggregate inside the

larvae. Using other isolation methods, for

example different isolation media or serial

dilutions of ground tissue, would also yield

other non-dominant bacteria and a greater

diversity of species. The results thus pertain

to the proportion of insects where the par-

ticular bacterial species are dominant,

rather than quantifying the actual propor-

tion of insects associated with the bacteria.

Some genotypic and phenotypic hetero-

geneity is reported for R. aquatilis (Brenner

et al. 1998, Pokhil 1998, Selenska-Pobell ef

al. 1995, Varbanets et al. 2004); the bio-

logical role of isolates found in MPB re-

mains an open question. For example, fur-

ther study would be needed to determine

whether the bacterium is pathogenic, or

nn

i)

J. ENTOMOL. SOC. BRIT. COLUMBIA 107, DECEMBER 2010

Table 3.

Bacterial incidence in living and dead larvae of Mountain Pine Beetle, where larvae in the

same bark sample were also infested with Rahnella aquatilis.

Species Class Order Isolates Site(s)'

Paenibacillus lentimorbus Bacilli Bacillales 2 S

Pantoea agglomerans GC Gammaproteobacteria Enterobacteriales l 8

subgroup

Enterobacter pyrinus Gammaproteobacteria Enterobacteriales l 6

Erwinia chrysanthemi Gammaproteobacteria Enterobacteriales 3 5,6. L0

biotype VI

Hafnia alvei Gammaproteobacteria Enterobacteriales l 10

Proteus vulgaris GC Gammaproteobacteria Enterobacteriales l

subgroup

Pantoea citrea Gammaproteobacteria Enterobacteriales l 3

Pseudomonas fluorescens Gammaproteobacteria Pseudomonadales l 10

biotype G

Pseudomonas syrningae var. Gammaproteobacteria Pseudomonadales 3 6

tabaci

Serratia liquefaciens Gammaproteobacteria Enterobacteriales 3,,6,8

Serratia plymuthica Gammaproteobacteria Enterobacteriales 7 3, 658

Sphingomonas paucimobilis _Alphaproteobacteria Sphingomonadales l 5

' Original collection sites listed in Table 1

Table 4.

The effect of bacterial species with fungal species assayed for growth inhibition.

Bluestain Fungus

O. montium R. aquatilis

S. liquefaciens

S. plymuthica

G. clavigera R. aquatilis

S. liquefaciens

S. plymuthica

Bacterial species

% Inhibition of fungal growth’

74.1 a

B2a

68.2 a

46.0 be

39

46.2 be

' Means in this column followed by the same letter are not significantly different according to

the Newman-Keuls multiple range test (p > 0.05). The test was performed on transformed data

(arcsinVY); actual %Inhibition is shown.

simply an opportunist that flourishes after

the larvae succumb to viruses, nematodes,

or other stresses not screened in this study.

The three most dominant bacteria inhib-

ited the growth of the bluestain fungi. None

of the bacterial species were significantly

more inhibitory versus either bluestain fun-

gal species. None of the bacterial species

aggregated on autoclaved mycelia of the

fungi. These results are interesting, in that

bacteria are thought to mediate or inhibit

the growth of fungi associated with two

other Dendroctonus spp.: D. frontalis

Zimmermann (southern pine beetle) and D.

rufipennis (Kirby) (spruce beetle) (Cardoza

et al. 2006, Scott et a/. 2008). Further tests

J. ENTOMOL. SOc. BRIT. COLUMBIA 107, DECEMBER 2010

53

Table 5.

The effect of chitin-containing substrates on the growth, adhesion and aggregation of Serratia

liquefaciens and Rahnella aquatilis.

Substrate S. liquefaciens' R. aquatilis' Density Adhesive Aggregating

(10° bacteria cultures cultures

mL") (no.) (no.)

None + 0.36a 0 0

0.32 ac 0 0

0.19 a 0 0)

G. clavigera 5.24 be 0) 0)

(autoclaved hyphae) 4.06 be 0 )

D222 OC 0 )

O. montium 7.88 ¢ 0 0

(autoclaved hyphae) 6.09 be 0 0

$.23:¢ 0 0

Crab shell 10.44 d 0 0

15.34 d 2 0

+ 16.69 d 3 3

' A plus sign (+) in these columns indicates cultures were inoculated with the species listed in the

column heading.

* Means in this column followed by the letter ‘a’ are not significantly different from the expected

control concentration of 0.29x10° bacteria mL’! using the Chi-square test (p > 0.05). Means fol-

lowed by the letter ‘b’ are not significantly different from the expected concentration for growth on

hyphae of G. clavigera (4.77x10° bacteria mL’'), means followed by the letter ‘c’ are not signifi-

cantly different from the expected concentration for growth on hyphae of O. montium (7.73 x10°

bacteria mL’'), and means followed by the letter ‘d’ are not significantly different from the expected

concentration for growth on Crab shell (15.167.73 x10° bacteria mL”), also using Chi-square tests

(p > 0.05).

would be necessary to understand the cause

of the inhibition. Nutrient depletion, pH

changes, or various secondary compounds

are all examples of possible inhibitory fac-

tors. Although R. aquatilis is reported to

grow well on wood (Kallioinen ef a/. 2003),

proliferation of bluestain fungi in the beetle

galleries suggests that the enteric bacteria

primarily inhabit the insect gut, where they

would have limited impact on the spread of

the fungi. Another study has shown that the

growth of O. montium is stimulated by mi-

crobes isolated from the galleries of MPB,

including three yeasts (Candida sp., Pichia

scolyti, and an unidentified basidiomycete)

and a bacterial species (Micrococcus sp.).

However, the same study also found that

Candida sp., the basidiomycete yeast, and

Micrococcus sp. were inhibitory to the

growth of G. clavigera. Isolated from fresh

phloem near MPB attacks, Bacillus pumilus

is also reported to be inhibitory to the

growth of both fungi (Adams ef a/. 2008).

There is probably a diverse range of antago-

nistic and mutualistic interactions among

the various microbes that associate with

MPB.

The adherence and aggregation of R.

aquatilis and Serratia spp. on arthropod

chitin could indicate a role for microbial

consortia in the observed beetle mortality.

Insect hindguts can possess chitinous struc-

tures that facilitate digestion through bacte-

rial aggregation (Hackstein and Stumm

1994). R. aquatilis and Serratia spp. are

involved in biofilm formation (Steidle ef al.

2001). Further research is needed to under-

stand the prevalence of R. aquatilis/Serratia

54 J. ENTOMOL. SOc. BRIT. COLUMBIA 107, DECEMBER 2010

spp. consortia in the environment and how

R. aquatilis becomes established in MPB

populations. The abundant growth of S.

liquefaciens and §S. plymuthica on auto-

claved mycelium of Ophiostoma spp.

agreed with the reported chitinolytic prop-

erties of these species (Berg ef al. 1999,

Joshi et al. 1988). Presumably, larval MPB

could benefit from any digestive action of

these bacteria on bluestain fungi. However,

other bacterial species could also persist in

the R. aquatilis/S. liquefaciens consortium,

potentially creating chronic stress in the

insect host or affecting cold-tolerance. In

this study, slabs producing specimens with

R. aquatilis also produced specimens con-

taining Pseudomonas spp., an ice-

nucleating species (Lee et al. 1998).

The association of Serratia spp. with

living beetles agrees with a previous study

of D. frontalis, where bacteria isolated from

the gut of healthy insects included Serratia

spp. (Moore, 1971). Earlier research on

southern pine beetle correlated the occur-

rence of several bacterial species with bee-

tle mortality, including Serratia marcescens

Bizio, Pseudomonas aeruginosa

(Schroeter) Migula, Pseudomonas fluores-

cens Migula, Bacillus thuringiensis Ber-

liner, Bacillus cereus Frankland & Frank-

land, and Flavobacterium sp. (Moore

1971). In further tests on southern pine bee-

tle, pathogenicity was demonstrated for S.

marcescens, the Pseudomonas spp., and the

Bacillus spp. (Moore 1972b).

Without further in vivo testing, it is dif-

ficult to assign a definite pathogenic or op-

portunistic role for R. aquatilis in larvae or

adults of MPB. A pathogenic form of R.

aquatilis might have utility as a biological

control agent, since the species is motile

and could potentially penetrate beetle gal-

leries during wet weather. On the other

hand, even if R. aquatilis is simply an op-

portunist that aggregates with species such

as S. liquefaciens, this mechanism of action

might still be exploited to reduce the cold

tolerance of the insect. Answers are needed

regarding the natural frequency of particu-

lar bacterial associations, but the deliberate

introduction of aggregating bacteria along

with ice-nucleating species could reduce the

cold tolerance of MPB. This could be espe-

cially beneficial if climate warming contin-

ues to exacerbate the beetle impacts. In any

eventual effort to develop biological control

agents, the occurrence and influence of

other pathogens in larvae (viruses, nema-

todes, etc.) should also be explored. The

geographic occurrence of R. aquatilis

should also be surveyed and its mode of

distribution within brood trees elucidated.

A more complete understanding of micro-

bial influences and constraints on MPB

may help us deal with future outbreaks.

ACKNOWLEDGEMENTS

Laboratory research was funded by the

Mountain Pine Beetle Initiative of Natural

Resources Canada and the Canadian Forest

Service Pest Management Methods Net-

work. Original beetle surveys were funded

by the BC Ministry of Forests and con-

ducted by Alta Vista Management of Tatla

Lake, BC. Beetle collection was funded and

conducted by Alta Vista Management. We

thank Jocelyn Joe-Strack for technical as-

sistance, Dr. Les Safranyik (PFC) for tech-

nical advice, and Dr. Colette Breuil (UBC)

for providing fungal cultures. We also

thank several anonymous reviewers for

their constructive comments.

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J. ENTOMOL. Soc. BRIT. COLUMBIA 107, DECEMBER 2010

Cereal leaf beetle, Oulema melanopus (L.) (Coleoptera:

Chrysomelidae), attraction to oat plantings of different ages

GEORGE D. HOFFMAN’” and SUJAYA RAO!

ABSTRACT

The cereal leaf beetle (CLB), Oulema melanopus, a serious pest in oats, barley and

wheat, is a relatively new pest on the west coast of North America. To determine if

adults showed a preference for oat stands of different ages, we examined adult and egg

densities in four sequentially planted oat stands in the Willamette Valley, Oregon, in

2005 and 2006. Adults moved from earlier to later plantings (from older to younger

oats) during the growing season, particularly once the flag leaf had emerged in earlier

plantings. In 2005, the seasonal pattern in egg counts tended to match that of adult

counts in the first three oat plantings. The egg to adult ratio was greater in the earlier

planted (older) oats, particularly the first planting. The egg to adult ratio was more vari-

able in 2006. Adults spent the most physiological time (degree-days) in the second oat

planting, and total egg numbers were highest in the second and third plantings. Data

suggest that delayed planting as a trap crop management tool for CLB is complex and

potentially ineffective.

Key Words: cereal leaf beetle, Ou/ema melanopus, plant age, host attraction, egg pro-

duction, management

INTRODUCTION

57

The cereal leaf beetle (CLB), Oulema

melanopus (L.) (Coleoptera: Chrysomeli-

dae), is a new pest in cereals in the western

United States (Rao et al. 2003). CLB was

first detected in Michigan in 1962 and soon

became a serious pest of small grains in the

Midwest, the Atlantic States, and eastern

Canada. Wheat (Triticum aestivum L.), bar-

ley (Hordeum vulgate L.), and oats (Avena

sativa L.) were damaged (Haynes and Gage

1981), with yield reductions in oats reach-

ing 30% (Wilson et al. 1969) to 48.8%

(Merritt and Apple 1969). Incorporation of

trichome resistance in experimental wheat

varieties reduced CLB numbers, although

effective trichome resistance was not pre-

sent in other grains (Haynes and Gage

1981). Biological control efforts were initi-

ated and, once the gregarious larval parasi-

toid Tetrastichus julis (Walker)

(Hymenoptera: Eulophidae) and the egg

parasitoid Anaphes flavipes (Foerster)

(Hymenoptera: Mymaridae) became estab-

lished, damaging populations requiring

chemical control measures were substan-

tially reduced (Haynes and Gage 1981).

CLB was first detected in Oregon and

Washington in 1999 (Rao eft al. 2003),

where it caused direct damage to cereal

crops. In addition, quarantine restrictions

were established on movement of hay and

forage from infested counties in Oregon

and Washington to neighboring California

and Canada due to potential transport of

adult CLB in baled straw. As the pest

moved into the region, attention was di-

rected towards developing improved moni-

toring tools (Rao et al. 2003) and control

tactics. The parasitoid 7. julis is not wide-

spread in the Pacific Northwest and 4.

flavipes has not yet established.

CLB has one generation per year. Eggs

are laid in spring and the larvae develop

through four instars by early summer. Pupa-

' Department of Crop and Soil Science, Oregon State University, Corvallis, Oregon 97331.

> E-mail: george. hoffman@oregonstate.edu

58 J. ENTOMOL. Soc. BRIT. COLUMBIA 107, DECEMBER 2010

tion occurs in earthen cells in the upper 5

cm of soil and adults emerge after two to

three weeks after pupation begins. The

overwintering adult population dies by mid-

June, several weeks before emergence of

the first summer adults. New adults feed to

a limited extent but do not mate. Adults

overwinter within fields if there is a large

amount of crop residue, or outside fields in

sheltered fencerows and woodlots (Haynes

and Gage 1981).

There is no consistent pattern in spring

movement of CLB in relation to the age of

the host plant. Adults typically move from

overwintering sites to grasses and winter

wheat and then to emerging spring-planted

grains. In winter wheat, which 1s planted in

fall, CLB adults prefer later plantings

(Casagrande et al. 1977) and more eggs and

larvae are found on these plants (Gage

1974). The opposite 1s found in oats, which

are planted in spring. Early plantings of

oats have more CLB adults (Casagrande et

al. 1977), eggs and larvae (Gage 1974). For

unknown reasons, adult preference for win-

ter wheat versus spring grains (primarily

oats) can change over several years

(Casegrande ef al. 1977), suggesting that

relationships between plant species, plant

age, and CLB adult preference are not static

(Haynes 1973).

In this study we examined the dynamics

of CLB adult and egg densities in oat stands

of four ages in the Willamette Valley, Ore-

gon. To assess the potential impact of CLB

in these stands, we determined the total

time adults spent in each stand and the total

number of eggs laid. The potential role of

host-plant selection behaviours in CLB

management strategies is discussed.

MATERIALS AND METHODS

The study was conducted in 2005 and

2006 at the Hyslop Field Laboratory, Ore-

gon State University, in a field of oats

(variety Cayuse). This field was adjacent to

an oat nursery used by USDA-APHIS and

Oregon Department of Agriculture (ODA)

for propagating 7. ju/is. Four stands of oats

(Cayuse) were planted sequentially on 15

March, 5 April, 26 April, 2 June, 2005; and

7 April, 20 April, 10 May and 6 June, 2006.

In 2005, the fourth planting was delayed

due to the high rainfall in May. In 2006,

limited rainfall in late April and May de-

layed germination of the second planting,

so the second and third plantings were irri-

gated until rains started again in late May.

In both years, the four treatments (called

planting dates PD1 through PD4 in chrono-

logical order) were set up as a randomized

complete block design with four replicates.

Each plot was 6.85 m wide and approxi-

mately 42.5 m long, and consisted of 45 oat

rows,

We monitored CLB adult and egg popu-

lations in the plots throughout the overwin-

tered adult activity period (mid-April

through late June). Sampling began as soon

as the first adults were seen in the plots and

continued at weekly intervals until adults

were no longer present in the field. In 2005,

adults and eggs were counted in ten ran-

domly located subsamples (30.5 cm of row)

per block. In 2006, we took five subsamples

per block, and low CLB populations neces-

sitated increasing the subsample area for

adults to five adjacent rows (30.5 cm sec-

tions), and two adjacent rows (30.5 cm sec-

tions) for eggs. The sampling regime ended

before the emergence of new adults of the

next generation. Adult females and males

were not differentiated in 2005. In 2006, a

representative number of adults was col-

lected from the plots for identification of

sex. In 2006, we recorded the plant devel-

opment stage using the Zadok’s Scale

(Zadoks et al. 1974) to allow for compari-

sons based on oat plant growth stage.

To calculate the total developmental

time adults spent in each of the planting

date treatments we transformed the data to a

centigrade degree-day (CDD) time scale.

Expressing the data on this scale eliminates

the effect of variable temperatures on be-

havior and oviposition, whether seasonal or

weekly. Cumulative centigrade degree-days

(CCDD) were calculated using temperature

J. ENTOMOL. Soc. BRIT. COLUMBIA 107, DECEMBER 2010

data from the Hyslop weather station, and

the CLB development thresholds of 7°C

(minimum) and 30°C (maximum) (Guppy

and Harcourt 1978). The biofix (start)

dates, 6 March in 2005, and 2 April in

2006, for CDD accumulation were based on

the date of first adult emergence as pre-

dicted by the CLB IPM weather model de-

rived from a synthesis of 10 data sets (IPPC

2009). Adult sample counts were plotted on

the CCDD scale, and the graphical method

(Southwood 1978) with 20-DD intervals

was used to plot the points. The values of

cumulative adult degree-days (CADD) per

30.5 cm of row were calculated by sum-

ming the area under the curve.

CLB eggs take about two weeks to

hatch at spring temperatures in the Wil-

lamette Valley. Hence, the same egg could

be counted at two or three weekly sampling

periods. To calculate the actual total num-

ber of eggs laid in each plot over the ovi-

positional period, egg counts were plotted

using the graphical method (Southwood

1978) at 20-DD intervals. The cumulative

egg degree-days was divided by the number

of degree-days it takes for an egg to hatch

(105 CDD; Guppy and Harcourt 1978) to

obtain the total number of actual eggs per

sample unit (Southwood 1978).

Egg production per adult was calculated

by dividing the number of eggs at a given

number of accumulated degree-days by the

number of 20-DD intervals over which the

eggs could have accumulated (5 intervals

maximum). This number was then divided

by the running average of adults over corre-

sponding time period. This value was not

calculated for the fourth planting because of

the minimal adult and egg density data.

Adults can move between plots, which

raised the question of whether these experi-

mental units were independent. This ques-

tion was addressed by examining the spatial

autocorrelation variogram for egg and adult

count data on each sampling date (SAS

Tel):

Repeated-measures analysis was used to

29

adjust the p-values for temporal autocorre-

lation present in the within-sampling-date

comparisons for differences in egg and

adult counts among planting dates. Because

counts of adults and eggs increase and then

decease over time we are not interested in

the main effects of Julian date and planting

date; and effects of planting date are more

appropriately analyzed using the cumula-

tive degree-day approach. Count data often

fit the Poisson distribution and the compari-

son of adult counts among planting dates

was conducted using PROC Genmod, with

parameter options link=log dist=poisson

(SAS 9.1). However the low power of this

analysis due to the small number of data

clusters, and the deviance from the Poisson

that occasionally resulted when using the

subsample means, led us to analyze the egg

count data differently. Egg counts were

transformed using the variance-stabilizing

transformation developed for CLB egg spa-

tial distributions (logl0 (counts + 0.13))

(Logan 1980) and analyzed using repeated

measures in PROC Mixed (SAS 9.1). The

lack of a similar stabilizing transformation

for adults, and the small number of adults in

2006, precluded our using the more power-

ful mixed model approach for the adult

data. For the PROC Mixed and PROC Gen-

mod analysis, we compared planting date

means within sampling dates using unad-

justed probabilities after first testing for the

main effect difference.

To adjust for the statistical problem of

modeling zero variance for adult and egg

counts where we recorded zero individuals,

we dropped these ‘planting date by sam-

pling date’ entries from the data sets before

analysis. Instead, we compared the other

counts on this date to zero counts by deter-

mining if the 95% confidence limits of the

means included zero.

Statistical comparisons of CADD and

total eggs among planting dates were per-

formed using ANOVA in PROC GLM with

the Tukey adjustment for the number of

comparisons (SAS 9.1).

60 J. ENTOMOL. SOC. BRIT. COLUMBIA 107, DECEMBER 2010

RESULTS

There was minimal spatial autocorrela-

tion in CLB stages among plots in both

years. Significant autocorrelation among

plots occurred on only one sampling date

for adults and one sampling date for egg

counts. On all other sampling dates there

was either no spatial autocorrelation or the

correlation was negative (plots at further

distances were more correlated than closer

plots). This indicates that the plots were

independent from each other and statistics

based on the assumption of independent

experimental units can be used. It also sug-

gests that significant movement was taking

place between the oat plots and adjacent

grain fields.

In 2005, overwintered adults appeared

after the first two plantings had emerged,

and most died by 15 June (Table 1). On

each sampling date, CLB adults had a

choice of oats of two or more planting date

(PD) treatments. No adults were counted in

the fourth planting (PD4) on 15 and 24

June, due to the late emergence of oats and

declining population of adults, however, a

few adults were seen outside the rows sam-

pled. Counts of adults in each planting date

treatment generally increased until the fol-

lowing planting date treatment became

more attractive, then decreased in the older

treatment. Statistical differences among

mean counts of adults per planting date

treatment occurred on six of eight sampling

dates where adults were present (Table 1).

Countering this general trend was the de-

cline in adults in the third planting and in-

crease in the second planting during the

third week of May (19 May). This was a

week of unusually intense rainstorms and

we speculate that the adults were seeking

refuge in areas of greater plant biomass. At

this time the plant height of PD2 was 31.4

cm versus 12.0 cm for PD3. The total num-

ber of adults decreased at this time (Table

1) suggesting that some were leaving the

oat plantings to seek refuge in other fields

or fencerows.

In 2006, sampling started just as the first

oat planting emerged. The 2006 adult popu-

lation was approximately one-tenth that in

2005 (Table 1). The rise and decline in

adults over time in each of the plantings

was similar to that in 2005, although the

trends were not as uniform, perhaps due to

the greater variability in the much smaller

populations. There were no statistical dif-

ferences among mean adult counts at P <

0.05. However trends in the 2006 count

data were similar to 2005 (Table 1). There

was a declining trend in adult counts in the

oat plantings on the 25 May sampling date.

Sampling that week occurred after three

days of high rainfall.

When adult population counts over cal-

endar time are transformed to a CDD scale,

the sequential movement of adults from

early- to late-planted oats through the sea-

son can be seen in 2005 and 2006 (Fig. La,

b). The decline in total adults in the plots

each year after several days of heavy rain in

late May is notable. It occurred at approxi-

mately 310 and 350 CDD in 2005 and

2006, respectively. Expressing the data on a

CDD scale also allows calculation of total

adult residency time, and total egg num-

bers, in stands planted at different times.

The analysis of CADD in 2005 showed that

PD2 had the highest CADD, followed by

PD3 and PDI (Table 2). In 2006, the rela-

tive CADDs of the four planting dates were

similar to 2005, although the differences

were not significant (Table 2). Total CADD

in 2005 was 13.88 versus 1.58 in 2006.

On a calendar scale, CLB adults first

appeared in the oat plots during the same

week in 2005 and 2006. In 2006, the large

movement of adults into PD3 coincided

with PDI and PD2 reaching the Zadok’s

scale of 43 and 33, respectively (boot stage

and 3rd node stage) (Table 3). On a degree-

day scale (developmental minimum of 7°C)

the adults appeared 76.4 CDD earlier in

2006 (Fig. 1). This translates to 12 days at

the daily temperatures at this time of year.

An even larger difference occurred, 17

days, when a developmental minimum of

9°C (Fulton and Haynes 1975) was used. In

2005, approximately 110 CDD occurred

between the time overwintering adults were

predicted to emerge, and when they ap-

J. ENTOMOL. Soc. BRIT. COLUMBIA 107, DECEMBER 2010 61

Table 1.

Weekly mean CLB adult counts in each sequentially planted oat stand.

Sampling dates 2005!

Planting 21-4 29-4 6-5 12-5 19-5 24-5 31-5 9-6 15-6 24-6

date

15 March 0.03b 0.13 0.15 0.18b 0.10b 0.10b 0.00b 0.00b 0.00 0.00

5 April 0.30a 0.30 0.30 0.28b 0.40a 0.25b 0.20b 0.00b 0.00 0.00

26 April 0.18 0.60a 0.38a 0.58a 1.33a 0.28a 0.00 0.00

2 June 0.00 0.00

Total 0.33 0.43 0.63 ~~ 1.06 0.88 0.93 153 0.28 0.00 0.00

Sampling dates 2006

21-4 26-4 2-5 10-5 18-5 = 25-5 1-6 8-6 15-6 23-6

7 April 0.02 0.03 0.03 0.02 0.04 0.02 0.00 0.00 0.00 0.00

20 April 0.02 0.04 0.10 0.01 0.05 0.01 0.01 0.00

10 May 0.06 0.09 0.03 0.03 0.00

6 June 0.00

Total 0.02 0.03 0.05 0.06 0.14 0.08 0.14 0.04 0.04 0.00

' Planting date means within year and sampling date with different letters are statistically dif-

ferent at P <0.05.

Table 2.

Cumulative adult degree-days and total egg numbers in the four oat planting dates

Planting date Cumulative adult degree-days (CADD)' Total number of eggs”

2005

15 March 1.53 + 0.40 b 49.0+5.5b

5 April hlot 150a 64.7+6.0a

26 April 4.49+ 0.38 b 63.042.7 a

2-June 000+ 0,00c 1.7+0.4¢

Total 13.88 178.4

2006

7 April 0.3720.10a (Al som eee

20 April 0.65 + :0.23:.a 5.1+0.7a

10 May 0.54+ 0.05 a 49+0.4a

6 June 0.00 + 0.00 b 0.3+0.1b

Total 1.56 16.3

' Means + SE of PD entries. Within years PDs with different letters are statistically different at

P3005:

> Total number of eggs per planting date estimated from the area under the curve analysis.

peared in the oat plots. In 2006, this interval

was 90 CDD (Fig. 1). Thus there are either

errors in the models or CLB adults are on

other hosts (winter grains) for several

weeks before migrating into the oats. We

did not monitor CLB adults on other hosts

early in the season.

In 2005, the temporal pattern in egg

62 J. ENTOMOL. Soc. BRIT. COLUMBIA 107, DECEMBER 2010

a. 2005

b 2006

0.10

CLB Adults per 30.5 cm of row

ai

0.05

000

0 100 200

Planting Date

15 March

ore 5 April

———— 26 April

2 June

Total

300 A400 500 600

Cumulative Centigrade Degree-days

Figure 1. The mean density of CLB adults expressed on a cumulative centigrade degree-day

scale, in oat stands of four age classes, 1.e., planting dates (PD) 1 — 4: (a) 2005; (b) 2006. The

biofix dates were 6 March in 2005, and 2 April in 2006, the dates of first adult emergence pre-

dicted by the CLB weather model (IPPC 2009).

density among planting dates was similar to

that of the adult populations, with egg

counts in each consecutive planting increas-

ing to a peak and then declining as counts

increased in the later plantings. There were

statistical differences in egg counts on eight

of the sampling days (Table 4). The similar

egg counts on 19 May are a consequence of

the movement of adults out of the younger

(smaller) oats, and movement into the older

(larger) oats which occurred that week.

Quantitative changes in egg density did not

always mirror changes in adult density. The

egg density in PD2 was only slightly

greater than that PD1 (Table 4), even

though the adult population was twice as

great in PD2 (Table 1). Similarly, even

though the population of adults in PD3 was

greater than that in PD2, the number of

eggs laid was approximately the same.

These relationships between egg and adult

counts resulted in the eggs per adult values

in PD1 being at times much greater than in

PD2 and PD3 (Table 5).

When expressed on a CDD scale, the

pattern of egg density over time in the four

planting date treatments in 2005 was simi-

lar to the calendar day scale (Fig. 2a). The

total number of eggs was greater for PD2

and PD3 compared to PD1, while few eggs

J. ENTOMOL. Soc. BRIT. COLUMBIA 107, DECEMBER 2010 63

Table 3.

Oat plant growth stages (Zadok’s scale) at three representative periods during the spring sam-

pling period in 2006

Planting date

Sampling date 7 April 20 April 10 May 6 June

2 May 14 t1.5 n/a! n/a

1 June 45 33 i n/a

23 June 87 55 33 14

' n/a - plants not yet emerged. Zadok’s Scale of 14 is a young plant with 4 leaves unfolded; 87

is the hard dough stage of the developing seed.

Table 4.

Weekly mean CLB egg counts in each sequentially planted oat stand

Sampling dates 2005!

21-4 29-4 6-5 12-5 19-5 24-5. 31-5 9-6 15-6 24-6

Planting

date

15 March 0.5b 16.1 30.0a 28.3b 17.5 16.06 4.0c 2.0c 0.2c 0.06

5 April 2.5a 18.4 37.4a 344a 21.0 22.9a 11.0b 3.9b 1.5b 0.1b

26 April 0.0 4.5b 9.6c 19.3 23.6a 40.3a 30.la 8.7a 0.3b

2 June 0.4c 2.0a

Sampling dates 2006'

21-4 26-4 2-5 10-5 18-5 25-5 1-6 8-6 15-6 23-6

7 April O35" * 19". 20a 26 dal De 10 O.1b 0.4b 0.0b

20 April 0.2b 0.9 3.7 1.9 6 O10 -0.2b —Ol1b

10 May 23 18 09a 2.5a 1.0a

6 June 0.3b

' Planting date means within year and sampling date with different letters are statistically dif-

ferent at P <0.05.

Table 5.

Mean CLB eggs per adult at 100 CDD intervals during the adult activity period!

Accumulative CDD from overwintering adult emergence

Planting date

2005

15 March

5 April

26 April

2006

7 April

20 April

10 May

160

Dpeealk

26.9

260

360

23.0

RS

9.5

i all

6.1

20.1

460

16.3

4.6

21

0.6

We

560

n/a

2.4

'5.3

' Number of eggs from Fig. 2 divided by the running average of adults from the previous five

20 CDD periods in Fig. | (adults that could have laid those eggs).

* ‘n/a - adults not found in plots

64 J. ENTOMOL. SOC. BRIT. COLUMBIA 107, DECEMBER 2010

60

40

5

ES

_ of

=

oO

Lr

am)

a]

_ oO

fab)

1.

wn

ra)

fa)

LL =

a]

mM

—I

OO)

4

*)

z:

0

0 100 200

Planting Date

—-—-— 15March

5 April

———— 26 April

2 June

Total

Planting Date

—-—-— 7 April

20 April

———— 10May

6 June

sUU 400 900 600

Cumulative Centigrade Degree-days

Figure 2. The mean density of CLB eggs expressed on a cumulative centigrade degree-day

scale, in oat stands of four age classes, 1.e., planting dates (PD) 1 — 4: (a) 2005; (b) 2006.

The biofix dates were 6 March in 2005, and 2 April in 2006, the dates of first adult emer-

gence predicted by the CLB weather model (IPPC 2009).

were laid in PD4 (Table 2).

In 2006, the temporal pattern of egg

counts did not follow adult densities as well

as in 2005. The sequential increase and

decrease in the three planting dates was not

as uniform. There were significant differ-

ences among planting dates in egg counts

on four sampling days, mostly at the end of

the CLB activity period when PD3 counts

were greater than those in the other planting

dates (Table 4). The movement of adults

from the younger (smaller) oats during the

rains of the week of 25 May disrupted the

sequential pattern of egg increases in the

younger oats. In 2006, the relative differ-

ence among the first three planting date

treatments in the eggs per adult value was

variable (Table 5), reflecting the lack of

correlation between adults and egg counts.

When 2006 egg densities are plotted on

a CDD scale (Fig. 2b), the most noticeable

difference is that the single peak in total

egg density separates into two peaks that

correspond to the peaks in total adults (Fig.

1b). There were no statistical differences in

the total number of eggs in the first three

planting date treatments (Table 2).

J. ENTOMOL. Soc. BRIT. COLUMBIA 107, DECEMBER 2010

65

DISCUSSION

When CLB adults have a choice of oat

stands of different age they tend to prefer

younger plants. Overwintered adults move

into newly emerged oats after spending

time on other hosts, and adult populations

increase over time. Populations on older

oats decline at the time adult counts are

increasing in younger oats. This behaviour

is particularly evident once the older oats

reach the flag leaf stage. The age of oat

leaves affects CLB oviposition, with a dra-

matic decline in oviposition occurring when

a plant approaches the flag leaf stage (GDH

unpublished data). This is in contrast to the

work of Casagrande et al (1977) that found

higher adult populations in older (earlier

planted) oats. This contrasting information

suggests that adult preference for young

plants is not static, and that factors such as

weather conditions and the relative differ-

ences in plant maturity and size can influ-

ence host preference.

There were some differences in the data

between 2005 and 2006, in part due to the

approximately 10-fold smaller CLB popula-

tions in 2006. The 45% parasitism rate of

late season larvae by 7. julis in 2005 (GDH

unpublished data) probably accounts for the

population decline in 2006. Samples in

2006 contained 100% parasitized CLB lar-

vae after the second generation of 7. julis

(R. Worth unpublished data). One differ-

ence between 2005 and 2006 that may have

influenced adult and egg counts is the

spring drought in 2006. While we used irri-

gation to get the second and third plantings

germinated and established, these later

planted stands were probably under greater

water stress than the deeper rooted first

planting. CLB adults may have been re-

sponding to a possible difference in plant

water status.

Observations during greenhouse studies

showed that many CLB adults leave the oat

plants and collect on cage sides between

1000 h and 1600 to 1700 h (unpublished),

and the present study documented periodic

disappearances of a portion of the within-

field population to other habitats. These

observations suggest that CLB adults are

moving within and between fields on a

regular basis and can respond to changing

host plant and environmental cues.

The eggs per adult values calculated

from the CDD data are much greater than

the eggs per female obtained from labora-

tory cage experiments. Laboratory data

ranged from 8.7 to 12.2 eggs per day at

26.7 °C (Wellso et al. 1973). Eight to 18

eggs per day were laid over the first half of

post-aestival adult life (Wellso et al. 1975).

The data from the present study, green-

house observations, and other studies

(Gutierrez et al. 1974, Casagrande ef al.

1977) indicate that CLB adults move fre-

quently within and between fields, and raise

the possibility that a significant portion of

the adults were not on the oat plants during

our mid-day sampling.

In 2005, the impact of CLB was greatest

on the second and third oat plantings. More

cumulative adult degree-days were re-

corded in the second planting, and the high-

est total egg numbers occurred in PD2 and

PD3. The fourth planting was minimally

affected by CLB due to its late emergence

in relation to adult phenology. Similar

trends were found in 2006.

While CLB adults were attracted to

younger oat plants, preference was not ab-

solute. The variation in adult attraction and

egg production in stands of different ages is

in part because older oats are not a uniform

resource for adults or their developing off-

spring. Older oats are actually a composite

of both young and old leaves, so adult CLB

can find young leaves in an older oat stand.

In a greenhouse study, the majority of eggs

were laid on the softer older leaves, or

young tiller leaves of older oat plants

(unpublished).

This study suggests that using oat plant-

ings of different ages as a trap crop to help

control damage for CLB infestations will be

unpredictable and potentially unprofitable.

Damage to the flag leaf causes the greatest

loss in yield (Yoshida 1972) so the opti-

mum timing of consecutively planted stands

66 J. ENTOMOL. Soc. BRIT. COLUMBIA 107, DECEMBER 2010

to draw CLB adults away from the primary

crop will need to be modeled using input

from oat plant growth models, CLB adult

preference related to oat phenology, and

CLB developmental thresholds. In addition,

late-planted oats are likely to have signifi-

cantly reduced yields compared to early

planted oats (Ciha 1983). Therefore, a late-

planted trap crop of oats, even if sprayed to

control CLB, will suffer yield loss.

ACKNOWLEDGEMENTS

Technical assistance in 2006 was pro-

vided by Erin Wycoff. The staff at the Ore-

gon State University Hyslop Experimental

Station planted and maintained the oat

appropriate statistical analyses. Funding for

this project was provided by USDA /

CREES, Grass Seed Cropping Systems for

Sustainable Agriculture.

plantings. Cliff Pereira advised us on the

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beetles. Annals of the Entomological Society of America 70: 19-30.

Ciha, A.J. 1983. Seeding rate and seeding date effects on spring seeded small grain cultivars. Agronomy

Journal 75: 795-799.

Fulton, W.C. and D.L. Haynes. 1975. Computer mapping in pest management. Environmental Entomology

4: 357-360.

Gage, S. H. 1974. Ecological investigations on the cereal leaf beetle, Ou/ema melanopus (L.), and the prin-

cipal larval parasite Tetrastichus julis (Walker). Ph.D. Thesis, Michigan State University.

Guppy, J. C. and D. G. Harcourt. 1978. Effects of temperature on development of the immature stages of

the cereal leaf beetle, Oulema melanopus (Coleoptera: Chrysomelidae). Canadian Entomologist 110:

257-263.

Gutierrez, A.P., W.H. Denton, R. Shade, H. Maltby, T. Burger and G. Moorehead. 1974. The within field

dynamics of the cereal leaf beetle (Oulema melanopus (L.)) in wheat and oats. Journal of Animal Ecol-

ogy 43: 627-640.

Haynes, D.L. 1973. Population management of the cereal leaf beetle. Pages 232-240. In P.W. Geier, L.R.

Clark, D.J. Anderson and H.A. Nix, editors Insects: studies in population management. Ecological Soci-

ety of Australia, Canberra.

Haynes, D.L. and S. H. Gage. 1981. The cereal leaf beetle in North America. Annual Review of Entomol-

ogy 26:259-287.

IPPC. 2009. Online cereal leaf beetle phenology model. Oregon State University Integrated Plant Protection

Center degree-days and phenology modeling website, http://uspest.org/cgi-bin/ddmodel.pl?spp=clb. Ac-

cessed January 26, 2010.

Logan, P.A. 1980. Spatial distribution of cereal leaf beetle (Oulema melanopus (L.)) eggs and larvae and

treatment of count data. Environmental Entomology 9: 186-189.

Merritt D.L., and J.W. Apple. 1969. Yield reduction of oats caused by the cereal leaf beetle. Journal of

Economic Entomology 62: 298-301.

Rao, S., A. A. Cossé, B. W. Zilkowski and R. J. Bartelt. 2003. Aggregation pheromone of the cereal leaf

beetle: Field evaluation and emission from males in the laboratory. Journal of Chemical Ecology 29:

2165-2175.

SAS 9.1. Copyright (c) 2002-2003 by SAS Institute Inc., Cary, NC, USA.

Southwood, T.R.E. 1978. Ecological Methods with particular reference to the study of insect populations.

Chapman and Hall, London.

Wellso, S.G., R.V. Connin and R.P Hoxie. 1973. Oviposition and orientation of the cereal leaf beetle. An-

nals of the Entomological Society of America 66: 78-83.

Wellso, S.G., W.G. Ruesink and S.H. Gage. 1975. Cereal leaf beetle: relationships between feeding, ovi-

position, mating, and age. Annals of the Entomological Society of America 68: 663-668.

Wilson, M.C., R.E. Treece, R.E. Shade, K.M. Day and R.K. Stivers. 1969. Impact of cereal leaf beetle lar-

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Yoshida, S. 1972. Physiological aspects of grain yield. Annual Review of Plant Physiology 23: 437-464.

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J. ENTOMOL. Soc. BRIT. COLUMBIA 107, DECEMBER 2010

Effect of sex pheromone and Kairomone lures

on catches of codling moth

ALAN L. KNIGHT!

ABSTRACT

Field studies were conducted in sex pheromone-treated apple orchards to evaluate the

performance of a clear 0.11 m/ vertical interception trap coated with oil and baited with

either (E,E)-8-10-dodecadien-l-ol (codlemone), ethyl (£, Z)-2,4-decadienoate (pear

ester), or both attractants (combo) for adult codling moth, Cydia pomonella (L.). Inter-

ception traps baited with codlemone or pear ester caught significantly more males only

or both sexes than unbaited traps, respectively. Interception and delta traps baited with

codlemone caught similar numbers of males. Interception traps baited with pear ester

caught up to 8-fold more males and 30-fold more females than similarly baited delta

traps, respectively. Seasonal catches of females did not differ between light and pear

ester-baited interception traps. Delta traps caught significantly more males, fewer fe-

males, and a similar number of total moths as the interception trap when both were

baited with the combo lure. These data suggest that new clear trap designs can be devel-

oped to increase catches of female codling moth which may enhance seasonal monitor-

67

ing and establish more useful predictive population models.

Key Words: apple, Cydia pomonella, traps, colour, monitoring

INTRODUCTION

Passive interception traps constructed of

clear plastic, coated with an oil film, and

hung vertical in the canopy were developed

to study the behaviors of male and female

codling moths Cydia pomonella (L.), in

orchards treated with sex pheromone

(Weissling and Knight 1994). While, moth

catches on individual interception traps

were not comparable to either sex phero-

mone-baited or light traps, two of the key

attributes of these passive traps were the

capture of nearly equal numbers of each sex

and in providing an unbiased estimate of

the proportion of mated females (Knight

2000). Passive interception traps have been

used to experimentally demonstrate the

occurrence of mating delay (Knight 1997)

and to estimate the level of mating in sex

pheromone-treated orchards (Knight 2006).

These traps have also been used to study

the distribution of moths within an orchard

canopy (Weissling and Knight 1995) and to

examine patterns of adult movement into

sex pheromone-treated orchards (Knight

2007a). In addition to their use as a research

tool, passive interception traps have been

evaluated as monitoring aides to predict the

seasonal phenology of female codling moth

(Knight 2000). Their use demonstrated that

female versus male moth captures can im-

prove the prediction of the start of egg

hatch and were more closely correlated

with levels of fruit injury at both mid-

season and prior to harvest.

Yet, despite these many benefits derived

from using interception traps to monitor

codling moth, a number of drawbacks have

limited their adoption by growers; such as

their relatively low moth capture rate com-

pared with sex pheromone-baited traps,

their non-specificity, the short useful life of

the oil coating, especially during hot or wet

periods, and an overall greater level of dif-

ficulty and higher cost of servicing these

'Yakima Agricultural Research Laboratory, ARS, USDA, 5230 Konnowac Pass Rd., Wapato, WA 98951.

Email: alan.knight@ars.usda.gov

68 J. ENTOMOL. SOC. BRIT. COLUMBIA 107, DECEMBER 2010

traps versus the standard plastic or card-

board traps. Alternative trap and lure de-

signs that could alleviate some of these

issues could enhance the benefits provided

to growers from monitoring female codling

moths within their orchards. One approach

may be to use the bisexual attractant, ethyl

(E, Z)-2, 4-decadienoate (pear ester) to fur-

ther increase the catch of female moths on

interception traps. Capture of female cod-

ling moth in delta traps baited with pear

ester have been reported to improve predic-

tion of first egg hatch and result in more

accurate action thresholds (Knight and

Light 2005a, b). However, the performance

of pear ester relative to codlemone with

standard traps has been inconsistent across

a number of geographical regions with a

broad range in its attractiveness for females

reported (loriatti et al. 2003, Thwaite et al.

2004, IF ichev 2004, Trimble and El-Sayed

2005, Kutinkova et al. 2005, Mitchell et al.

2008).

Growers within the western United

States have widely adopted a lure (combo

lure) loaded with both pear ester and (E£, £)-

8-10-dodecadien-1-ol (codlemone) because

of its higher male and total moth catch than

codlemone lures (Knight et al. 2005). Un-

fortunately, the combo lure catches a low

proportion of female moths and few pest

managers have been willing to identify the

sex of trapped moths (Hawkins 2008).

Thus, the full potential value of utilizing

pear ester to monitor female codling moth

has not been realized. Studies are reported

here that evaluated the effectiveness of bait-

ing interception traps with codlemone, pear

ester, or both attractants. Results suggest

that opportunities exist with codling moth

to develop more efficacious monitoring

systems that include adult female densities.

MATERIALS AND METHODS

General methods. Studies were con-

ducted in 2003 and 2006 in a 20-ha com-

mercial apple orchard, Malus domestica

(Borkhausen) situated near Moxee, WA

(46° 33’ N, 120° 23’ W). This orchard was

a mixed planting of ‘Delicious’ and

‘Golden Delicious’ with a 4.0 — 4.5 m can-

opy height, and a 4.8 x 5.5 m (tree x row)

spacing. The orchard was certified organic

and no supplemental insecticide sprays

were applied during either season, except

for the use of 2 — 6 applications of 1.0%

horticultural oil (Orchex 796, Exxon, Hous-

ton, TX). The orchard was treated with 500

— 1,000 Isomate™ C-Plus dispensers ha 7

loaded as per label with 182 mg of a

53:30:6 blend of codlemone, dodecanol,

and tetradecanol (Pacific Biocontrol, Van-

couver, WA).

Interception traps (0.33 x 0.33 m) were

cut from rolls of 0.25 mm semi-rigid UV-

stabilized film (#10SR36150, W. J. Dennis

Co., Elgin, IL). A 0.5 x 2.0 cm slit was cut

in the top center of each trap 1.5 cm from

the edge. A 17.0 cm piece of 1.4-cm wide

yellow _tie-strapping (Postal Products

Unlimited, Milwaukie, WI) was threaded

through this slit and used to attach each trap

to an orange plastic clip (Suterra LLC,

Bend, OR). Traps were coated with oil

(STP Oil Treatment, STP, Fort Lauderdale,

FL) using a standard paint roller (smooth

texture). Interception traps were replaced

every 3-7 d during studies. Interception

traps were baited with proprietary

codlemone, pear ester, or pear ester and

codlemone lures provided by manufactur-

ers. Septa were attached to interception

traps by piercing lures with a standard pa-

per clip and hooking the clip to the yellow

strapping ca. 1-4 cm above the center top

edge of the trap. The membrane lure was

attached to the strapping with an adhesive

pad provided on the back surface of the

lure. White delta-shaped traps (28.5 x 20.0

cm) with sticky inserts (17.0 x 17.0 cm)

were included in these studies for compari-

son (Trécé Inc., Adair, OR). Sticky liners

were replaced either weekly or up to a 4-wk

interval depending on their condition. Lures

were replaced after 8 wks during the two

seasonal studies in 2003 and 2006. All traps

were placed in the upper third of the can-

opy, ca. 3-m. Interception traps were

J. ENTOMOL. Soc. BRIT. COLUMBIA 107, DECEMBER 2010

clipped with the use of a pole to small

branches, while delta traps were attached to

a 1.3 m schedule 40 pvc pipe (Knight et al.

2006). Traps within each study were evenly

randomized and spaced 15 — 30 m apart in a

grid. Moths were removed from traps in the

field and sexed with the aid of a microscope

in the laboratory.

Baiting interception traps. Two tests

were conducted to evaluate the attractive-

ness of interception traps baited with

codlemone during 2003. The first test was

conducted from 10 — 18 July with a red

rubber septum loaded with 10.0 mg

codlemone (Pherocon® CM 10X, Trécé

Inc.). The baited delta-shaped trap was rep-

licated 6-times and 10 replicates of baited

and unbaited interception traps were in-

cluded. In addition, six unbaited delta traps

were included in the study, but none of

these traps caught any moths and these data

were not included in the analysis. Delta trap

liners and interception traps were replaced

on 15 July. A second study was conducted

from 19 — 29 July using a proprietary plas-

tic membrane lure (Biolure® 10X, Suterra

LLC, Bend, OR). Baited delta traps were

replicated nine times and 15 baited and

unbaited interception traps were included.

Delta trap liners and interception traps were

replaced on 22 July.

Seasonal evaluation, 2003. A portion

of the orchard was subdivided into eight

100 x 100 m replicate blocks. Five baited

(pear ester) and unbaited interception traps,

one baited (pear ester) delta-shaped trap,

and one light trap (6 W blacklight bulb)

baited with Dichlorvos (18.6% active ingre-

dient, No-pest Strip™, United Industries,

St. Louis, MO) were randomly placed in a

grid with a 25 x 25 m spacing within each

block. The study was initiated on 13 June

and all traps were checked 21 times (2-7 d

69

intervals) until 29 August. Data were sum-

marized across dates based on the accumu-

lation of degree days (lower threshold of 10

°C) from first moth flight (5 May) to the

completion of the first (456 degree days)

and second moth flight (1044 degree days)

(Knight 2007b). Moth catch recorded after

10 July was included in the second flight

period.

Seasonal evaluation, 2006. The orchard

was divided into six 100 x 100 m blocks.

Unbaited interception traps and interception

and delta-shaped traps baited with either

pear ester or the combo lure were com-

pared. One delta-shaped trap with each lure

and three interception traps of each type

were placed within each block in a grid

with a 30 x 30 m spacing. Traps were ini-

tially placed in orchards on 13 June and

checked 20 times during the season. Cumu-

lative moth counts for each flight were

based on the accumulation of degree days

from the start of moth flight (4 May). Moth

catch after 6 July was included in the sec-

ond flight.

Statistical analysis. The mean moth

catches from each group of interception

traps placed within each block (5 traps per

block in 2003 and 3 traps per block in

2006) were calculated and used in the sub-

sequent comparison with other trap types.

Count data were transformed with a square

root transformation and proportional data

with the angular transformation to stabilize

variances (Snedecor and Cochran 1967).

Analysis of variance (ANOVA) was used to

compare the main treatment effect for the

various trap and lure combinations

(Analytical Software 2003). Tukey’s

method was used to detect significant (P <

0.05) pair-wise comparisons within signifi-

cant ANOVA’s.

RESULTS

Baiting interception traps. Significant

differences in catches of both sexes and

total numbers of moths occurred among the

three trap-lure combinations in tests with

two different codlemone lures (Table 1).

Codlemone-baited delta and interception

traps caught similar numbers of male and

total moths. Both traps caught significantly

more male and total moths than the un-

baited interception trap. The baited and

70

Comparison of mean (SE) codling moth catches in (2003) unbaited and baited interception and

J. ENTOMOL. Soc. BRIT. COLUMBIA 107, DECEMBER 2010

Table 1.

baited delta traps using high-load codlemone lures.

Mean (SE) moth catch per d'

Lure Trap Male Female Total

Red septa Baited delta 5.8 (0.5)a 0.0 (0.0)b 5.8 (0.5)a

Unbaited interception 1.4 (0.3)b 1.0 (O.1)a 2.4 (0.3)b

Baited interception 5.6 (1. 7a 0.9 (0.1)a 6.5 (1.7)a

ANOVA: F= 8.75, F=111.5 F = 6.62,

df = 2,23 P=001 P<0.0001 P<0.01

Membrane _ Baited delta 4.8 (0.4)a 0.02 (0.01)b 4.8 (0.4)a

Unbaited interception 0.5 (0.1)b L.1(2)a 1.6 (0.3)b

Baited interception 3.4 (0.9)a 0.7 (0.2)a 4.1 (1.l)a

ANOVA: F = 33.7 F=24.2 F=10.5

df = 2,36 P< 0.0001 P<(0.0001 P=0,00n

‘Column means for each lure followed by a different letter were significantly different, P <

0.05, Tukey’s.

unbaited interception traps caught signifi-

cantly more females than the baited delta

trap. Results were similar in tests using

either a rubber septum or membrane lure

(Table 1).

Seasonal evaluation, 2003. Significant

differences in the cumulative male, female,

and total moth catches during each moth

flight occurred among four trap-lure combi-

nations (Table 2). Light traps caught sig-

nificantly more male and total numbers of

codling moth than interception and delta

traps baited with pear ester and unbaited

interception traps. Pear ester-baited inter-

ception and light traps caught similar num-

bers of females. The baited interception

traps caught significantly more female and

total moths than the pear ester-baited delta

traps. The unbaited interception trap caught

significantly more moths than the delta trap

during the first but not the second flight.

The interception traps baited with and with-

out pear ester caught similar numbers of

male moths in the first flight but the baited

trap caught significantly more female and

total moths in the second flight. The pro-

portion of females caught by the different

lure-trap combinations varied significantly,

F 3, 28 = 3.45, P < 0.05. The light trap

caught a significantly lower proportion of

female moths than the baited interception

trap over the entire season. The unbaited

interception and delta traps caught an inter-

mediate proportion of female moths.

Seasonal evaluation, 2006. Significant

differences in the catches of male, female,

and total codling moths occurred between

unbaited interception and interception and

delta traps baited with either pear ester or

the combo lure during both flights in 2006

(Table 3). The combo-baited delta trap

caught significantly more male moths than

all other trap types during both flights. The

other four traps did not differ during the

first moth flight in male moth captures.

However, during the second moth flight the

baited interception traps caught signifi-

cantly more males than the unbaited inter-

ception and the pear ester-baited delta traps.

The unbaited interception trap also caught

significantly more male moths than the pear

ester-baited delta trap.

The baited interception traps caught

significantly more female codling moth

than the baited delta traps during both

flights (Table 3). The unbaited interception

trap caught similar numbers of female

moths as the pear ester-baited interception

J. ENTOMOL. Soc. BRIT. COLUMBIA 107, DECEMBER 2010 71

Table 2.

Comparison of seasonal codling moth catches (2003) in interception and delta traps baited with

pear ester and unbaited interception and light traps, n = 8.

Cumulative mean (SE) moth catch per trap’

Ist moth flight

2nd moth flight

Lure - trap Male Female Total Male Female Total

Unbaited 15.8(1.9)b 10.8(1.5)a 26.5 (2.5)b 74.4 (6.30bce 44.4 (5.9)b 118.8 (7.9)be

interception

Baited 18.1(1.3)b 19.9(2.5)a 38.0(2.7)b 159.1 (5.6)b 128.9(15.3)a_ 288.0 (16.6)b

interception

Baited delta 19(1.0)c 0.8(0.4)b 2.6(0.9)c 21.0(4.4)c = 16.3 (4.5)b 37.3 (8.3)¢

Unbaited 71.6 (13.3)a 27.1 (8.1L)a 98.8 (18.6)a 526.1 (138.9)a 172.1 (43.7)a 698.3 (179.0)a

light

ANOVA: F=33.8 F=169 F = 34.2 F=22.4 F=17.8 F=22.4

df=3,28 P<0.0001 P<0.001 P<0.0001 P<0.0001 P <0.0001 P <0.0001

' Column means followed by a different letter were significantly different, P < 0.05, Tukey’s .

Table 3.

Comparison of seasonal moth catches (2006) in unbaited interception and baited interception

and delta traps with pear ester and pear ester + codlemone (combo) lures, n = 6.

Cumulative mean (SE) moth catch per trap’

lst moth flight

2nd moth flight

Lure - trap Male Female Total Male Female Total

Unbaited 0.6 (0:2)b 0.8(0.2)be 1.3(0.3)be 9.0(1.2)c 6.1(0.8)b = 15.1 (1.9)b

interception

Pear ester-baited 1.0(0.2)b 1.6(0.4)ab 2.7(0.5)ab 17.5(1.7)b 27.4(2.8)a 44.9 (4.2)a

interception

Combo-baited 1.3(0.3)b 1.8(0.4)a 3.2(0.6)ab 17.9(1.7)b 23.5(2.5)a 41.4 (4.0)a

interception

Pear ester-baited 0.0(0.0)b 0.2(0.2)c 0.2 (0.2)c 2.0 (0.7)d 0.8 (0.5)c 2.8 (0.9)¢

delta

Combo-baited PxAS)ar O2(02Qe 2 7.7 Q04)a 32.7(63)a 3.70.1I)e 36.3 (7.5)a

delta

ANOVA: F=9.06 F=5.27 F=9.01 F=23.2 F= 45.0 F=30.2,

df= 4,25. P= 0.0001). P<0001-. P<0,0001 P<0.0001 P<0.0001 P<0.0001

‘Column means followed by a different letter were significantly different, P < 0.05, Tukey’s.

and both delta traps during the first moth

flight. The unbaited interception trap caught

significantly more female moths than either

delta trap during the second flight.

The combo-baited delta trap caught

similar numbers of total moths as the baited

interception traps in both moth flights. The

pear ester-baited delta caught significantly

fewer total moths than these three traps

during both moth flights. The unbaited in-

terception traps caught an _ intermediate

number of total moths: fewer moths than

the combo-baited delta in the first flight and

fewer moths than the baited interception

and the combo-baited delta, but signifi-

cantly more moths than the pear ester-

72 J. ENTOMOL. Soc. BRIT. COLUMBIA 107, DECEMBER 2010

baited delta in the second flight. The pro-

portion of females caught by the different

lure-trap combinations varied significantly,

F454 = 4.56, P < 0.01. The combo-baited

delta trap caught a significantly lower pro-

portion of female moths than either of the

baited interception trap during 2006. The

unbaited interception and pear ester-baited

delta traps caught an intermediate propor-

tion of female moths.

DISCUSSION

The clear, oil-coated unbaited intercep-

tion trap has proved to be an effective tool

to monitor the mating status of female cod-

ling moths and the density, distribution, and

movement of both sexes in experimental

orchards (Weissling and Knight 1994,

1997; Knight 1997, 2000, 2006, 2007a).

Studies reported here demonstrate that bait-

ing the interception trap with codlemone

can increase male catches to levels compa-

rable to standard delta traps and with the

use of pear ester creates a more effective

trap than the delta for monitoring female

codling moth. Further studies with the inter-

ception trap should evaluate the use of the

more potent, acetic acid and pear ester com-

bination lure (Landolt et al. 2007).

Trap effectiveness is strongly influenced

by the anemotactic flight and close-range

behaviors of adult moths to both the lure

and the physical structure of the trap (Foster

and Muggleston 1993). Trap reflectance

and moth vision appear to be critical factors

influencing the capture of male codling

moths in traps of various colours (Knight

and Miliczky 2003, Knight and Fisher

2006). Multiple field observations of adult

codling moth inside screened cages suggest

that both sexes fly accidently into the clear

interception trap while moving within and

through tree canopies (unpubl. data). Sur-

prisingly, an unbaited interception trap

caught significantly more total moths than a

delta trap baited with pear ester.

Flight tunnel studies have revealed that

a significant proportion of male codling

moths orienting to codlemone lures placed

inside of various white sticky traps land on

the outside of the trap first, walk inside, and

then become stuck (Knight et al. 2002).

Switching from white to orange-colour

traps increased male moth catches in the

field, and flight tunnel assays suggested this

was primarily caused by increasing the pro-

portion of males that flew directly inside

the orange versus white trap, especially

under low light conditions (Knight and

Fisher 2006). Interestingly this difference in

moth behavior between trap colours became

greater as the light level was increased.

This may reflect the male’s response period

to both codlemone and pear ester occurring

primarily during scotophase (Knight and

Light 2005c).

Female codling moths respond to trap

colour differently than males. For example,

orange and white delta traps baited with

pear ester had similar catches of females in

both field and flight tunnel experiments

(Knight and Fisher 2006). While, direct

observations of female’s orientation and

contact with pear ester-baited delta traps

have not been reported, the diurnal response

of females to pear ester-baited traps begins

in the late afternoon and occurs on average

earlier than the response of males (Knight

and Light 2005c), and coincides with the

peak timing of oviposition, 1800 — 2200 h

(Ried! and Loher 1980). Thus, trap’s reflec-

tance over the UV or visible spectrum may

be a more critical factor affecting female

than male capture on interception traps.

The relatively high cost of maintaining

interception traps likely will continue to

interfere with grower adoption despite the

enhanced benefits which can be derived

from monitoring female codling moth. Re-

ducing the size of traps and placing them

lower in the canopy would improve their

handling and servicing but would also sig-

nificantly reduce moth catches (loriatti et

al. 2003, Knight and Light 2005c). One

alternative that should be explored is the

use of clear delta traps. The operational

advantages of a clear delta versus intercep-

tion trap are that its profile is smaller so that

J. ENTOMOL. Soc. BRIT. COLUMBIA 107, DECEMBER 2010

traps can more easily be placed in the can-

opy and fewer non-target insects are caught,

and that the sticky liner is placed horizon-

tally inside the trap so the coating does not

run off and is protected from precipitation.

Clear delta traps with clear liners are avail-

able from at least one supplier in Europe

(PRI, Wageningen, The Netherlands), but

its use with pear ester for codling moth has

not been reported. Studies are needed to

assess whether a smaller horizontal sticky

surface placed inside of a clear trap with a

restricted opening would be as effective as

the larger vertical surface of the intercep-

tion trap.

ACKNOWLEDGEMENTS

We would like to thank Brad

Christianson, Chey Temple, and Duane

Larson, USDA, ARS, Wapato, WA for

their help in setting up the field trials, and

Tom Larsen (Suterra LLC, Bend, OR) and

Bill Lingren (Trécé Inc., Adair, OK) for

providing traps and lures. Helpful reviews

were provided by Ally Hari, The Volcani

Center, ARO, Bet Dagan, Israel; Mitch

Trimble, Agriculture and Agric-Food Can-

Contreras, Facultad de Ciencias Agrarias,

Universidad de Talca, Talca, Chile; Greg

Loeb, Cornell University, Geneva, NY;

Maritza Reyes Carreno, Universidad Aus-

tral de Chile, Valdivia, Chile; and Claudio

loriatti, IASMA_ Research Center, San

Michele alto Adige, Italy. This research

was partially funded by the Washington

Tree Fruit Research Commission, We-

natchee, WA.

ada, Vineland, Ontario; Eduardo Fuentes-

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and untreated plots. Entomologia Experimentalis et Applicata 77: 271-275.

J. ENTOMOL. Soc. BRIT. COLUMBIA 107, DECEMBER 2010 2D

Aphids (Hemiptera: Aphididae) associated with rhubarb

(Rheum spp.) in the Matanuska Valley, Alaska: species

composition, seasonal abundance, and potential virus vectors

ALBERTO PANTOJA!”, AARON M. HAGERTY’,

SUSAN Y. EMMERT/', JOSEPH C. KUHL’, KEITH PIKE’,

JUAN M. ALVAREZ and ANDREW JENSEN®

ABSTRACT

Culinary rhubarb, Rheum spp., is one of the priority crop species curated by the United

States Department of Agriculture (USDA) Agricultural Research Service (ARS) in

Palmer, Alaska. Water-pan traps in commercial rhubarb in the Matanuska-Susitna River

Valley near Palmer and in the USDA ARS Rheum germplasm collection caught aphids

belonging to eight species: Aphis helianthi Monell; Chaitophorus neglectus Hottes and

Friso, Euceraphis betulae (Koch); Hayhurstia atriplicis (L.); Macrosiphum euphorbiae

(Thomas); Myzus persicae (Sulzer); Pemphigus spp.; and Rhopalosiphum padi (L.). Only

three of the species (M. euphorbiae, M. persicae, and R. padi) collected in water- pan

traps were also handpicked from rhubarb plants. The bird cherry-oat aphid, R. padi, was

the most abundant species collected in water-pan traps and from rhubarb plants. Based on

their disease transmission capability, 4. helianthi, M. euphorbiae, M. persicae, and R.

padi, can be considered to be of potential economic importance to rhubarb production in

Alaska.

Key Words: aphids, rhubarb, Rheum, Alaska, vectors, germplasm

INTRODUCTION

Culinary rhubarb, Rheum _ spp.

(Polygonaceae), is one of the priority crop

species curated at the Subarctic Agricultural

Research Unit (SARU) of the United States

Department of Agriculture (USDA) Agri-

cultural Research Service (ARS). This site

in Palmer, Alaska, is the primary rhubarb

repository for the USDA ARS National

Plant Germplasm System (NPGS 2010)

which maintains a diverse collection of

plant genetic material. Currently, the SARU

Rheum collection has 41 clonal accessions

(Kuhl and DeBoer 2008), some of which

are infected with Turnip mosaic. virus

(TuMV) (Robertson and Ianson 2005), one

of the most important diseases affecting

rhubarb in Britain (Tomlinson and Walkey

1976) and in Alaska. Turnip mosaic virus

has a large plant host range and a world-

wide distribution (Stobbs and Sterling

1990, Walsh and Jenner 2002, Plant V1-

ruses on Line 2009). The virus spreads me-

' United States Department of Agriculture, Agricultural Research Service, Subarctic Agricultural Research

Unit, P.O. Box 757200, Fairbanks, AK 99775 USA

* Corresponding author. Email: Alberto.Pantoja@ars.usda.gov

>United States Department of Agriculture, Agricultural Research Service, 533 East Fireweed Ave,

Palmer, AK 99645; Current address: Department of Plant, Soil, and Entomological Sciences, University

of Idaho, Moscow, ID 83844-2339 USA

* Washington State University, Irrigated Agriculture Research and Extension Center, 24106 N. Bunn Rd,

Prosser, WA 99350 USA

? University of Idaho, Aberdeen Research and Extension Center, 1693 S 2700 W, Aberdeen, ID 83210

USA

° Washington State University, Pullman, WA 99164 USA

76 J. ENTOMOL. SOC. BRIT. COLUMBIA 107, DECEMBER 2010

chanically and by aphid transmission. The

insect fauna associated with Rheum spp. is

little known and there is no consensus on

the aphid species associated with this plant.

In a bibliography of rhubarb and other

Rheum species, Marshall (1988) listed nine

aphid species from seven genera affecting

rhubarb throughout the world. Other au-

thors reported three (Capinera 2001), or

nineteen species (Blackman and Eastop

2006) of aphids associated with Rheum

species.

There are no known published reports

on aphids associated with rhubarb in

Alaska. The present work was initiated to

identify the aphids associated with rhubarb

in the Matanuska-Susitna River Valley,

Alaska, USA and to identify potential vec-

tors of TuMV.

MATERIALS AND METHODS

Aphids associated with rhubarb were

surveyed on a commercial farm in the Ma-

tanuska-Susitna River Valley near Palmer

(N 61.53°, W 149.08°), Alaska. Samples

were also taken from the SARU Rheum

collection in Palmer (N 61.57°, W 149.25°).

Habitat types surrounding field sites varied.

The commercial farm is located in a devel-

oped rural area adjacent to large-scale vege-

table production. The SARU collection is

located in an isolated area surrounded by

grassland and forest. Rhubarb foliage and

stems were harvested weekly on the farm,

while no harvesting occurred in the SARU

germplasm collection. Rhubarb inflorescen-

ces were removed at both sites.

To construct a voucher collection,

aphids were collected from both sites. Sam-

ples were taken weekly or bi-weekly by

examining rhubarb plants selected at ran-

dom from fields on the commercial farm

(2005 to 2007) and in 2008 by inspecting

all plants in the SARU Rheum collection.

Collected aphids were placed in 95% etha-

nol, and stored for slide mounting and

eventual identification by the authors using

various references (Palmer 1952, Foottit

and Richards 1993, Foottit and Maw 1997,

Pike et al. 2003, Blackman and Eastop

2000, 2006) and museum vouchers. The

abaxial and adaxial sides of the top three

leaves of every plant in the collection were

inspected every seven days during the

months of August and September 2008.

Aphids were also captured in water pan

traps similar to those described by Stoltz ef

al. (1997). Traps were constructed by plac-

ing a 7-mm thick, yellow-green acrylic

square (10 x 10 cm, Yellow 2037, United

States Plastic Corp., Lima, OH, USA) in a

750-ml plastic Rubbermaid® dish (Newell

Rubbermaid Company, Fairlawn, OH,

USA) filled with a 0.05% soap solution

(Ultra Dishwashing Liquid, Planet®, Victo-

ria, BC, Canada). Traps were maintained at

canopy height with the aid of adjustable

stands (Villanueva and Pena 1991; Stoltz et

al. 1997). Traps were placed around field

perimeters just prior to rhubarb emergence

and maintained until all plants were har-

vested (commercial field) or at first frost

(SARU Rheum collection). A total of 33

trap stations were set (six traps/year in the

commercial field and five traps/year in the

germplasm collection) from May to Octo-

ber, 2005-2008. Traps were changed

weekly and brought back to the laboratory

where insects were strained from the soap

solution and preserved in 95% ethanol for

identification. The numbers of aphids per

trap per week were combined to calculate

the total number of aphids per 14-day pe-

riod over the three years.

Additionally, the University of Alaska

Museum of the North (UAM) insect collec-

tion was examined for aphids. The UAM

collection includes the Washburn insect

collection (Washburn 1972; UAM 2009),

which was compiled by USDA entomolo-

gists J.C. Chamberlin, R.H. Washburn, and

others during the 1940’s and 1950’s. This

collection is considered to be the only large

general insect collection maintained in the

state (Pantoja ef al. 2009).

J. ENTOMOL. Soc. BRIT. COLUMBIA 107, DECEMBER 2010

ri

RESULTS AND DISCUSSION

A total of 3,325 specimens representing

eight species and genera were collected

from water-pan traps in commercial rhu-

barb and the SARU Rheum collection

(Table 1). The species include: Aphis heli-

anthi Monell; Chaitophorus neglectus Hot-

tes and Frison; Euceraphis betulae (Koch);

Hayhurstia atriplicis (L.); potato aphid,

Macrosiphum euphorbiae (Thomas); green

peach aphid, Myzus persicae (Sulzer); Pem-

phigus spp.; and bird cherry-oat aphid,

Rhopalosiphum padi (L.). All species were

collected in both locations, but more aphids

(72% of total) were collected from the

SARU Rheum collection than from the

commercial field. Approximately 18% of

the aphids collected could not be identified.

Two species, R. padi (34.1%) and Pemphi-

gus spp. (21.3%) represented 55% of the

overall number of aphids collected. Rhopa-

losiphum padi was the most abundant spe-

cles, representing 26% and 37% of the

aphids collected from the commercial field

and the SARU Rheum collection, respec-

tively. The difference in aphid counts be-

tween sites can be explained: by crop asso-

ciation. The SARU Rheum collection is

located in an isolated site surrounded by

forest and grasses, while the commercial

rhubarb field was surrounded by vegetables

providing additional alternate hosts for the

aphids. There were no aphid colonies on

rhubarb plants, indicating that this crop

does not support development and is not a

preferred host for the aphid species reported

here.

Examination of the UAM insect collec-

tion revealed a total of 38 specimens repre-

senting nine identified species from eight

genera, but none of the specimens were

associated with Rheum spp. (Table 2). To

our knowledge, the present study represents

the first report on aphids from Rheum spp.

in Alaska.

The seasonal abundances of the two

most prevalent species and of three less

numerous species that are potential virus

vectors (discussed below) are shown in

Figures 1 and 2. Both R. padi and Pemphi-

gus spp. were trapped from late June until

mid October, with Pemphigus reaching a

peak in early July, and R. padi peaking in

late August (Fig. 1). Aphis helianthi, M.

euphorbiae and M, persicae were trapped

from early June until mid October, with A.

helianthi peaking in early July, and the

other two species being present at low num-

bers throughout (Fig. 2).

The majority of the species collected in

our study probably represent migratory

aphids moving from other plant species.

The second most abundant genus, Pemphi-

gus, 1S represented by several species not

easily identifiable (Foottit and Maw 1997).

Although the Pemphigus spp. complex is

commonly collected in agricultural fields in

Alaska (Stoltz et al. 1996, 1997), the distri-

bution and biology of the complex is poorly

known and there are no reports on virus

transmission studies with this group (Stoltz

et al. 1997). Pemphigus spp. was the preva-

lent species collected in potato fields in the

Matanuska-Susitna River Valley of Alaska

representing 23% of the water-pan trap

catches (Stoltz et al. 1997). An unidentifi-

able species of the Pemphigus spp. complex

has been reported affecting rhubarb (R.

rhaponticum) roots in New Zealand

(Savage 1982). The agricultural importance

of the Pemphigus species complex needs

attention and revision (Savage 1982, Foottit

and Maw 1997, Stoltz et al. 1997, Black-

man and Eastop 2000).

Although present in low numbers, 4.

helianthi, M. euphorbiae, and M. persicae

are of potential economic importance to

rhubarb production. These three species

along with R. padi are known vectors of

potyviruses (Kortier and Grafius 1994,

Foottit and Maw 1997, Blackman and Eas-

top 2000). Myzus persicae 1s a known vec-

tor of TuMV on Cruciferae (Dombrovsky ef

al. 2005). Aphis helianthi has been associ-

ated with crops of the Compositae and Um-

belliferae families, but its biology and vec-

tor capacity are not well known (Kortier

and Grafius 1994, Blackman and Eastop

2006). Macrosiphum euphorbiae and M.

persicae have been associated with Rheum

spp., suggesting that they might be vectors

78 J. ENTOMOL. SOc. BRIT. COLUMBIA 107, DECEMBER 2010

Table 1.

Sums and percentages of aphids captured in water-pan traps in a commercial rhubarb field and

in the USDA ARS Rheum germplasm collection at the Subarctic Agricultural Research Unit in

Palmer, Alaska, USA, during 2005-2008.

Commercial Collection

Species Sum % Sum %

Aphis helianthi Monell 29 Del 79 3.3

Chaitophorus neglectus Hottes and Frison 18 [2 94 329

Euceraphis betulae (Koch) 167 i 247 10.0

Hayhurstia atriplicis (L.) 33 Se 128 5.4

Macrosiphum euphorbiae (Thomas) a2 3.4 54 2

Myzus persicae (Sulzer) pa Za 24 1.0

Pemphigus spp. 189 20.1 520 21.8

Rhopalosiphum padi (L.) 244 Pia ie 89] 37.4

Unknown 209 222 346 14.5

Total 942 2383

Table 2.

Sums and percentages of aphid species present at the University of Alaska Museum of the

North in Fairbanks, Alaska, USA.

Species

Aphis helianthi Monell

Aphis varians Patch

Bornerina variabilis Richards

Euceraphis sp.

Macrosiphum euphorbiae (Thomas) ]

Nearctaphis bakeri (Cowen)

Nearctaphis yohoensis Bradly

Pterocoma populifoliae (Fitch)

7

l

l

l

7

Nasovonia sp. 2

l

l

4

Rhopalosiphum padi (L.) l

2

Unknown

Total 38

of viruses on rhubarb plants (Marshall

1988, Capinera 2001, Blackman and Eastop

2000, 2006). To our knowledge, our report

represents the first time A. helianthi and R.

padi are linked with rhubarb.

Fifty-nine alate specimens representing

three species, M. euphorbiae (n =18), M.

persicae (n = 6), and R. padi (n =31), were

handpicked from rhubarb plants, suggesting

that these aphid species might serve as virus

Sum

% Host

18.4

2.6 Epilobium angustifolium L.

2.6 Alnus crispa (Aiton) Turrill

Cornus stolonifera Michx.

2.6 Betula resinifera Britton

44.7

5.3. Delphinium sp.

2.6 Malus sp.

2.6 Sorbus sp.

10.5

2.6 Prunus padus L.

Malus sp., Lactuca sativa L.

Populus sp.

5.3. Lonicera tatarica L., Cornus sp.

vectors (Kortier and Grafius 1994, Foottit

and Maw 1997, Blackman and Eastop

2000).

Although not collected in this study, the

melon aphid, Aphis gossypii Glover, and

the turnip aphid, Lipaphis pseudobrassicae

(Kaltenbach), have been previously re-

ported in Alaska (Stoltz et al. 1997). Both

species have been associated with over 50

plant viruses, including TuMV

J. ENTOMOL. SOC. BRIT. COLUMBIA 107, DECEMBER 2010 79

9€R. padi -®Pemphigusspp.

Aphids/14 days

—

wi

So

100

50

0

ey - ¥ - ,

CP vw PNY

ms x

es S a ou oe

a’ ay Yo f v

x

on

>

Figure 1. Sums of aphids per 14 days for the two most abundant species, R. padi and Pemphi-

gus spp. collected with water-pan traps at two sites in proximity to Palmer, Alaska, USA, from

2005 to 2008.

@ <A. helianthi

Aphids/14 days

FM. cuphorbiae

ee Vi. persicae

Figure 2. Sums of aphids per 14 days for the three aphid species that are potential virus vec-

tors, A. helianthi, M. euphorbiae, and M. persicae, collected with water-pan traps at two sites

in proximity to Palmer, Alaska, USA, from 2005 to 2008.

(Dombrovsky ef al. 2005, Blackman and

Eastop 2006). Pantoja (unpublished data)

collected 48 specimens of the cabbage

aphid, Brevicoryne brassicae (L.) in water

pan traps from commercial rhubarb in

Palmer, Alaska, in 2004, after a nearby cab-

bage field was harvested. Brevicoryne bras-

sicae 1s another potential vector of TuMV

to rhubarb (Blackman and Eastop 2000). To

our knowledge, the presence of B. brassi-

cae on rhubarb in the Palmer area repre-

sents a new record for agricultural crops in

the state as this species is not listed by pre-

vious reports (Chamberlin 1949, Washburn

1974, Robinson 1979, Stoltz et al. 1996,

1997, UAM 2009) from agricultural set-

tings in Alaska.

Future research should investigate the

80 J. ENTOMOL. SOC. BRIT. COLUMBIA 107, DECEMBER 2010

correlation between the aphid species pre-

sent in Alaska and their potential associa-

tion with TuMV in rhubarb fields, alternate

hosts of the abundant species, and overwin-

tering habits of the economically important

species. Research is also needed to establish

the potential contribution of aphids to the

spread of TuMV in the SARU germplasm

collection in Palmer.

ACKNOWLEDGEMENTS

We thank Ted Pyrah for access to his

rhubarb farm. Technical assistance in the

field and laboratory was provided by Steve

Lillard, Richard Ranft, Bethany Sweet, Bob

Torgerson, Candice Flint, Cary Curlee, Eric

Brandi Fleshman, Jonathan Nigg, Septem-

ber Martin, and Dan Hall. Critical com-

ments on an earlier draft of this manuscript

were provided by D. Fielding, D. Fleming,

and J. Munyanesa, USDA ARS.

Dotseth, James Malapanis, Nettie Jenkins,

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list and keys. J. Wiley & Sons, Chichester, England.

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pends on suitability of the helper component and the N terminus of the coat protein. Archives of Virology

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and J.A. Downes (eds.), Insects of the Yukon, Biological Survey of Canada Monograph Series No. 2,

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of Canada (Homoptera: Aphidoidea and Phylloxeroidea). Agriculture Canada Research Branch, Ottawa.

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celery. Great Lakes Entomologist 29: 69-77.

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for Horticultural Science 133: 587-592.

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ture, National Agricultural Library, Agricultural Research Service, Bibliographies and Literature of Agri-

culture No. 62. Beltsville, Maryland.

NPGS 2010. http://www.ars-grin.gov/npgs/stats/sitesummary.html (accessed October 2010).

Palmer. M.S. 1952. Aphids of the Rocky Mountain region. Thomas Say Foundation,

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associated with potatoes in Alaska: species composition, seasonal abundance, and potential phytoplasma

vectors. American Journal of Potato Research 86: 68-75.

Pike, K.S., L.L. Boydston and D.W. Allison. 2003. Aphids of western North America north of Mexico.

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Robertson, N. and D. Ianson. 2005. First report of Turnip mosaic virus in rhubarb in Alaska. Disease Notes

89: 430.

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J. ENTOMOL. SOC. BRIT. COLUMBIA 107, DECEMBER 2010

83

SCIENTIFIC NOTE

New distribution records for United States Lygaeoidea

(Hemiptera: Heteroptera)

G.G.E. SCUDDER!

New state records are given for 17 ly-

gaeoid Heteroptera species in the United

States. Henry and Froeschner (1988) sum-

marized the known state distribution for the

species of Heteroptera in Canada and the

continental United States. A number of new

state records have been noted during recent

identification of specimens from various

sources. These are detailed below. I am

indebted to the Museum curators listed for

loan of specimens

Data cited are those on specimen labels.

Museum abbreviations used in the text are

as follows:

CNC: Canadian National Collection of

Insects, Agriculture and Agri-Food Canada,

Ottawa, ON (R.G. Foottit).

ROM: Royal Ontario Museum, Toronto,

ON (D. Currie).

UCB: Essig Entomological Collection,

University of California, Berkeley, CA (C.

Barr).

UBC: Spencer Entomological Collec-

tion, Beaty Biodiversity Museum, Univer-

sity of British Columbia, Vancouver, BC

(K.M. Needham).

UIM: W.F. Barr Entomological Collec-

tion, University of Idaho, Moscow, ID

(W.F. Barr).

USNM: National Museum of Natural

History, Smithsonian Institution, Washing-

ton, DC (T.J. Henry).

UNITED STATES NEW RECORDS.

Family GEOCORIDAE

Geocoris atricolor Montandon

MT: 19, Missoula, 9.vi.1985 (G.G.E.

Scudder) [USNM].

Family HETEROGASTRIDAE

Heterogaster flavicosta Barber

AR: Yell Co:, Mt..Nebo St. Pk., ca 10

mi SW Russellville on Ark. Hwy. 155,

along trail, 1800', 9.v.1984 (R. Jaagumagi,

R. Vineyard) [ROM #84001 5c].

Family LYGAEIDAE

Nysius angustatus Uhler

MT: 19, Apgar, Glacier, 9.1x.1966

(G.G.E. Scudder) [USNM]; 14, Bozeman,

19.vii.1929 (W. Downes) [UBC]; 34 39,

id., 17.vii.1936 (W. Downes) [UBC]; 39,

Logan Pass, 6664', 9.1x.1966 (G.G.E. Scud-

der) [CNC].

Nysius raphanus Howard

GA: 14, Millen, 25.viii.1957 (J.G.

Chillcott) [CNC]. NH: 19, Lakes of the

Clouds, Mt. Washington, 5000", 4.viii.1954

(Becker, Munroe & Mason) [CNC]; 10, id.,

9.vii.1954 (Becker, Munroe & Mason)

[CNC]. SC: 19, Aiken, at light, 11.vi.1957

(J.R. Vockeroth) [CNC]; 14, id.,

12.vi.1957 (J.R. Vockeroth) [CNC]; 19,

id., 12.vi.1957 (W.R.M. Mason) [CNC]; 1d

12, id., 23.vi.1957 (W.R.M. Mason)

[CNC]; 19, Kirksey, 24.vi.1957 (W.R.M.

Mason) [CNC]; 9d 69, Seneca,

20.viii.1957 (L.A. Kelton) [CNC]; 2¢ 29,

id., (W.R. Richards) [CNC].

Xyonysius californicus (Stal)

TN: 19, Indian Gap, 5200', Great

Smoky Mt. N.P., 8.vii.1957 (W.R.M. Ma-

son) [CNC].

Family OXYCARENIDAE

Crophius scabrosus Uhler

NV: 43 49, Carson City, 25.vi.1929

(R.L. Usinger) [UCB]; 64 8°, Washoe

County, Pyramid, Juniperus sp., 4-

5.vil.1947 (R.L. Usinger) [UCB].

Family RHYPAROCHROMIDAE

Subfamily PLINTHISINAE

Plinthisus americanus Van Duzee

ID: 19, Lemhi Co., 8 mi S. Tendoy,

' Beaty Biodiversity Centre and Department of Zoology, University of British Columbia, 6270 University

Boulevard, Vancouver, BC V6T 1Z4

84 J. ENTOMOL. Soc. BRIT. COLUMBIA 107, DECEMBER 2010

10.1x.1969 (W.F. Barr) [UIM].

Subfamily RHYPAROCHROMINAE

Tribe DRYMINI

Scoloposthethus diffidens Horvath

NV: 1¢ 192, Lake Tahoe, Zephyr Cove,

1900 m., 9.xi1.1986 (A. Smetana) [CNC].

Scoloposthethus thomsoni Reuter

NV: 1¢ 12, Lake Tahoe, Zephyr Cove,

1900 m., 9.x11.1986 (A. Smetana) [CNC].

Tribe GONIANOTINI

Trapezonotus arenarius Linnaeus

ID: 12, Kootenai Co., Worley, sweep-

ing alfalfa, 30.1v.1953 (W.F. Barr) [UIM].

Tribe LETHAEINI

Xestocoris nitens Van Duzee

AR: 14, Logan Co., Cove Lk., 9 mi SE

Paris, ex debris at edge of lake, 25.v.1986

(J.M. Campbell) [CNC].

Tribe MEGALANOTINI

Megalonotus sabulicola (Thomas)

MT: 1°, Glacier Nat. Park., L. McDon-

ald, 13.vi.1985 (G.G.E. Scudder) [CNC];

1¢ 12, Missoula, 9.vi.1985 (G.G.E. Scud-

der [CNC].

Spragisticus nebulosus (Fallén)

VES 16.9 Essex Jee

Brimley) [CNC].

Tribe MYODOCHINI

Ligyrocoris sylvestris (Linnaeus)

NM: 192, Jemez Sprs., 23.viii.1972

(L.A. Kelton) [CNC]. WA: 12, Newport,

Pioneer Park, 8.1x.1966 (G.G.E. Scudder)

[CNC].

Perigenes constrictus (Say)

MN: 2¢ 12, Minneapolis, 28.viii.1969

(A.B. Acton) [USNM].

Tribe RHYPAROCHROMINI

Cordillonotus stellatus Scudder

ID: 14, Ada Co., Black’s Cr. Res.,

30.v1.1963 (A.R. Gittins) [UIM].

Tribe UDEOCORINI

Neosuris castanea (Barber)

WA: 16, Oroville, E. Osoyoos L., 48°

S8'N 119°25'W, Purshia assoc., AN

BGxhl, Pitfall trap 04-4, 5.v-30.v.1994

(G.G.E. Scudder) [CNC]; 19, id., trap 04-5

(G.G.E. Scudder) [CNC]; 1, id., trap 05-1

(G.G.E. Scudder) [CNC]; 19, id., trap 04-4,

2.vill-6.1x.1994 (G.G.E. Scudder) [CNC].

IS.x.1952

REFERENCES

Henry, T.J. and R.C. Froeschner (eds.). 1988. Catalog of the Heteroptera, or True Bugs, of Canada and the

Continental United States. E.J. Brill, Leiden. 958 pp.

J. ENTOMOL. Soc. BRIT. COLUMBIA 107, DECEMBER 2010

85

SCIENTIFIC NOTE

The Schizopteridae (Hemiptera), a family new to Canada

G.G.E. SCUDDER!

The Schizopteridae is a very small fam-

ily of minute bugs belonging to the Infraor-

der Dipsocoromorpha (Stys 1995). These

insects are typically extremely small, and

have antennae with the first two segments

very short, while the third and fourth anten-

nal segments are longer and thinner, with

many long, thin, erect or semi-erect setae.

Keyed by both Slater and Baranowski

(1978) and Stys (1995), the Schizopteridae

are 0.8 to 2.00 mm long. The forewings are

convex, strongly sclerotized and beetle-like,

but they overlap slightly along the midline.

Characteristically, these bugs have the pros-

ternum inflated and produced ventrally so

as to enclose the fore coxae and the ventral

surface of the head. The hind coxae are also

peculiar in having the inner surface pro-

vided with a pair of roughened pads which

are used in conjunction with a metasternal

spine as a Jumping organ.

The family has a worldwide distribution

and is primarily tropical or subtropical, with

at least 35 genera and about 120 described

species (Stys 1995). Only four genera and

four species are reported to occur in North

America (Henry 1988; Henry et al. 2010).

Males have three tarsal segments on each

leg, while females have two segments on

the fore and middle legs, and three seg-

ments on the hind legs. The four known

North American taxa have been keyed by

Baranowski and Slater (1978).

While Glyptocombus saltator Heide-

mann is reported from several of the United

States (AR, DC, GA, MD, MI, TN, VA,

WA), Corixidea major McAtee & Malloch

is known from Arkansas, Florida, Okla-

homa, Tennessee and Virginia, while Nan-

nocoris arenarius Blatchley is recorded

from Florida, Georgia, North Carolina and

Virginia (Henry et al. 2010). Schizoptera

bispina McAtee & Malloch is restricted to

Florida in North America (Henry 1988).

This note records the first occurrence of the

family in Canada. The record is based on

14 with the data: “CAN: BC: Vancouver,

Pacitic Spirit’ Pr,.Pk.;, 26.41.1997, Colls.. J.

Lea, A. Klimaszewski, ex forest edge”. The

specimen (Fig. 1) 1s 1.33 mm long and is

deposited in the Canadian National Collec-

tion of Insects, Agriculture and Agri-Food

Canada, Ottawa, ON.

This British Columbia specimen is not

any of the genera or species keyed in Slater

and Baranowski (1978). Dr. T.J. Henry has

examined the specimen and informs me that

it appears to be new to science. However,

not being an expert on the family involved,

I am inclined not to describe it at the pre-

sent time. The elucidation of the correct

identity must remain a future task.

Elsewhere, the Schizopteridae most fre-

quently occur in damp soil and in forest

litter. Little is known of their biology and

feeding habits, but they are thought to be

predators (Slater and Baranowski 1978).

I am indebted to Dr. T.J. Henry for his

advice. Don Griffiths kindly took the photo-

graph presented as Figure |. Launi Lucas

processed this note.

REFERENCES

Henry, T.J. 1988. Family Schizopteridae Reuter, 1891. The Schizopterids. Pp. 682-683. /n 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.

' Beaty Biodiversity Centre and Department of Zoology, University of British Columbia, 6270 University

Boulevard, Vancouver, BC V6T 124

86 J. ENTOMOL. Soc. BRIT. COLUMBIA 107, DECEMBER 2010

Henry, T.J., G.F. Hevel and S.W. Chordas III. 2010. Additional records of the little-known Corixidea

major (Heteroptera: Schizopteridae) from Arkansas and Oklahoma. Proc. Entomol. Soc. Wash. 112:

475-477.

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.

Stys, P. 1995. Schizopteridae. Pp. 80-83. Jn R.T. Schuh and J.A. Slater. True Bugs of the World

(Hemiptera: Heteroptera). Classification and Natural History. Cornell University Press, Ithaca and Lon-

don.

Figure 1. Family Schizopteridae, 1¢, CAN: BC: Vancouver, Pacific Spirit Pr. Pk., 26.vi.1997,

Colls. J. Lea, A. Klimaszewski, ex forest edge. Photograph by D. Griffiths.

J. ENTOMOL. Soc. BRIT. COLUMBIA 107, DECEMBER 2010

87

SCIENTIFIC NOTE

Range expansion and hosts of Crenophthalmus pseudagyrtes

Baker (Siphonaptera: Ctenophthalmidae) in central Alaska

G.E. HAAS!, N. WILSON’, J.R. KUCERA®, T.O. OSBORNE’,

J.S. WHITMAN? and W.N. JOHNSON?

Traub (1980, 1985) noted that Eocene mam-

mal data account for Ctenophthalmus in the

western hemisphere. The Nearctic flea Cr. pseu-

dagyrtes originated from an African ancestor

that accompanied its host rafting in the south

Atlantic from Africa to South or Middle Amer-

ica while continents were much closer together

in the early Eocene epoch (Traub 1980: pp. 144-

145, 161; 1985: pp. 368-406). Traub (1980: p.

119) commented “The host versatility exhibited

by Ctenophthalmus is exceptional... ”. He gave

examples of “Extra Ordinal fleas within Two or

More Orders. of Host” - including

”9

“Ctenophthalmus (C.) agyrtes, etc.” on

“microtines, murids, soricids, etc.” (Traub 1985:

Table 8.15). Traub (1980: pp. 142-143) had

observed “The range of its main host, Microtus,

greatly exceeds that of C. (N.) pseudagyrtes.”

Traub (1980: pp. 142-145) was especially puz-

zled by the anomalous range of the only

Ctenophthalmus species in North America being

limited in the West by the Rocky Mountains and

the North by Southern Canada. He and Holland

overlooked collections in central Alaska, Ari-

zona, and New Mexico. Only preliminary re-

cords for Alaska were published (Haas ef al.

1989). Five new Alaska Ct pseudagyrtes and

two new host Myodes rutilis (Pallas) records are

given in New Material Examined and Fig. | of

the present report.

New material examined: USA, Alaska,

Galena, 15 km WNW, N bank Yukon River: 1°

ex Sorex cinereus, 21-vii-1988, T. O. Osborne

4520. McGrath vicinity, Kuskokwim River wa-

tershed, 14 ex My. rutilis, 29-vi-1989, J. S.

Whitman; 1d same data but pool of 8 My.

rutilis, 7-21-vii-1989. Nowitna NWR, NW bank

Little Muddy River, Hades Lake vicinity,

Nowitna River watershed, 64°38’N, 154°00’W,

13, 12 ex Microtus xanthognathus (Leach), 9-

1x-1995, W. N. Johnson 421; same data but 19,

9-ix-1996, WNJ 383.

"PO Box 60985, Boulder City, NV, USA 89006

One interior Alaska watershed with Cr. pseu-

dagyrtes reported by Haas et al. (1989, Fig. 1)

increased to three, i.e. from north bank Yukon

River west of Galena to Nowitna River and Kus-

kokwim River above McGrath. Tributaries of

the latter two rivers interdigitate at elevations

below 305 m to provide potential for small

mammal populations to move between these

watersheds.

This topography can account for the range of

Mi. xanthognathus having a southwest marginal

record near the mouth of the Takotna River (Hall

1981, Map 460). That location is directly across

from the Kuskokwim River oxbow in which

McGrath is located. The ectoparasite study area

was within 1.6 km of the city.

The habitat was second-growth deciduous

forest interspersed with upland grassy areas.

Understory was a heavy grass/forb type. At the

Nowitna NWR 1985 burn study areas, Mi. xan-

thognathus habitat was “tall shrub-sapling stage

of early successional forest,” with some patches

of mature conifers, moss and herbs (Paragi ef al.

1996). Similarities between the two study areas

suggest that both provided habitats favored by

taiga voles for their post-fire vegetation

(Conway and Cook 1999).

Myodes rutilis stands apart from the other

hosts by being amphiBeringean and without

previous Ct. pseudagyrtes records. However, the

southern-ranging Nearctic congener southern

red-backed vole, My. gapperi (Vigors) is listed

as a major host of Ct. pseudagyrtes in Canada by

Holland (1949, 1985) with 43 collections. In

contrast, the northern red-backed vole ranges

from Alaska across Canada to Hudson Bay north

of the southern red-backed vole range (Holland

1985, Map 86), without any record of being a

host in Canada (Holland 1985, p. 489, Map 34).

Updated distribution maps of the two red-backed

voles are in MacDonald and Cook (2009: Maps

ZO 21):

> Department of Biology, University of Northern Iowa, Cedar Falls, IA, USA

* Associated Regional and University Pathologists, Inc., Salt Lake C ity, UT, USA

* PO Box 22, Okaukuejo, Via Outjo 9000, Namibia

> Alaska Department of Fish and Game, Anchorage, AK, USA

88 J. ENTOMOL. SOC. BRIT. COLUMBIA 107, DECEMBER 2010

%K cr. PSEVDAGYATES

Figure 1. Locations of three study areas in Alaska where Ctenophthalmus pseudagyrtes specimens were

collected. Upper left symbol marks north bank Yukon River, 15 km WNW of Galena, lower symbol marks

McGrath, Kuskokwim River and upper right symbol marks Hades Lakes, Nowitna NWR.

Ctenophthalmus pseudagyrtes had a large

literature with many genera and species of small

mammal hosts, mainly Nearctic shrews, mice

and voles in North America north of Mexico

(Benton 1980, Holland 1985, Haas et al. 1989,

Fagerlund et a/. 2001, etc.) and in certain bio-

geographic provinces of Mexico (Morrone ef al.

2000). Sorex cinereus and especially Microtus

pennsylvanicus are hosts that range widely

across the continent. The taiga vole, however,

despite ranging from Hudson Bay across much

of Alaska (Hall 1981, Map 460; MacDonald and

Cook 2009: Map 19) only has Cr. pseudagyrtes

records in Alaska. Adult specimens of Ct. pseu-

dagyrtes were scarce and no larvae were found.

Yet 160 of 291 vole nests were positive for 2420

adult fleas (Haas 1982). These were vole fleas of

14 taxa with inclusion of 618 adults of 10 taxa

reared from 29 of the nests. Specimens of Ct.

pseudagyrtes were not found.

New records of three more Cr. pseudagyrtes

ex two taiga voles in Nowitna NWR remind us

of the hypothesis (Haas ef a/. 1989) that fossil

records of Zakrzewski (1985) show where taiga

voles shifted from northeast and Midwest US

states northwest to Alaska and could have car-

ried Cr. pseudagyrtes with them. A recent re-

view of Mi. xanthognathus fossils in Alaska and

a new taiga vole distribution map are presented

by MacDonald and Cook (2009: pp. 97-98, Map

19).

REFERENCES

Benton, A.H. 1980. An atlas of the fleas of the eastern United States. Marginal Media, Fredonia, NY, 177 pp.

Conroy, C.J. and J.A. Cook. 1999. Microtus xanthognathus. Mammalian Species 627: 1-5.

Fagerlund, R.A., P.L. Ford and P.J. Polechla, Jr. 2001. New records for fleas (Siphonaptera) from New Mexico with notes

on plague-carrying species. Southwestern Naturalist 46: 94-96.

Haas, G.E. 1982. Fleas (Siphonaptera) from vole nests in subarctic Alaska. Canadian Journal of Zoology 60: 2157-2161.

Haas, G.E., N. Wilson, T.O. Osborne, R.L. Zarnke, L. Johnson and J.O. Wolff. 1989. Mammal fleas (Siphonaptera) of

Alaska and Yukon territory. Canadian Journal of Zoology 67: 394-405.

Hall, E.R. 1981. The mammals of North America. Vol. 2, 2™ edition, John Wiley & Sons, Inc., NY.

Holland, G.P. 1949. The Siphonaptera of Canada. Canada Department of Agriculture Technical Bulletin 70.

Holland, G.P. 1985. The fleas of Canada, Alaska and Greenland (Siphonaptera). Memoirs of the Entomological Society of

Canada 130.

MacDonald, S.O. and J.A. Cook. 2009. Recent Mammals of Alaska. University of Alaska Press, Fairbanks. 387 pp.

Morrone, J.J., R. Acosta and A.L. Gutierrez. 2000. Cladistics, biogeography, and host relationships of the flea subgenus

Ctenophthalmus (Alloctenus), with the description of a new Mexican species (Siphonaptera: Ctenophthalmidae). Jour-

nal of the New York Entomological Society 108: 1-12.

Paragi, T.F., W.N. Johnson, D.D. Katnik and A.J. Magoun. 1996. Marten selection of postfire seres in the Alaskan taiga.

Canadian Journal of Zoology 74: 2226-2237.

Traub, R. 1980. The zoogeography and evolution of some fleas, lice and mammals. Pp. 93-172. /n Fleas. Proceedings of

the International Conference on Fleas, Ashton Wold, Peterborough, U.K., 21-25 June 1977. R. Traub & H. Starcke

(eds.). A.A. Balkema, Rotterdam. 420 pp.

Traub, R. 1985. Coevolution of fleas and mammals. Pp. 295-437. /n K.C. Kim (ed.). Coevolution of Parasitic Arthropods

and Mammals. John Wiley & Sons, NY. 800 pp.

Zakrzewski, R.J. 1985. The fossil record. Jn Biology of New World Microtus. R.H. Tamarin (ed.). Special Publication

American Society of Mammalogists No. 8. Pp. 1-51.

NOTICE TO CONTRIBUTORS

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Submission deadline for Vol. 108 is September 1, 2011. Submit contributions to:

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Journal of the

Entomological Society of British Columbia

olume 107 Issued December 2010 ISSN #0071-0733

Directors of the Entomological Society of British Columbia, 2010-2011 ......:.....2

G.G.E. Scudder. Melacoryphus admirabilis (Uhler) (Hemiptera: Lygaeidae) new

to Canada, with additional Canadian provincial records for other Heteroptera .

COOH EHEHHEEHEEE HEHEHE HHH EHEEHHEEH EEE HEHEHE EESHEHEEEHEEEEEEHEHEEEHHEHEEHEEHE HEHEHE HEHEHE EHEEHEEEEEEHEEHEESEHEHEEEHEEEEEHES

K.G.A. Hamilton and Yong Jung Kwon. Taxonomic changes in Dicraneura

Hardy, Colladonus’ Ball and Macrosteles Fieber (Homoptera-

Auchenorrhyncha) in the Montane Cordilleran Ecozone ..................ss0eeeeeeeees 11

L.M. Humble, E. John, J. Smith, G.M.G. Zilahi-Balogh, T. Kimoto and M.K.

Noseworthy. First records of the banded elm bark beetle, Scolytus schevyrewi

Semenov (Coleoptera: Curculionidae: Scolytinae), in British Columbia ....... 24

Jeremy R. deWaard, Leland M. Humble and B. Christian Schmidt. DNA barcod-

ing identifies the first North American records of the Eurasian moth, Eupithe-

cia pusillata (Lepidoptera: Geometridae). .......:..5.2.-22-¢.cancuceoeene ee a

Robert A. Cannings, Mare A. Branham and Robert H. McVickar. The fireflies

(Coleoptera: Lampyridae) of British Columbia, with special emphasis on the

light-flashing species and their distribution, status and biology .................... a5

Richard S. Winder, Donna E. Macey and Joe Cortese. Dominant bacteria associ-

ated with broods of mountain pine beetle, Dendroctonus ponderosae

(Coleoptera: Curculionidae, Scolytinae): .:..:.u..208,.0ses se 43

George D. Hoffman and Sujaya Rao. Cereal leaf beetle, Oulema melanopus (L.)

(Coleoptera: Chrysomelidae), attraction to oat plantings of different ages ....57

Alan L. Knight. Effect of sex pheromone and kairomone lures on catches of cod-

ling moth (Lepidoptera: Tortricidae) on clear interception traps .................. 67

Alberto Pantoja, Aaron M. Hagerty, Susan Y. Emmert, Joseph C. Kuhl, Keith

Pike, Juan M. Alvarezand Andrew Jensen. Aphids (Homoptera: Aphididae)

associated with rhubarb in the Matanuska Valley, Alaska: species composition,

seasonal abundance, and potential virus VeCtors .:.2.........c.sssedereneeae eee eee

NOTES

G.G.E. Scudder. New distribution records for United States Lygaeoidea

(Hemiptera: Heteroptera) s..2......cccéccccckk.ctavuouscneadensesesaee ee Sane eae 83

G.G.E. Scudder. The Schizopteridae (Hemiptera), a family new to Canada ...... 85

G.E. Haas, N. Wilson, J.R. Kucera, T. O. Osborne, J.S. Whitman and W.N. John-

son. Range expansion and hosts of Ctenophthalmus pseudagyrtes Baker

(Siphonaptera: Ctenophthalmidae) in central Alaska ...............::ccccssseeceeeeseees

NOTICE TO CONTRIBUTORS .).:.cc450.3 cee